transcriber's note: this book is heavily illustrated; references to the illustrations have been removed from this text version. please look for the fully illustrated html version at http://www.gutenberg.net.] the tale of mr. jeremy fisher by beatrix potter _author of_ _"the tale of peter rabbit," &c._ frederick warne & co., inc. new york copyright, 1906 by frederick warne & co for stephanie from cousin b. once upon a time there was a frog called mr. jeremy fisher; he lived in a little damp house amongst the buttercups at the edge of a pond. the water was all slippy-sloppy in the larder and in the back passage. but mr. jeremy liked getting his feet wet; nobody ever scolded him, and he never caught a cold! he was quite pleased when he looked out and saw large drops of rain, splashing in the pond-"i will get some worms and go fishing and catch a dish of minnows for my dinner," said mr. jeremy fisher. "if i catch more than five fish, i will invite my friends mr. alderman ptolemy tortoise and sir isaac newton. the alderman, however, eats salad." mr. jeremy put on a macintosh, and a pair of shiny goloshes; he took his rod and basket, and set off with enormous hops to the place where he kept his boat. the boat was round and green, and very like the other lily-leaves. it was tied to a water-plant in the middle of the pond. mr. jeremy took a reed pole, and pushed the boat out into open water. "i know a good place for minnows," said mr. jeremy fisher. mr. jeremy stuck his pole into the mud and fastened the boat to it. then he settled himself cross-legged and arranged his fishing tackle. he had the dearest little red float. his rod was a tough stalk of grass, his line was a fine long white horse-hair, and he tied a little wriggling worm at the end. the rain trickled down his back, and for nearly an hour he stared at the float. "this is getting tiresome, i think i should like some lunch," said mr. jeremy fisher. he punted back again amongst the water-plants, and took some lunch out of his basket. "i will eat a butterfly sandwich, and wait till the shower is over," said mr. jeremy fisher. a great big water-beetle came up underneath the lily leaf and tweaked the toe of one of his goloshes. mr. jeremy crossed his legs up shorter, out of reach, and went on eating his sandwich. once or twice something moved about with a rustle and a splash amongst the rushes at the side of the pond. "i trust that is not a rat," said mr. jeremy fisher; "i think i had better get away from here." mr. jeremy shoved the boat out again a little way, and dropped in the bait. there was a bite almost directly; the float gave a tremendous bobbit! "a minnow! a minnow! i have him by the nose!" cried mr. jeremy fisher, jerking up his rod. but what a horrible surprise! instead of a smooth fat minnow, mr. jeremy landed little jack sharp the stickleback, covered with spines! the stickleback floundered about the boat, pricking and snapping until he was quite out of breath. then he jumped back into the water. and a shoal of other little fishes put their heads out, and laughed at mr. jeremy fisher. and while mr. jeremy sat disconsolately on the edge of his boat--sucking his sore fingers and peering down into the water--a _much_ worse thing happened; a really _frightful_ thing it would have been, if mr. jeremy had not been wearing a macintosh! a great big enormous trout came up--ker-pflop-p-p-p! with a splash--and it seized mr. jeremy with a snap, "ow! ow! ow!"--and then it turned and dived down to the bottom of the pond! but the trout was so displeased with the taste of the macintosh, that in less than half a minute it spat him out again; and the only thing it swallowed was mr. jeremy's goloshes. mr. jeremy bounced up to the surface of the water, like a cork and the bubbles out of a soda water bottle; and he swam with all his might to the edge of the pond. he scrambled out on the first bank he came to, and he hopped home across the meadow with his macintosh all in tatters. "what a mercy that was not a pike!" said mr. jeremy fisher. "i have lost my rod and basket; but it does not much matter, for i am sure i should never have dared to go fishing again!" he put some sticking plaster on his fingers, and his friends both came to dinner. he could not offer them fish, but he had something else in his larder. sir isaac newton wore his black and gold waistcoat, and mr. alderman ptolemy tortoise brought a salad with him in a string bag. and instead of a nice dish of minnows--they had a roasted grasshopper with lady-bird sauce; which frogs consider a beautiful treat; but _i_ think it must have been nasty! the end proofreading team. [illustration: a frog he would a-wooing go. r. caldecott picture books] a frog he would a-wooing go [illustration] [illustration] a frog he would a-wooing go, _heigho, says_ rowley! whether his mother would let him or no. _with a rowley-powley, gammon and spinach_, _heigho, says_ anthony rowley! [illustration] so off he set with his opera-hat, _heigho, says_ rowley! and on his way he met with a rat. _with a rowley-powley, gammon and spinach_, _heigho, says_ anthony rowley! [illustration] "pray, mr. rat, will you go with me," _heigho, says_ rowley! "pretty miss mousey for to see?" _with a rowley-powley, gammon and spinach_, _heigho, says_ anthony rowley! [illustration] [illustration] now they soon arrived at mousey's hall, _heigho, says_ rowley! and gave a loud knock, and gave a loud call. _with a rowley-powley, gammon and spinach_, _heigho, says_ anthony rowley! [illustration] "pray, miss mousey, are you within?" _heigho, says_ rowley! "oh, yes, kind sirs, i'm sitting to spin." _with a rowley-powley, gammon and spinach_, _heigho, says_ anthony rowley! [illustration] [illustration] [illustration] "pray, miss mouse, will you give us some beer?" _heigho, says_ rowley! "for froggy and i are fond of good cheer." _with a rowley-powley, gammon and spinach_, _heigho says_, anthony rowley! [illustration] [illustration] [illustration] "pray, mr. frog, will you give us a song? _heigho, says_ rowley! "but let it be something that's not very long." _with a rowley-powley, gammon and spinach,_ _heigho, says_ anthony rowley! [illustration] [illustration] "indeed, miss mouse," replied mr frog, _heigho, says_ rowley! "a cold has made me as hoarse as a hog." _with a rowley-powley, gammon and spinach_, _heigho, says_ anthony rowley! [illustration] "since you have caught cold," miss mousey said. _heigho, says_ rowley! "i'll sing you a song that i have just made." _with a rowley-powley, gammon and spinach_, _heigho, says_ anthony rowley! [illustration] but while they were all thus a merry-making, _heigho, says_ rowley! a cat and her kittens came tumbling in. _with a rowley-powley, gammon and spinach_, _heigho, says_ anthony rowley! [illustration] [illustration] the cat she seized the rat by the crown; _heigho, says_ rowley! the kittens they pulled the little mouse down. _with a rowley-powley, gammon and spinach_, _heigho, says_ anthony rowley! [illustration] this put mr. frog in a terrible fright; _heigho, says_ rowley! he took up his hat, and he wished them good night. _with a rowley-powley, gammon and spinach_, _heigho, says_ anthony rowley! [illustration] but as froggy was crossing a silvery brook, _heigho, says_ rowley! a lily-white duck came and gobbled him up. _with a rowley-powley, gammon and spinach_, _heigho, says_ anthony rowley! [illustration] [illustration] so there was an end of one, two, and three, _heigho, says_ rowley! the rat, the mouse, and the little frog-gee! _with a rowley-powley, gammon and spinach_, _heigho, says_ anthony rowley! [illustration: randolph caldecott's picture books "the humour of randolph caldecott's drawings is simply irresistible, no healthy-minded man, woman, or child could look at them without laughing." * * * * * _in square crown 4to, picture covers, with numerous coloured plates._ 1 john gilpin 2 the house that jack built 3 the babes in the wood 4 the mad dog 5 three jovial huntsmen 6 sing a song for sixpence 7 the queen of hearts 8 the farmer's boy 9 the milkmaid 10 hey-diddle-diddle and baby bunting 11 a frog he would a-wooing go 12 the fox jumps over the parson's gate 13 come lasses and lads 14 ride a cock horse to banbury cross, &c. 15 mrs. mary blaize 16 the great panjandrum himself _the above selections are also issued in four volumes, square crown 4to, attractive bindings red edges. each containing four different books, with their coloured pictures and numerous outline sketches._ 1 r. caldecott's picture book no. 1 2 r. caldecott's picture book no. 2 3 hey-diddle-diddle-picture book 4 the panjandrum picture book _and also_ _in two volumes, handsomely bound in cloth gilt, each containing eight different books, with their coloured pictures and numerous outline sketches._ r. caldecott's collection of pictures and songs no. 1 r. caldecott's collection of pictures and songs no. 2 * * * * * miniature editions. _size 5-1/2 by 4-1/2 art boards, flat back._ -----four volumes entitled r. caldecott's picture books nos. 1, 2, 3, and 4. _each containing coloured plates and numerous outline sketches in the text._ * * * * * _crown 4to, picture covers._ randolph caldecott's painting books. three volumes _each with outline pictures to paint, and coloured examples_. ----_oblong 4to. cloth_. a sketch book of r. caldecott's. _containing numerous sketches in colour and black and white_. * * * * * frederick warne & co--ltd london & new york _the published prices of the above picture books can be obtained of all booksellers or from the illustrated catalogue of the publishers._ printed and copyrighted by edmund evans, ltd., rose place, globe road, london, e.i.] [illustration] [illustration] how freckle frog made herself pretty _by_ charlotte b. herr _designs_ frances beem published in the shop of p. f. volland & co. chicago u.s.a. * * * * * this little story is told and the little pictures were drawn for a good little child named --------- copyright 1913 p f. volland & co chicago, u. s. a. * * * * * [illustration] how freckle frog made herself pretty once upon a time there was a little girl named marian, and she had a doll called big mary. marian loved big mary, and meant to be very good to her. but sometimes she was not. santa claus had brought big mary one snowy christmas night, and he had brought also a great many pretty clothes for her to wear. there were three dresses, a warm red one for winter, and a white one, very thin, for summer, and still another, of beautiful blue silk with lace on it, for best. then, also, there were little skirts, and tiny stockings, and pretty little shoes with shiny buckles and real heels, and there was a pink parasol, and, best of all, a dear little muff, made of soft white fur, to keep big mary's hands warm in cold weather. at first little marian loved to dress big mary in all these pretty things, and she would put on first the warm red dress, and then the thin white one, and then the one of blue silk with the beautiful lace. and she would raise the big parasol and put it over big mary's head. but she hardly ever gave big mary the little white muff to hold, because that was for very, _very_ best. little marian's own mamma had said so. but when marian's birthday came, grandma gave her a doll's trunk, and after that the days were not so pleasant for big mary. it was so much fun to pack the trunk that little marian often took off all the clothes big mary had on to put them away in the trunk. many a time poor big mary had to sit for hours all undressed, and she would shiver and shake, until at last one time when little marian had left her lying all night on the floor without any clothes on, she took a dreadful cold and became very ill. then little marian was very sorry for what she had done, and she put big mary to bed and sent for dr. prince. when the doctor came he looked at big mary's tongue, and felt her pulse. and then he shook his head and looked very grave. he said that big mary must take some medicine every day, and must sit out in the fresh air, and always wear her best clothes all the time; for she was a very sick doll indeed. so little marian dressed big mary in the blue silk trimmed with lace, because that was her very best dress, and she raised the pink parasol and put it over her head and she gave big mary the white muff to hold, because that was for very, _very_ best. then she carried big mary out to the gray rock in the back yard where the nasturtiums grow, to sit in the fresh air all day long. now little miss freckle frog lived under the big rock. she was ugly, as all frogs are, but she loved pretty things, perhaps because she was not pretty herself. but although she was not pretty, she was a kind-hearted little body, and all her friends liked her. [illustration] every day when big mary sat in the sunshine, freckle frog crept out from under the rock, and hid in the grass, and watched her. she thought big mary was wonderful, but she thought that the blue silk dress and the pink parasol were more wonderful still, and the little soft muff,--that was the most wonderful of all! and poor little freckle frog wished that she had a blue silk dress with lace, and a pink parasol like big mary. but most of all she wished that she had a little soft muff. now it happened, too, that it was just about the time for mr. robin redbreast to give his big party in the orchard, and little freckle frog had been invited, and more than that, her own cousin, billy bullfrog, had promised to sing, and of course she wished to look just as nice as she could. so early one fine day, she went to see the morning glory ladies who live near the back porch and always wear such beautiful dresses, and she said to them: "oh, dear morning glory ladies, your dresses are always so beautiful! but have you seen big mary's blue silk trimmed with lace? it is more beautiful still, the loveliest dress in the whole world! would you mind making me one like that to wear to robin redbreast's party? my cousin, billy bullfrog, is to sing, and i wish so very much to look just as nice as i can. i am not one bit pretty like big mary, but clothes always help a great deal, you know. would you mind lending me one for the party?" [illustration] but the morning glory ladies were angry because they had not been invited, and they would not help her. poor little freckle frog felt very badly to think she had hurt their feelings. she almost cried about it. but just then little black spider, who was a good friend of hers, peeped out from under a leaf and said: "i wouldn't mind them. they are a conceited lot anyway. it is a hot day, too, and they are apt to be cross on hot days. i will spin you all the lace you want." and so he did. he wove it all that day in his web, and the next morning he brought her a long piece of the loveliest spider-lace as fine as a cobweb. little freckle frog was very grateful to him. [illustration] "but what shall i do for a parasol?" she asked. "oh, i'll tell you!" called a soft little voice, and when she looked up she saw a tiny white butterfly resting on a flower. "i know where there is the dearest little mushroom. it kept the rain off of me the other day, and it is just as soft and pink as big mary's parasol." so she showed freckle frog where to find the mushroom, and it was very soft and pink, just as she had said, and freckle frog was very happy about it. "now if i only had a muff," she sighed, "i could look just as beautiful as big mary at the party!" just then there was a great noise in the tree near the rock and robin redbreast himself flew out from among the leaves, but right at her feet dropped a little white caterpillar. he was so frightened that he curled himself up into a ball and lay very still. he made big mary laugh, but freckle frog had a bright idea. [illustration] "oh what a splendid muff you would make!" she cried. "would you mind if i wear you to the party just this once?" the poor little caterpillar uncurled himself. "if you will promise to take care of me and not let robin redbreast eat me," he answered, "i shall be only too glad to be your muff." so freckle frog went to the party and wore the cobweb lace, and carried the mushroom parasol, and held the soft little white caterpillar for a muff. she even bought a sweet-pea bonnet to please the morning glory ladies. then robin redbreast said she looked better than anybody else at his party, and big mary, who was well enough by that time to go also, said so, too. [illustration] now robin redbreast, as you must know, always had his parties just at twilight. he himself was always in better voice then, he said, and so he felt sure that billy bullfrog and all the other singers must be, too. then the world was lovelier at that time than it was through the long, hot day, when sensible people like birds and frogs, and sometimes even babies and dolls, took naps and did not stir out at all. at twilight one could always depend upon the sky to grow very soft and pink, and the fairies never failed to hang the leaves with dewdrops, all to make his parties beautiful! the cherries tasted better then, too, and later still, when it began to grow dark, the katy-dids would play if any one cared to dance. so robin redbreast always gave beautiful parties, but even he had never given so beautiful a one before. little freckle frog was very happy. every one admired her beautiful lace, and she told them all how kind little black spider had been. and by and by, when it came to be time for refreshments, she ate a whole cherry. she never had tasted one before, but as she told mr. sparrow, who had brought it to her, she really never had dreamed how delicious a big red cherry could be. then, when the katy-dids began to play, she danced with her cousin, billy bullfrog, until it was time to go home. [illustration] there was only one thing that troubled her, and that was that the morning glory ladies were still angry with her. for little freckle frog wanted to be friendly with everybody. but at last another idea came into her head. she would give a party herself, just as beautiful a one as robin redbreast's, and have it early in the morning so that the morning glory ladies could come. [illustration] so that very evening, before she went home, she told big mary all about it, and big mary promised to help all she could. robin redbreast said that he would surely come, and so did billy bullfrog and all the rest. freckle frog invited little black spider, too, and even the little white caterpillar. "and you needn't be a muff this time," she said, "but just eat cherries, and have a good time." then, early the next morning, before any one else was up, she went to invite the morning glory ladies, for they are always good-natured then, and never frown and scowl at people until the sun is hot. "please, dear morning glory ladies," said freckle frog, "will you come to my party? i want you more than any one else." then the morning glory ladies fluttered with joy, for they loved parties, and they smiled and answered her: "yes, indeed, we shall come, little freckle frog, and wear our best dresses, too." then at last freckle frog was perfectly happy, and she laughed to herself and said: "it really doesn't matter about my being pretty any more, for every one likes me now!" [illustration] * * * * * produced from images generously made available by the university of florida, the internet archive/children's library) [illustration: cover] one shilling or on cloth eighteen pence the frog who would a wooing go [illustration] by charles bennett london: routledge, warne, and routledge. [illustration: the frog who would a wooing go charles bennett] a frog he would a-wooing go, whether his mother would let him or no. off he set with his opera-hat. on the road he met with a rat. [illustration] [illustration] "pray, mr. rat, will you go with me, kind mrs. mousey for to see?" they soon arrived at mousey's hall. they gave a loud tap, and they gave a loud call. [illustration] [illustration] "pray, mrs. mouse, are you within?" "yes, kind sirs, and sitting to spin." "pray, mrs. mouse, now give us some beer, that froggy and i may have good cheer." [illustration] [illustration] "pray, mr. frog, will you give us a song? let the subject be something that's not very long." "indeed, mrs. mouse," replied the frog, "a cold has made me as hoarse as a hog." [illustration] [illustration] "since you have caught cold, mr. frog," mousey said, "i'll sing you a song that i have just made." as they were in glee and merrymaking, a cat and her kittens came tumbling in. [illustration] [illustration] the cat she seized the rat by the crown, the kittens they pulled the little mouse down. this put mr. frog in a terrible fright, he took up his hat, and he wished them good night. [illustration] [illustration] as froggy was crossing it over a brook, a lilywhite duck came and gobbled him up. so here is an end of one, two, three- the rat, the mouse, and little froggy. [illustration: the end] [illustration: song: the frog who would a wooing go] the frog who would a-wooing go. by charles bennett. on the quiet sedgy bank of a stagnant pool, and under the shadow of rank reeds and bulrushes, sat two frogs. they had retired from the shoal, who were disporting themselves in the water, and were earnestly talking. the elder of the two, an old matron, addressing the younger, who, by-the-by, was her son, said,-"my dear froggy, you had better stop quietly with me; you do not know what dangers you may encounter, if you leave your secluded home." "croak, croak!" said froggy. "ah, my son!" continued the old lady, "i see that, like most young frogs, you are very obstinate, and will not listen to reason. but why on earth you should wish to go gadding after a poor, hungry little mouse, is more than i can tell--you with your beautiful legs and speckled coat, born to a splendid estate of reeds and water, the heir of nine bulrushes and a water-lily. i thought you were more of a frog." "croak, croak!" said froggy again. "have you thought of the boys who throw stones?" "croak!" "or the birds with long beaks?" "croak!" "or the ducks?" "croak!" "if you want to go a-wooing, there are frogs in your own station in life; indeed, with your personal appearance, you might even aspire to an eft or a lizard." "croak!" persisted the sulky little frog. "you are no better than a tadpole!" said his mother, getting very angry at last; and no sooner were the words out of her lips, than up jumped master froggy in a passion, and taking his opera-hat under his arm, off he went at a rapid pace, singing at the top of his voice, so as to hide his rage,- "rowley, powley, gammon and spinach. 'heigh o!' says anthony rowley." froggy had not walked very far before he saw, jogging on before him, a brown little fellow in a long-tail coat and blucher boots, who carried an old cotton umbrella. "dear me," thought the frog, "that looks like my friend mr. rat;" and sure enough so it was. "how do you do?" asked master froggy, when he had overtaken him. "pretty well!--how's your self?--where are you going?--fine day!--squeak!" replied mr. rat, in a succession of short, shrill sentences. "i'm going," said froggy, "to see the pretty little widow mousey, who lives in that snug cottage yonder. pray come with me, for i feel rather bashful at going by myself." "with all my heart," replied mr. rat; and off they went together. they soon arrived at the cottage; and the rat having given a loud knock, while the frog gave a loud "croak," mrs. mousey put away her spinning-wheel in a great hurry, and admitted her guests. "good morning, mrs. mouse," said the frog; "we were out walking, and thought we would give you a call." "you are very kind, i am sure," replied mrs. mousey. "pray sit down and rest yourselves; i dare say you are tired." "and here--i say--squeak!--mrs. mousey--some beer!--we're thirsty," said mr. rat. "croak--let's enjoy ourselves while we can," observed the frog. "certainly," said mousey. "i'll draw you some of the last brew." so mousey drew some beer, and they sat down very cosily; and soon were chatting so comfortably, that master froggy thought he should soon get rid of his bashfulness, and then should be able to ask pretty mrs. mousey to marry him. presently their little hostess proposed a song, and called upon froggy to oblige; but, "really," he replied, "i must be excused, for the fog last night gave me such a cold that i'm as hoarse as a hog." he didn't forget that he had been singing "rowley, powley," as he came along, but he was afraid that his voice was not good enough for his company. "well," said mousey, laughing, "i am sorry for that; but if you won't sing, i will." so she sang a pretty little song she had just composed, and a very charming ditty it was--rather shrill, perhaps, but very well sung indeed. after this, i need hardly tell you that they enjoyed themselves amazingly. perhaps mr. rat drank rather too much beer; but altogether it was a very pleasant little party, and froggy had so far got over his bashfulness as to squeeze mrs. mousey's paw once or twice rather tenderly. but while they were thus happily employed, a terrible old cat who lived in the neighbourhood, and went by the name of "browzer," was tying on her shawl--calling to her kittens, and saying, "come, my children, it is a fine day--let us go for a walk. make haste, for something tells me we shall find some dinner on our way." and sure enough they did; for after looking after little birds, and trying the windows of all the pantries they knew,-"b-row!" said the cat, snuffing the air; "do you know--i'm not quite certain--but--yes, really--i smell mouse." "mew!" said the kittens; "we're so glad." "and i think, also, rat." "oh, come along!" said the kittens. "don't make a noise," whispered the cat. slowly and cautiously they crept on towards mrs. mousey's cottage, till at an unexpected moment, and just as mrs. mouse was going to get a fresh mug of beer, in tumbled the cat and her kittens. down went the rat under the cat's paw--up in a corner the two kittens got the mouse. "m-row-ow, fit-z-z!" and rat and mouse were killed. "holla!" says the frog; "this won't do! perhaps they'll be after me in a minute. i must be off home to my mother." and sure enough off he went (trembling like a leaf), but as rapidly as he could. "oh, why did i ever leave home?" said this foolish frog: "i should have been safe enough with my mother. i'll never leave home again. never! never! never!" "quack, quack!" observed a duck who had been watching him. "oh, my goodness gracious!" said the frog; "what shall i do now? there's the very duck that ate up my uncle who went abroad! now, if i can't cross over this brook in a single jump, i shall never get home alive. here goes!" but, alas! since it must be told, he could not cross the brook in one jump. in he fell--splash! up came the duck. "quack, quack! gobble, gobble, gobble!" and the poor frog never got home at all. we are all sorry for his untimely end, and wish that the duck had not gobbled him up: but we must not forget that if he had been less self-willed and obstinate, if he had only paid attention to what his mother told him, he might have been safe at home--perhaps, in due course, married to an amiable frog, and the father of a large family of innocent little tadpoles. [illustration: edm evans. engraver & printer. raquet court. fleet st.] * * * * * transcriber's notes: obvious punctuation errors repaired. there was some variation between the hyphenation of the text itself and of the text of the illustrations. these variations were retained. [illustration] the ducks and the frogs by ff boston joseph.h.francis mdcccxlix. [illustration] the ducks & the frogs, a tale of the bogs. by fanny fire-fly the ducks and the frogs, a tale of the bogs. by fanny fire-fly. with engravings by hartwell, from designs by billings. boston: joseph h. francis. m dccc xlix. entered according to act of congress, in the year 1848, by alonzo hartwell, in the clerk's office of the district court of the district of massachusetts. white & potter, printers, j. w. wilcox, electrotyper a. hartwell, wood engraver. littleton, mass. [illustration] the ducks and the frogs it chanced upon a certain day, when cheerful summer, bright and gay, had brought once more her gift of flowers, to dress anew her pleasant bowers; when birds and insects on the wing made all the air with music ring; when sunshine smiled on dell and knoll, two ducks set forth to take a stroll. 'twas morning; and each grassy bank of cooling dew had deeply drank- each fair young flower was holding up its sweet and freshly painted cup, filled with bright dew drops, every one; gay, sparkling treasures for the sun, who bears them lightly to the sky, holds them as vapor far on high, till with his rays in dazzling tints, the rainbow on the cloud he paints. but our two ducks we'll not forget, they were not troubled by the wet; they rambled on, and soon they took the path that led them to a brook, [illustration] whose sparkling waters danced along, with a gushing, rushing, rippling song. the ramblers, when they reached the brink, stepped down to bathe, and take a drink. they loved to frolic, dive and dash beneath the water with a splash. they washed and smoothed each glossy feather, then said, "let's have a swim together!" as moving gracefully, they went, they heard loud tones of sad lament. they listened, and did sharply look for cause of woe in that sweet brook; and soon espied beneath some bushes, among the reeds and tall, green rushes, a company of long-faced frogs, a delegation from the bogs; sitting with their up-turned faces, in attitudes to please the graces, around a stone, on which was speaking a member of this grave marsh meeting. the ducks were pleased; they knew them all, for very often they did call at that sweet brook, to hear them sing; they thought their music quite the thing. "and now," said they, "we will draw near," for much they wished to see and hear what was this fuss and noise about, so joined the party to find out. the frogs received them with a smirk, and gave their hands with nervous jerk. bowing and smiling in return, the ducks prepared themselves to learn [illustration] from what the orator might say, the cause of all their friends' dismay. now the chief speaker in this scene, dressed in a suit of bottle green, folding his arms across his breast, again the meeting thus addressed: "my friends," said he, "i'm rather hoarse, and must be brief in my discourse; but as these ducks have joined our band, i wish to have them understand we have not come to this fair spot, to break the peace or hatch a plot; but we have met to form a plan to waken in the heart of man, pity for our sad condition. we would present a grave petition, beseeching of the men who rule, that we, lone dwellers of the pool, may be permitted to reside in safety, with our scanty tribe. we humbly say there's no occasion, to send an army of invasion into our loved and quiet bogs, to murder happy, harmless frogs. take our own dear sons and daughters, drag them from their winter quarters, then, when no heart with pity melts, to cut them up as food for smelts! think what a very shocking fate, caught and killed, and used as bait, to take those harmless little fishes to multiply man's dainty dishes." now, as the frog this sentence spoke, _each brother gave a solemn croak._ the gentleman in bottle-green was quite exhausted by his theme; he paused a moment, wiped his brow; then said, "i think you will allow we've been a persecuted race, since first on earth we had a place. there is, i'm told, a land called france, where all the people sing and dance- and they acquire their easy grace by living on our helpless race; and though i say it with a sigh, 'tis this that makes them all so spry." puffing for breath, the speaker stopped and quickly from the stone he hopped. the ducks, while listening to this tale, had felt their very hearts turn pale. at length, the largest of the two, a handsome drake, in green and blue, arose, and opening wide his beak, _bowed, coughed_, and then began to speak. "neighbors, i'm not a coward bird- but the sad story i have heard, would cause the boldest one to quake, and makes my every feather shake. i like the plan that you propose, to write a list of these your woes, and ask for mercy from these men; but have it done by some smart pen; if stated by some able writer, i think your fortunes may be brighter." [illustration] just at this moment, up there sprung a frog quite pert, for one so young; said he, "i vote for emigration, 'twill save us all this botheration!" our proud drake turned, in great surprise, while grave rebuke flashed from his eyes. said he, "it makes my blood run cold, to see young folks so smart and bold. there's not a duckling of my brood, that would presume to be thus rude; young sir, i will a lesson give, that may be useful while you live: wait till your counsel others seek, and then think twice before you speak! for you, the elders of this tribe, i hope you here will still reside. in every pleasant brook and marsh, you'll meet with cares and trials harsh; if you'll but try to be contented, much that's wrong will be prevented. my lady duck and i 'tis plain, are wiser than when here we came. we thought our lot was very hard, when shut within the poultry yard; although 'tis large, and well supplied with water, and all else beside for happiness and comfort too, yet much we wished for something new. our wings are clipped, we cannot fly, and this too costs us many a sigh. we seldom pass our owner's gate, he keeps his poultry rather straight. we should not have been out to-day, but duck and i just ran away; and as we came to bathe this morn, fretful we felt, and quite forlorn; we thought our lot in life so sad, and all our troubles quite too bad. could we have got our brood away, we had quit town this very day. as gloomily we stepped along, the air was filled with many a song from happy creatures, gay and bright, rejoicing in the morning light. the dew, o'er flowers and trees was flung, like diamonds pure, in drops it hung; all nature seemed reproaching us, for making all this dismal fuss. but we grew calmer as we walked, of all these cheering things we talked. and hearing all your griefs and sighs, much better feelings did arise. for let me tell you, friends and brothers, listening to the woes of others, and pitying their deep distress, will ever make our own seem less. then patience whispers, (pray regard her,) your lot though hard, might still be harder. now, gossips, i am tired of speaking, our ducklings too we must be seeking; although it makes our heart-strings quiver, to see yon bright and pleasant river; and hearing its cool waters splashing, we long beneath them to be dashing. yet we must close this visitation, and without farther hesitation, resist our very strong desire, and cheerful to our homes retire. our kindest wishes rest with you, so, now good friends, we'll bid adieu." the ducks then smoothed each ruffled feather, and gracefully walked off together. the frogs with courtesy arose, and stretched themselves high on their toes; and so far conquered all their fears, they gave their friends three parting cheers! then as they sank upon the grass, this resolution they did pass: "here, now, before we separate, we pledge ourselves, to bear our fate with patience; and if ill betide, we'll try to find some brighter side. our homes with cheerful tones shall ring, and over every care _we'll spring_." they stopped; each folded his green dress about him with much cheerfulness; shook hands all round, and said "good day," then merrily they _hopped away_. [illustration] when these bright people all were gone, and i sat musing quite alone, out of this their simple preaching, came the lesson they'd been teaching. each little reader too can see what seems so very clear to me. * * * * * 'tis this: that dark-browed discontent must from our hearts be quickly sent; whate'er may be our daily lot, think all is well, and grumble not; a generous pity feel for all, and charity for great and small. one other hint we also find, that children all should bear in mind, treat aged people--strangers too, with reverence; it is their due. take warning from that frog so young, and keep a bridle on the tongue! these teachings seem so very plain, we hope they are not given in vain. [illustration: the end.] [illustration: boston joseph. h. francis mdcccxlix.] university of kansas publications museum of natural history volume 15, no. 3, pp. 175-181, 1 fig. march 7, 1962 a new species of frog (genus tomodactylus) from western méxico by robert g. webb university of kansas lawrence 1962 university of kansas publications, museum of natural history editors: e. raymond hall, chairman, henry s. fitch, theodore h. eaton, jr. volume 15, no. 3, pp. 175-181, 1 fig. published march 7, 1962 university of kansas lawrence, kansas printed by jean m. neibarger, state printer topeka, kansas 1962 29-1923 a new species of frog (genus tomodactylus) from western méxico. by robert g. webb thirteen specimens of frogs collected in the summers of 1960 and 1961 in the mexican states of durango and sinaloa represent a heretofore unnamed species. the specimens have been deposited in the museum of natural history of the university of kansas (ku) and in the museum of michigan state university (msu). the species may be named and described as follows: tomodactylus saxatilis new species _holotype._--ku 63326 (fig. 1); obtained eight miles west of el palmito, sinaloa, approximately 6100 feet, on 23 june 1961; original field number, 2354 of robert g. webb. _paratypes._--a total of 12 specimens: ku 63327-33, same data as holotype, 23-25 june 1961; msu 4085-88, two miles north of pueblo nuevo, durango, approximately 6000 feet elevation, 24 july 1960; msu 4089, one half mile west of revolcaderos, durango, approximately 6600 feet, 29 july 1961. _diagnosis._--a species of _tomodactylus_ possessing the following combination of characters: (1) tips of two outer fingers truncate, about twice width of narrowest part of digit; (2) tympanum small, less than one half diameter of eye; (3) ventral surfaces smooth; (4) contrasting marbled pattern on back and top of head, and (5) venter whitish, lacking dark marks. _description of holotype._--adult male; snout-vent length, 31.5 (measurements are in millimeters and were taken by means of dial calipers reading to one tenth of a millimeter); width of head, 11.2; length of head, 10.3; horizontal diameter of eye, 3.1, and of tympanum, 1.2; distance from eye to nostril, 3.8; internarial width, 2.9; interorbital width, 4.1; width of eyelid, 2.5; lumbar gland (left side), 7.0 x 2.6; distance from axilla to groin, 15.2; tibial length, 12.7; length of foot, 13.1. head slightly wider than body; tip of snout rounded, slightly truncate; canthus rounded; tympanum small, less than one half diameter of eye; tympanum having posterior margin ill-defined, separated from eye by distance about equal to diameter of tympanum; diameter of eye slightly less than distance from eye to nostril; width of eyelid about two thirds interorbital width; paratoid gland indistinct; lumbar glands high, separated from insertion of leg by about one millimeter; back and sides of body having low, scarcely elevated pustules; top of head, limbs and venter smooth; few low, whitish pustules below and behind tympanum, and low on sides of body; posterior surface of thighs and anal region pustulate; one pair of whitish postanal spots; ventral disc attached near insertion of legs, lacking conspicuous transverse fold; skin loose on throat, chest and abdomen. digits not webbed; tips of two outer fingers truncate, having terminal transverse grooves, about twice width of narrowest part of digit; digits of first and second fingers slightly expanded; fingers from shortest to longest, 1-2-4-3, first only slightly shorter than second; three palmar tubercles; inner palmar tubercle about one third size of large median tubercle; outer tubercle about one tenth size of large median tubercle; four supernumerary palmar tubercles; tips of toes slightly wider than narrowest part of digits; toes from shortest to longest, 1-2-5-3-4, second only slightly shorter than fifth; inner metatarsal tubercle about four times size of small outer metatarsal tubercle; supernumerary tubercles on foot small; no tarsal fold; heels touching when tibiae adpressed to thighs; tibiotarsal articulation reaching eye when leg adpressed to side of body. contrasting marbled pattern on back and top of head; contrasting, mostly barred, pattern on limbs; ventral surfaces whitish, lacking dark marks, but having minute dark peppering; marbling of dorsal surfaces blackish and whitish in preservative. vomerine teeth lacking; internal choanae lateral, partly concealed by maxillaries; tongue smooth, elongate, shallowly notched distally, free for about half its length; vocal sac median; internal vocal slits large and near angle of jaw. _variation._--twelve males closely resemble the holotype. two specimens from pueblo nuevo are soft and not well preserved. the ranges of variation (means in parentheses) for the 13 males comprising the type series are: snout-vent length, 25.5-31.5 (27.9); width of head, 9.7-11.2 (10.5); diameter of eye, 2.9-3.6 (3.2); horizontal diameter of tympanum, 1.1-1.8 (1.4); length of tibia, 11.1-13.1 (12.1); length of foot, 11.0-13.1 (12.1). the pale ground color of the marbled pattern in most specimens is least extensive on the back and arms, but most extensive on the legs. the lumbar glands are slightly elevated and conspicuous, and in ku 63328 are extremely protuberant, or (ku 63330) evident on left side but flattened and indistinct on right side. the back is rough having low, scarcely elevated pustules, but becomes less rough anteriorly and most of the top of head is smooth. the three specimens from pueblo nuevo, durango, differ slightly from the other specimens examined in lacking pairs of postanal white spots, and in having smooth backs (slightly pustulate in msu 4088). the tibiotarsal articulation fails to reach the eye in ku 63330. the small inner palmar tubercle is continuous with the large median tubercle on the right hand of ku 63330, and lacking on both hands of ku 63329 and on the left hand of ku 63328. the tip of the tongue is entire in some specimens and in others has an irregular margin. _coloration of living specimens._--marbled pattern on back and top of head of dark brown to blackish on yellowish-gold; pattern slightly less contrasting on limbs than on back, consisting of brown to grayish on pale yellow; side of head and body grayish sometimes having pale yellow to whitish spots; iris blackish having fine reticulation of yellowish to greenish-gold; venter dirty white. _habitat._--the three records of occurrence for _tomodactylus saxatilis_ are in a mixed boreal-tropical habitat, which is transitional between a pine-oak forest at higher elevations and a tropical deciduous forest at lower elevations. the mixed boreal-tropical habitat is most conspicuous at elevations between approximately 7800 and 5500 feet on southerly exposed slopes of barrancas and arroyos of the dissected plateau of the sierra madre occidental. the mixed boreal-tropical habitat occurs for approximately 30 miles along the paved highway (mexican highway 40) between cd. durango, durango, and mazatlán, sinaloa. the records of occurrence in those states that are along this highway are separated by 14.5 miles (_via_ road). [illustration: fig. 1. _tomodactylus saxatilis_ new species, adult male, ku 63326, holotype (× 2), dorsal view.] the terrain consists of occasional level areas, but is mostly of steep hillsides. dominant trees are large oaks and pines; a characteristic pine is the sad or drooping-needle pine, locally called "pino triste." the vegetational cover is usually open, including grasses, small oaks and pines, broad-leaved shrubs and herbs, prickly pears, magueys, thorny acacias, bracken fern, and epiphytes in trees. ferns occur in moist protected places, and orchids are occasional, sometimes in trees. outcrops of rock, boulder-strewn areas, and occasional rock slides (talus) also characterize the terrain. _tomodactylus saxatilis _seems to be restricted to rocky habitats. the individuals collected were detected when they called at night from within crevices of rocks or from exposed perches on rocks and boulders; some calling frogs were out of reach on steep rock walls. the call is a single, loud, high peep. _comparisons._--dixon (1957) recognized six species of _tomodactylus_ (_nitidus_, _dilatus_, _albolabris_, _angustidigitorum_, _fuscus_ and _grandis_) in his revision of the genus. another species (_rufescens_) was subsequently described by duellman and dixon (1959). _tomodactylus saxatilis_ differs from all the species named immediately above by the combination of characters given in the diagnosis. _tomodactylus saxatilis_ differs from _nitidus_, _angustidigitorum_ and _grandis_ in having the tips of the two outer fingers widened and truncate; _saxatilis_ differs from _dilatus_, _albolabris_, _fuscus_ and _rufescens_ in having a smooth venter (not pustulate), a contrasting pale and dark marbled pattern on the back, and a lack of "flash" colors on the femora. _tomodactylus saxatilis_, having lumbar glands, also resembles three species referred to the genus _syrrhophus_. _tomodactylus macrotympanum_ was described by taylor (1940:496, 497) as having a large, moderately distinct lumbar gland; the species was referred to the genus _syrrhophus_ by dixon (_op. cit._:384). according to firschein (1954:55), _syrrhophus smithi_ and _s. petrophilus_ have elongate lumbar glands shaped like those in _tomodactylus_. _tomodactylus saxatilis_ resembles _macrotympanum_, _smithi_ and _petrophilus_ more than it does other species; all four attain large maximal sizes, and have lumbar glands, mostly smooth ventral surfaces, three palmar tubercles (sometimes absent in _saxatilis_), and usually contrasting dorsal patterns (reduced to flecks and spots in all species except _saxatilis_). _tomodactylus saxatilis_ differs from _macrotympanum_ in having an extensive marbled dorsal pattern and a small tympanum, and differs from _smithi_ and _petrophilus_ in having a marbled dorsal pattern and the tips of the outer two fingers widened and truncate. _tomodactylus saxatilis_ differs from all other named species of _syrrhophus_ in having conspicuous lumbar glands and in lacking inguinal glands. _remarks._--the characteristics delimiting the genera _tomodactylus_ and _syrrhophus_ are not agreed upon by all workers (see discussions by firschein, 1954:50; langebartel and shannon, 1956:164; and dixon, 1957:383). i have referred _saxatilis_ to the genus _tomodactylus_ on the basis of a lumbar gland, which was considered a distinguishing character for the genus by smith and taylor (1948:46) and langebartel and shannon (1956:165). lumbar glands are longer than broad, at least one third the distance from axilla to groin, lateral and usually high, and often conspicuous and protuberant. the elevation or flatness of the lumbar glands seems to be due to individual variation; living specimens in the field had conspicuous and protuberant, or non-elevated, indistinct lumbar glands. lumbar glands are not to be confused with inguinal glands, which are roundish, often yellowish, sometimes diffuse, lateral but low, often inconspicuous, and usually not protuberant. inguinal glands occur in the genus _microbatrachylus_ and in some species of _eleutherodactylus_, and have been described as flat, or low, or small, or indistinct for most species of _syrrhophus_. for financial assistance with field work i am grateful to rollin h. baker, and those individuals who administer the michigan state university development fund and the bache fund of the national academy of sciences (grant no. 463). i am grateful also to j. keever greer, donald f. switzenberg, and rudolph a. scheibner for aid in the field, to edward h. taylor, james r. dixon, and william e. duellman for profitable discussions, and to thomas sweringen for figure 1. the specific name alludes to the habitat (latin, _saxatilis_ = found among rocks). literature cited dixon, j. r. 1957. geographic variation and distribution of the genus tomodactylus in mexico. texas journ. sci., 9(4):379-409, 5 figs., 1 map, december. duellman, w. e., and dixon, j. r. 1959. a new frog of the genus _tomodactylus_ from michoacan, mexico. texas journ. sci., 11(1):78-82, 1 fig., 1 table, march. firschein, i. l. 1954. definition of some little-understood members of the leptodactylid genus _syrrhophus_, with a description of a new species. copeia, 1954, 1:48-58, february 19. langebartel, d. a., and shannon, f. a. 1956. a new frog (_syrrhophus_) from the sinaloan lowlands of mexico. herpetologica, 12(3):163-165, 2 figs., september 1. smith, h. m., and taylor, e. h. 1948. an annotated checklist and key to the amphibia of mexico. bull. u. s. nat. mus., no. 194, pp. iv + 118. taylor, e. h. 1940. herpetological miscellany. univ. kansas sci. bull., 26(15):489-571, 10 pls., 7 figs., november 15. 29-1923 university of kansas publications museum of natural history volume 18, no. 1, pp. 1-10 september 24, 1968 the genera of phyllomedusine frogs (anura: hylidae) by william e. duellman university of kansas lawrence 1968 university of kansas publications, museum of natural history editors of this number: frank b. cross, philip s. humphrey, j. knox jones, jr. volume 18, no. 1, pp. 1-10 published september 24, 1968 university of kansas lawrence, kansas printed by robert r. (bob) sanders, state printer topeka, kansas 1968 32-3687 the genera of phyllomedusine frogs (anura: hylidae) by william e. duellman one of the most distinctive phyletic lines among the diverse neotropical hylid frogs is composed of a group of 40 species placed in the genus _phyllomedusa_ (funkhouser, 1957) or in two or three different genera (goin, 1961; lutz, 1966). these species differ from all other neotropical hylids by possessing a vertical, instead of horizontal, pupil. the only other hylids having a vertical pupil belong to the papuan genus _nyctimystes_. goin (1961) erroneously stated that _nyctimantis_ and _triprion_ have vertical pupils. although limited information is available on the cytotaxonomy of hylids, the data show that phyllomedusine species have _n_=13 (2_n_=26) chromosomes. _acris_ has _n_=11 (2_n_=22) (cole, 1966). members of the _hyla leucophyllata_, _microcephala_, and _parviceps_ groups have _n_=15 (2_n_=30), _gastrotheca ceratophrys_ has a haploid number of 14, the papuan hylid genus _nyctimystes_ and all but one of the australo-papuan _hyla_ for which the numbers are known have a haploid number of 13, and all other new world hylids studied have _n_=12 (2_n_=24) (duellman and cole, 1965; duellman, 1967). cei (1963) and cei and erspamer (1966) noted that phyllomedusine frogs differ notably from other neotropical hylids on the basis of the amines and polypeptides in the skin. all species of phyllomedusines deposit their eggs in a gelatinous mass on leaves or branches above water. although this type of egg deposition is characteristic of some rhacophorines and apparently all centrolenids, it is known among hylids only in the phyllomedusines and in two species of _hyla_. the distinctive combination of morphological, physiological, chromosomal, and behavioral characteristics is strongly suggestive that these frogs represent an early phyletic divergence within the hylidae. günther (1859) proposed the familial name phyllomedusidae for _phyllomedusa bicolor_ (boddaert). i suggest the recognition of the group as a subfamily. the following classification of the phyllomedusines is based on my own knowledge of the middle american and some south american species and on evidence from the literature on those south american species with which i am not personally familiar. subfamily phyllomedusinae günther, 1859 phyllomedusidae günther 1859 [type genus, _phyllomedusa_ wagler, 1830]. _definition._--moderately small to large hylids having vertical pupils, _n_=13 (2_n_=26) chromosomes, skin containing large amounts of powerful bradykinin-like and physalaemin-like polypeptides, eggs suspended from vegetation above water, and tadpoles have a ventral spiracle sinistral to midline. _range._--low and moderate elevations in south and middle america, including trinidad, from northern argentina and northwestern ecuador to veracruz and southern sonora, méxico. _content._--three genera, one of which probably is composite. genus =agalychnis= cope, 1864. _agalychnis_ cope, 1864 [type species, _hyla moreletii_ duméril, 1853, by subsequent designation]. _definition._--fingers and toes at least half webbed; terminal discs large; first toe shorter than second and not opposable to others; skin smooth, lacking osteoderms; parotoid glands, if present, poorly developed and diffuse; palpebral membrane reticulate (except in _a. calcarifer_); iris red or yellow; skull shallow, depth less than 40 per cent of length; nasals large; frontoparietal fontanelle large; quadratojugals reduced; prevomerine teeth present. _range._--central veracruz and northern oaxaca, méxico, southeastward through central america to northwestern ecuador; one species disjunct in amazonian ecuador. _content._--eight species [synonyms in brackets]: _annae_ (duellman, 1963); _calcarifer_ boulenger, 1902; _callidryas_ (cope, 1862) [_helenae_ cope, 1885; _callidryas taylori_ (funkhouser, 1957)]; _craspedopus_ (funkhouser, 1957); _litodryas_ (duellman and trueb, 1967); _moreleti_ (duméril, 1853) [_holochroa_ (salvin, 1861)]; _saltator_ taylor, 1955; _spurrelli_ boulenger, 1913. _remarks._--savage and heyer (1967) provided evidence that _a. callidryas taylori_ (funkhouser) and _a. helenae_ cope were junior synonyms of _a. callidryas_ (cope). genus =pachymedusa=, new genus type species, _agalychnis dacnicolor_ cope, 1864. _definition._--fingers and toes having basal webs and lateral fringes; terminal discs large; first toe shorter than second and not opposable to others; skin smooth or shagreened, lacking osteoderms; paratoid glands present, diffuse; palpebral membrane reticulate; iris golden yellow with black reticulations; skull deep, depth more than 50 per cent of length; nasals large; frontoparietal fontanelle moderately large; quadratojugal robust; prevomerine teeth present. _range._--pacific slopes and lowlands from southern sonora to the isthmus of tehuantepec, méxico. _content._--monotypic: _dacnicolor_ cope, 1864 [_alcorni_ taylor, 1952]. _remarks._--the generic name is derived from the greek _pachy_ meaning thick and the greek _medousa_ (latin, _medusa_) in reference to _phyllomedusa_; the sense implied is the heavy body of _pachymedusa dacnicolor_. genus =phyllomedusa= wagler, 1830 _phyllomedusa_ wagler, 1830 [type species, _rana bicolor_ boddaert, 1772]. _pithecopus_ cope, 1866 [type species, _phyllomedusa azurea_ cope, 1862 (=_phyllomedusa hypochondrialis_ daudin, 1803), by original designation]. _hylomantis_ peters, 1872 [type species _hylomantis aspera_ peters, 1872, by monotypy]. _phrynomedusa_ miranda-ribeiro, 1923 [type species, _phrynomedusa fimbriata_ miranda-ribeiro, 1923, by subsequent designation]. _bradymedusa_ miranda-ribeiro, 1926 [type species, _bradymedusa moschada_ miranda-ribeiro, 1926 (=_phyllomedusa rohdei_ mertens, 1926) by subsequent designation]. _definition._--fingers and toes having greatly reduced webbing or lacking webs; terminal discs small; first toe shorter than, equal to, or longer than second, opposable or not; skin smooth or rugose having osteoderms or not; parotoid glands present, in most species, usually distinct and elevated; palpebral membrane not reticulate; iris uniformly silvery white to orange-bronze with black reticulations; skull moderate to deep, depth more than 38 per cent of length; nasals moderately small; frontoparietal fontanelle present, variable in size; quadratojugal reduced in some species; prevomerine teeth present or absent. _range._--low and moderate elevations in south america east of the andes from the caribbean (including trinidad) to northern argentina; costa rica and panamá in central america. _content._--thirty-one species [synonyms in brackets]: _aspera_ (peters, 1872); _ayeaye_ (b. lutz, 1966); _bahiana_ a. lutz, 1925; _bicolor_ (boddaert, 1772) [_scleroderma_ cope, 1868]; _blombergi_ funkhouser, 1957; _boliviana_ boulenger, 1902; _buckleyi_ boulenger, 1882; _burmeisteri burmeisteri_ boulenger, 1882; _burmeisteri distincta_ b. lutz, 1950; _centralis_ bokermann, 1965; _cochranae_ bokermann, 1966; _coelestis_ (cope, 1874); _edentula_ andersson, 1945; _feltoni_ shreve, 1935; _fimbriata_ (miranda-ribeiro, 1923) [_appendiculata_ a. lutz, 1925]; _guttata_ a. lutz, 1925; _hypochondrialis_ (daudin, 1803) [_azurea_ cope, 1862; _megacephala_ (miranda-ribeiro, 1926)]; _iheringi_ boulenger, 1885; _lemur_ boulenger, 1882; _loris_ boulenger, 1912; _medinae_ funkhouser, 1962; _nicefori_ barbour, 1926; _orcesi_ funkhouser, 1957; _pailona_ shreve, 1959; _perlata_ boulenger, 1882; _rohdei_ mertens, 1926 [_moschada_ (miranda-ribeiro, 1926)]; _sauvagei_ boulenger, 1882 [_rickettsii_ günther, 1897]; _tarsius_ (cope, 1868); _tomopterna_ (cope, 1868) [_palliata_ peters, 1872]; _trinitatis_ mertens, 1926, _vaillanti_ boulenger, 1882, _venusta_ duellmann and trueb, 1967. _remarks._--_phyllomedusa_ includes 1) a series of large species (_bicolor-burmeisteri_) showing progressive specialization of the feet; 2) a series of small species having grasping feet (_ayeaye_, _centralis_, _cochranae_, _guttata_, _hypochondrialis_, and _rohdei_); 3) a series of small, relatively unspecialized species (_lemur_, _loris_, and _medinae_); and 4) several other species of questionable affinities. lutz (1966) resurrected cope's (1866) _pithecopus_ for 12 species (_ayeaye_, _boliviana_, _burmeisteri_, _coelestis_, _hypochondrialis_, _nicefori_, _rohdei_, _sauvagei_, _tarsius_, _tomopterna_, _trinitatis_, and _vaillanti_). adequate material is not available for detailed study of all south american species; consequently, a firm classification cannot be established at this time. nevertheless, it is obvious that lutz's arrangement is unnatural. if subsequent investigations show, as seems likely, that the small specialized phyllomedusines are a natural phyletic unit, the generic name _pithecopus_ is available. however, species such as _boliviana_, _burmeisteri_, _nicefori_, and _trinitatis_ do not belong in _pithecopus_. as noted by funkhouser (1962), the small, relatively unspecialized species (_lemur_, _loris_, and _medinae_) form a natural group; possibly this group should be accorded generic recognition. until more evidence on the interspecific relationships is acquired, the maintenance of the current classification is desirable. discussion noble (1931) considered the species of _phyllomedusa_ having opposable digits, reduced terminal discs, and no webbing to be advanced and such species as _agalychnis moreleti_, _calcarifer_, and _spurrelli_ to be primitive. funkhouser (1957) followed noble's suggestion and attempted to explain the evolution of the species of _phyllomedusa_ (_sensu lato_) by assuming that they evolved from an advanced _hyla_-like ancestor. therefore, she placed those species having large, fully webbed hands and feet near the base of her phylogenetic scheme and hypothesized that evolutionary sequences involved stages of reduction and eventual loss of webbing, followed by the development of grasping toes. such an evolutionary history is highly unlikely. the _agalychnis_ phyletic line has one kind of specialization for an arboreal existence. it is contrary to evolutionary theory that a specialized group would evolve into a generalized form and then evolve new kinds of specializations to meet the needs imposed by the same environmental conditions affecting the earlier specialized group. a more reasonable hypothesis is that the evolution of opposable digits took place in a phyletic line that had as its ancestral stock a frog with generalized hands and feet. if this assumption is correct, _phyllomedusa_ and _agalychnis_ represent different phyletic lines; each exhibits divergent modes of adaptation for arboreal habits, whereas _pachymedusa_ probably remains relatively little changed from the basic phyllomedusine stock. on the basis of modern distribution and areas of diversification alone (no fossils are known), it is evident that _phyllomedusa_ underwent its adaptive radiation in south america, _agalychnis_ evolved in central america, and _pachymedusa_ ended up in western méxico. if we follow the matthewsian concepts of the american herpetofauna outlined by dunn (1931) and modified by schmidt (1943) and stuart (1950), _pachymedusa_ represents a "hanging-relict" of a group that moved southward. according to savage's (1966) interpretation of the origins and history of the american herpetofauna, _agalychnis_ and _pachymedusa_ are members of the mesoamerican fauna, and _phyllomedusa_ is part of the neotropical fauna. perhaps the phyllomedusines arose in south america; from there a primitive stock spread northward and survived as _pachymedusa_ in méxico, whereas the stock in central america and south america evolved into _agalychnis_ and _phyllomedusa_, respectively. evidently the primitive phyllomedusines evolved the habit of arboreal egg deposition and a walking gait; the latter is best developed in the small, highly specialized species of _phyllomedusa_ (lutz, 1966). probably the other divergent arboreal adaptations resulted from environmental stresses and competition. the generalized _pachymedusa_ inhabits relatively dry areas characterized by low forest. throughout its range it coexists with no more than five other arboreal hylids. the species of _agalychnis_ live in rain forests and humid montane forests. in any given area one species of _agalychnis_ occurs sympatrically with no more than a dozen other arboreal hylids. with few exceptions the species of _agalychnis_ are more arboreal in their habits than are other hylids. the species of _phyllomedusa_ live in the same kinds of habitats as do those of _agalychnis_, but throughout the ranges of most of the species of _phyllomedusa_ the diversity of arboreal hylids is much greater than in central america. in the upper amazon basin as many as 35 hylids occur sympatrically. many groups of _hyla_ in this area (for example, the _hyla boans_ and _hyla marmorata_ groups) are equally as arboreal in their habits as are the species of _agalychnis_ in central america. conceivably, competition within this array of tree frogs resulted in selection for modification of the extremities, thereby bringing about a different mode of climbing in _phyllomedusa_. the walking gait already present in phyllomedusines provided a source for further modification, which resulted in the development of opposable digits and the associated lemuroid manner of climbing. the known life histories of most species of _phyllomedusa_, all species of _agalychnis_, and that of _pachymedusa_ are similar. characteristically the tadpoles are generalized pelagic types that develop in ponds, but at least some of the small specialized _phyllomedusa_ in southeastern brazil have stream-adapted tadpoles with funnel-shaped mouths (cochran, 1955; bokermann, 1966). knowledge of the life histories of the other species of _phyllomedusa_ should aid in the interpretation of the phylogenetic relationships of the several groups of frogs now assigned to that genus. acknowledgments i am grateful to linda trueb who provided the osteological data included, and who helped me in formulating some of the ideas expressed in the discussion. this paper is a result of investigations on hylid frogs supported by the national science foundation (nsf-gb-5818). literature cited bokermann, w. c. a 1966. a new _phyllomedusa_ from southeastern brazil. herpetologica, 22:293-297. cei, j. m 1963. some precipitin tests and preliminary remarks on the systematic relationships of four south american families of frogs. bull. serological mus., 30:4-6. cei, j. m. and v. erspamer 1966. biochemical taxonomy of south american amphibians by means of skin amines and polypeptides. copeia, no. 1:74-8. cochran, d. m 1955. frogs of southeastern brazil. bull. u. s. natl. mus., 206; xvi + 423 pp. cole, c. j 1966. the chromosomes of _acris crepitans blanchardi_ harper (anura: hylidae). copeia, no. 3:578-580. cope, e. d 1864. contributions to the herpetology of tropical america. proc. acad. nat. sci. philadelphia, 16:166-181. 1866. on the structures and distribution of the genera of arciferous anura. jour. acad. nat. sci. philadelphia, ser. 2, 6:67-112. duellman, w. e 1967. additional studies on chromosomes of anuran amphibians. syst. zool., 16:38-43. duellman, w. e. and c. j. cole 1965. studies of chromosomes of some anuran amphibians (hylidae and centrolenidae). syst. zool., 14:139-143. duméril, a. h 1853. memoire sur les batraciens anoures de la famille des hylaeformes ou rainettes comprenant la description d'un genre nouveau et de onze espèces nouvelles. ann. sci. nat., ser. 3, 19:135-179. dunn, e. r 1931. the herpetological fauna of the americas. copeia, no. 3:106-119. funkhouser, a 1957. a review of the neotropical tree-frogs of the genus _phyllomedusa_. occas. papers nat. hist. mus., stanford univ., 5:1-90. 1962. a new _phyllomedusa_ from venezuela. copeia, no. 3:588-590. goin, c. j 1961. synopsis of the genera of hylid frogs. ann. carnegie mus., 36:5-18. günther, a. c. l. g. "1858" [1859], catalogue of die batrachia salientia in the collection of the british museum. taylor and francis, london, xvi + 160 pp., 12 pls. lutz, b 1966. _pithecopus ayeaye_, a new brazilian hylid with vertical pupils and grasping feet. copeia, no. 2:236-240. miranda-ribeiro, a. de 1923. as phyllomedusas do museu paulista. bol. mus. nac, rio de janeiro, 1:3-6. 1926. notas para servirem ao estudo dos gymnobatrachios (anura) brasileiros. arch mus. nac., rio de janeiro, 27:1-227, pls. 1-22. noble, g. k 1931. the biology of the amphibia. mcgraw-hill, new york, 577 pp. peters, w. c. h 1872. über eine, zwei neue gattungen enthaltende, sammlung von batrachiern des hrn. dr. o. wucherer aus bahia, so wie übereinige neue oder weniger bekannte saurier. monatsb. akad. wiss. berlin, 1872:768-776. savage, j. m 1966. the origins and history of the central american herpetofauna. copeia, no. 4:719-766. savage, j. m. and w. r. heyer 1967. variation and distribution in the tree-frog genus _phyllomedusa_ in costa rica, central america. beitr. neotrop. fauna, 5:111-131. schmidt, k. p 1943. corollary and commentary for "climate and evolution." amer. midl. nat., 30:241-253. stuart, l. c 1950. a geographic study of the herpetofauna of alta verapaz, guatemala. contr. lab. vert. biol., univ. michigan, 45:1-77, pls. 1-9. wagler, j. g 1830. näturliches system der amphibien, mit vorangehender classification der saügethiere und vögel. münchen, vi + 354 pp., 9 pls. _transmitted april 18, 1968._ university of kansas publications museum of natural history volume 17, no. 10, pp. 493-502, 3 figs. october 27, 1966 systematic status of a south american frog, allophryne ruthveni gaige by john d. lynch and howard l. freeman university of kansas lawrence 1966 university of kansas publications, museum of natural history editors: e. raymond hall, chairman, henry s. fitch, frank b. cross volume 17, no. 10, pp. 493-502, 3 figs. published october 27, 1966 university of kansas lawrence, kansas printed by robert r. (bob) sanders, state printer topeka, kansas 1966 31-5378 systematic status of a south american frog, allophryne ruthveni gaige by john d. lynch and howard l. freeman gaige (1926) described _allophryne ruthveni_ as a new genus and species of diminutive bufonid from british guiana. noble (1931) considered _a. ruthveni_ to be a toothless relative of _centrolenella_ and placed the genus in the hylidae. gallardo (1965) suggested that _allophryne_ is a leptodactylid of uncertain affinities. other references to the monotypic genus have consisted only of a listing of the name or of its inclusion in a key. to date the holotype and one paratype (both females) have been reported (gaige, 1926), and the family position of the genus remains unsettled. a male of _allophryne ruthveni_ is among the amphibians and reptiles collected in southern british guiana by william a. bentley in january, 1962, and deposited in the museum of natural history at the university of kansas (ku). four additional specimens (females) are in the american museum of natural history; only one of the latter has definite locality data. _acknowledgments._--we are grateful to dr. ernest e. williams, museum of comparative zoology (mcz) and dr. richard g. zweifel, american museum of natural history (amnh) for the loan of specimens. we are further indebted to dr. zweifel for permission to clear and stain one specimen. dr. william e. duellman and linda trueb offered many constructive criticisms. miss trueb executed the drawings of the skull and finger bones. mr. martin wiley provided x-ray photographs of _allophryne_. methods and materials six of the seven known specimens were available for study. measurements were taken in the manner described by duellman (1956). one specimen was cleared and stained, using the technique of davis and gore (1936), in order to study the skeleton. x-ray photographs were made of another specimen for comparison. _specimens examined._--six, as follows: british guiana, _dist. demarara_: marudi creek, amnh 44749; _dist. equibo_: tumatumari, mcz 11790 (paratype); _dist. rupununi_ (_berbice_): wai wai country, n of acarahy mountains, west of new river (2°n, 58°w), ku 69890. also, 3 specimens from "probably british guiana," amnh 70108-10 (70110 cleared and stained). systematic account the availability of additional material and the new information pertaining to osteology permit an amplification of gaige's (1926) description. genus ~allophryne~ gaige _allophryne_ gaige, occas. papers mus. zool., univ. michigan, 176:1, oct. 14, 1926. crawford, annals carnegie mus., 21(1):29, 32, nov. 14, 1931. noble, the biology of the amphibia. mcgraw-hill, p. 510, 1931. ruthven, herpetologica, 1:3, july 11, 1936. gallardo, papéis avulsos, 17:79, jan. 1, 1965. _type species._--_allophryne ruthveni_ gaige. _diagnosis and definition._--a genus of diminutive frogs; vomers, maxillae, and premaxillae edentate; skin of head strongly anchored to connective tissue on cranium; prepollical spine absent in males; disk of third finger larger than tympanum, smaller than eye; no humeral hook in either sex; ilia extending anteriorly beyond sacral expansions; adults attaining snout-vent length of 31 mm.; male having darkened external subgular vocal sac; skin of dorsum pustulate. ~allophryne ruthveni~ gaige _allophryne ruthveni_ gaige, occas. papers mus. zool., univ. michigan, 176:1-3, pl. i, oct. 14, 1926. crawford, annals carnegie mus., 21(1):32, nov. 14, 1931. ruthven, herpetologica, 1:3, july 11, 1936. barbour and loveridge, bull. mus. comp. zool., 96(2):64, feb., 1946. peters, occas. papers mus. zool., univ. michigan, 539:10, sept. 19, 1952. _holotype._--university of michigan museum of zoology 63419, adult female, from tukeit hill, below kaiteur falls, equibo district, british guiana; obtained in may, 1924, by e. n. clarke. _diagnosis._--fingers free; toes two-thirds webbed; no supernumerary tubercles on soles or palms; no tarsal fold; elongate anal sheath, anal opening on lower surface of thighs; head broad, interorbital space 2.5 times width of upper eyelid; snout subacuminate in dorsal profile, strongly sloping in lateral profile; tympanum visible in males, concealed in females; venter areolate. _external morphology._--(fig. 1) _additional features not mentioned in diagnoses_: head wider than long, about as wide as body; supratympanic fold present; canthus rostralis rounded, loreal region slightly concave, nearly vertical; nostril at tip of snout; pupil horizontal; no teeth on maxillary, premaxillary, or vomer; tongue small, round, thick, not notched behind, free posteriorly for one-sixth of length; choanae large, only partly visible from directly below; males having darkened subgular vocal sac; vocal slits present in male. axillary membrane lacking or but slightly developed; no tubercles or ridge under forearm; two palmar tubercles; subarticular tubercles small, simple, round, flattened; tips of fingers slightly expanded, t-shaped, with prominent transverse groove; first finger shorter than second (stated as longer than second in diagnosis by gaige, 1926:2); folds extending laterally from anus for a short distance, then downward to venter of thighs; no appendage on heel, no inner or outer tarsal folds or tubercles; inner metatarsal tubercle oval, about twice as long as wide; outer metatarsal tubercle nearly absent; no supernumerary tubercle on sole; subarticular tubercles on foot small, round, simple, and diffuse; toes t-shaped, slightly wider than digit; toes about two-thirds webbed (fig. 1d). skin of venter coarsely areolate; skin of flanks, throat, chest, undersurfaces of arms, tibia, tarsi, dorsal surfaces of thighs, tarsi, hands, and feet smooth; skin of dorsal surfaces of tibia, forearm, back, and top and sides of head having large horny pustules (sharply spinous in male). [illustration: fig. 1. _allophryne ruthveni_, male (ku 69890); (_a_) dorsum. (_b_) thenar view of right hand. (_c_) lateral profile of head. (_d_) plantar view of right foot. × 3.5.] _color._--dorsum gray with irregular network of black lines and elongate blotches; flanks and labial region black with large white ocelli; dorsal surfaces of limbs gray, marked as follows: two large, elongate white spots on each thigh, concealed white spot on base of upper arm, black-edged gray transverse bars on forearms and shanks, white spot on each knee and elbow; ventral surfaces pale gray; black-edged white spot on ventral surface of thigh on each side of anal opening; chin and throat dark gray with white spots; vocal sac in male black (fig. 1a and c). gaige (1926) briefly described the color, which conforms to the above in all particulars. the paratype (mcz 11790) has lost the gray color after 40 years in preservation; now (1966) the ground-color is cream-brown, and the dorsal spotting, noted by gaige as being black, is now brown. the spots on the feet, tarsi, knees, thighs, flanks and upper arm are white in preservative, but in life possibly were red or yellow. these colors usually fade to white in preservative. red or yellow spots are common aposematic colors in frogs. _variation._--eight measurements were taken on each specimen and four ratios were computed; these are summarized in table 1. gaige's illustration of the holotype shows that it has a greatly reduced pattern, whereas the paratype and three of the other five known specimens have relatively large and numerous spots. the male (ku 69890) and one female (amnh 70108) have a reduced pattern intermediate between that of the holotype and the four other specimens. table i.--variation in measurements and proportions of allophryne ruthveni. (ranges in parentheses below means.) --------------------------+----------+---------------- character | male (1) | females (5) --------------------------+----------+----------------snout-vent (in mm.) | 20.6 | 23.6 | | (18.4-31.0)[a] | | tibia/snout-vent | 0.43 | 0.43 | | (0.41-0.47) | | tympanum/head width | 0.12 | 0.15 | | (0.14-0.16) | | eyelid/interorbital space | 0.55 | 0.53 | | (0.49-0.56) | | tympanum/eye length | 0.40 | 0.46 | | (0.42-0.50) --------------------------+----------+----------------[footnote a: holotype is reported to be 31 mm. snout-vent length (gaige, 1926). the largest measured by us was 26.2 mm. snout-vent.] the dorsal spinules are most pronounced and extensive on the male (fig. 1) and less so in all other specimens examined. the illustration of the holotype suggests that it has equally prominent, but fewer, spinules (gaige, 1926). the holotype, a gravid female, is the largest known specimen (31 mm., snout-vent length). another gravid female (amnh 70108) has a snout-vent length of 26.2 mm. _distribution._--all known specimens have been found in the foothills of the northeastern face of the guiana massif in british guiana. family position the following characters of _allophryne_ are those generally held to be useful in determining family relationships: 1. presacral vertebrae procoelus, eight in number. 2. parahyoid absent. 3. free ribs lacking. 4. bidder's organ absent. 5. intercalary cartilages present in digits; phalangeal formulae 3-3-4-4 and 3-3-4-5-4. 6. coccyx articulating with sacrum by two condyles. 7. tarsal bones not fused. 8. pectoral girdle arciferal. 9. epicoracoidal horns present, free. 10. terminal phalanges t-shaped. 11. sacrum procoelus and diapophyses expanded. 12. maxillae, premaxillae, and prevomers edentate. 13. cranial roofing bones well ossified. griffiths (1959) accorded considerable taxonomic weight to the presence or absence of epicoracoidal horns in showing relationships among the genera placed in the brachycephalidae [= atelopodidae; dendrobatidae; and leptodactylidae (in part)] by noble (1931). _allophryne_ possesses well-developed, free epicoracoidal horns, such as those found in the hylidae, centrolenidae, leptodactylidae and bufonidae. the presence of intercalary elements in the digits is characteristic of the centrolenidae, hylidae, phrynomeridae, pseudidae, and the rhacophorine ranids (including the hyperoliidae). this element is bony in the pseudids and cartilaginous in the other families. phrynomerids and rhacophorine ranids lack epicoracoidal horns and have firmisternal pectoral girdles. centrolenids are small, delicate, arboreal frogs having poorly ossified skulls and fused tarsal bones, but agree with _allophryne_ in having t-shaped terminal phalanges. [illustration: fig. 2. dorsal (_a_) and lateral (_b_) views of distal phalanges of third finger of _allophryne_. × 40.] only the presence of intercalary cartilages (fig. 2) suggests relationship of _allophryne_ to the hylidae. the t-shaped terminal phalanges suggest affinities with centrolenids, elutherodactyline leptodactylids, or certain "brachycephalid" frogs. griffiths (1959) clearly showed that noble's brachycephalidae was a polyphyletic assemblage. no hylid genus is edentate, and none has either t-shaped terminal phalanges or the unusual dorsal spinules. perhaps the presence of intercalary cartilages is not indicative of relationship but instead is a parallelism (or convergence) in _allophryne_ and genera of the centrolenidae. cranial osteology the skull of _allophryne_ (fig. 3) is distinctive among anurans; it does not closely resemble the skulls of either hylids or centrolenids, both of which have generally more delicate (except for casque-headed hylids, such as _corythomantis_, _diaglena_, _osteocephalus_, _triprion_) and generalized skulls. _allophryne_ on the other hand has a strongly ossified central region (cranial roofing bones and sphenethmoid complex) and a weak peripheral zone. the peripheral elements are reduced (maxilla, pterygoid, and squamosal) or absent (quadratojugal), whereas the frontoparietals, nasals, sphenethmoid, proötics, and exoccipitals form a compact central zone. an elongate frontoparietal fontanelle is present. [illustration: fig. 3. dorsal view of skull of _allophryne_ (amnh 70110). × 12.] dorsally (fig. 3), the premaxillae are not visible. the proportionally gigantic septomaxillae are visible anterior to the nasals. the moderate-sized nasals are separated medially and in broad contact with the ethmoid posteriorly. the palatine process of the nasal does not meet the frontal process of the maxilla. a large frontoparietal fontanelle is evident between the frontoparietals. the tegmen tympani are much reduced and maintain only cartilaginous contact with the posterior arms of the squamosals. the foramen magnum, occipital condyles, and exoccipitals show no unusual features. the _pars facialis_ and frontal process of the maxilla are greatly reduced. the maxilla and premaxilla are articulated. the high, narrow alary processes of the premaxillae extend dorsally about two-thirds of the height of the snout. a cartilaginous internasal septum is illustrated (fig. 3), but sectioning is necessary to determine the true nature and extent of this element. ventrally, the skull lacks palatines. the maxillae, premaxillae, and prevomers are edentate. the parasphenoid is large with relatively short, stout alary (lateral) processes. the sphenethmoid is extensive in ventral aspect and forms the major supporting structure in the anterior part of the skull. the pterygoid has a broad articulation with the maxilla, a tenuous contact with the squamosal, but is not attached to the proötic. the anterior (zygomatic) process of the squamosal is greatly reduced (only about one-third the length of the posterior process). discussion the skull of _allophryne_ is definitely non-hylid. most of the post-cranial features do not help to clarify relationships. _allophryne_ shares several osteological features with the dendrobatidae: t-shaped terminal phalanges, general cranial morphology and procoelus vertebrae. but, the dendrobatids possess firmisternal pectoral girdles and lack epicoracoidal horns. also, no dendrobatid has intercalary elements in the digits. we are, therefore, left with a taxonomic enigma. in one or more characters generally regarded as important, _allophryne_ differs from all presently defined families of frogs. the hylidae and dendrobatidae are the only currently recognized families in which the genus might be placed. the function and taxonomic importance of the large septomaxillae are unknown and are probably associated with the modification of the sphenethmoid-prevomer area. a more detailed study of the cranial osteology of _allophryne_, especially the structural relationships of the sphenethmoid-prevomer area may elucidate the relationships of _allophryne_. the relationships of _allophryne_ cannot be understood without a re-analysis of some of the features used as major criteria in frog classification (the nature of an intercalated cartilage; the nature of the sternal complex; the relative value of cranial osteology; the vertebral structure; and the thigh musculature). some of these features have been investigated by other workers, most notably griffiths, but others have not and need re-examination. a re-analysis of some of the major criteria used in frog classification is in progress (callison, lynch, and trueb) and upon completion of that study we think the relationships of _allophryne_ will become apparent. a more comprehensive study of the cranial anatomy of certain hylids, leptodactylids, dendrobatids, and atelopodids along with that of _allophryne_ is needed to clarify the relationships of _allophryne_, and might indicate that the recognition of a fifth family is necessary. conclusion among currently recognized families of frogs, _allophryne_ is least different from the hylidae although it is our opinion that inclusion of this genus in the hylidae probably represents an unnatural classification. however, the present evidence suggesting that _allophryne_ should be in another family is less convincing than evidence suggesting it should be in the hylidae. we tentatively place _allophryne_ in the hylidae. literature cited davis, d. d. and gore, u. r. 1936. clearing and staining skeletons of small vertebrates. fieldiana: technique, 4:1-16. duellman, w. e. 1956. the frogs of the hylid genus _phrynohyas_ fitzinger, 1843. misc. publs. mus. zool., univ. michigan, 96:1-47, february 21. gaige, h. t. 1926. a new frog from british guiana. occas. papers mus. zool., univ. michigan, 176:1-3, october 14. gallardo, j. m. 1965. a propósito de los leptodactylidae (amphibia anura). papéis avulsos, 17:77-87, january 1. griffiths, i. 1959. the phylogeny of _sminthillus limbatus_ and the status of the brachycephalidae (amphibia: salientia). proc. zool. soc. london, 132:457-87, may. noble, g. k. 1931. the biology of the amphibia. mcgraw-hill, new york, vii + 577 pp. _transmitted august 2, 1966._ 31-5378 * * * * * transcriber's notes italicized text is shown within _underscores_. bold text is shown within ~tildes~. table 1 and figs. 2 and 3 have been moved slightly to avoid breaking up the paragraphs of text. university of kansas publications museum of natural history volume 13, no. 8, pl. 25, figs. 1-2, pp. 349-357 april 27, 1961 descriptions of two species of frogs, genus ptychohyla studies of american hylid frogs, v by william e. duellman university of kansas lawrence 1961 university of kansas publications, museum of natural history editors: e. raymond hall, chairman, henry s. fitch, robert w. wilson volume 13, no. 8, pl. 25, figs. 1-2, pp. 349-357 published april 27, 1961 university of kansas lawrence, kansas printed in the state printing plant topeka, kansas 1961 28-6442 descriptions of two new species of frogs, genus ptychohyla studies of american hylid frogs, v by william e. duellman field studies on hylid frogs in southern méxico and northern central america have resulted in the collection of numerous specimens of _ptychohyla_, a genus of hylid frogs heretofore poorly represented in museum collections. experience with the living frogs in their natural habitats has been helpful in defining the species and in formulating ideas concerning their relationships. taylor (1944) proposed the generic name _ptychohyla_ for a new species of frog, _ptychohyla adipoventris_ [= _ptychohyla leonhard-schultzei_ (ahl)--_fide_ duellman, 1960] from agua del obispo, guerrero. taylor defined the genus as having large ventrolateral glands and horny nuptial spines in males. stuart (1954:169) discussed the generic characters and pointed out that both the ventrolateral glands and horny nuptial spines were seasonal in their development, being found only in breeding males. stuart then went on to describe _ptychohyla schmidtorum_, a species characterized by the absence of horny nuptial spines in breeding males. my investigations of these frogs have revealed the presence of two groups of species. in both groups breeding males have large ventrolateral glands, but the two groups are easily separated by four characters. the first group contains, among others, _ptychohyla leonhard-schultzei_, _euthysanota_, _spinipollex_, and another species in the mesa central of chiapas to which i tentatively apply the name _ptychohyla macrotympanum_ (tanner), 1957. this group of species is characterized by horny nuptial spines in breeding males, presence of a tarsal fold, a call consisting of a single long note, and tadpoles having lips not greatly expanded. the second group, as recognized here, is characterized by the absence of horny nuptial spines in breeding males, lack of a tarsal fold, a call consisting of a series of short notes, and tadpoles having greatly expanded lips. in this group belong _ptychohyla schmidtorum_ and the two species described below. only the descriptions of the new species are given in this paper; detailed comparisons, descriptions of osteological features, analyses of calls, and discussions of relationships are reserved for a forthcoming review of the entire genus. in the spring of 1959, collections of amphibians and reptiles were made in the cloud forests on the northern slopes of the sierra madre oriental in northern oaxaca. among the hylids found, two specimens of a heretofore unnamed species of _ptychohyla_ have brilliant red flash-colors on the groin and thighs; in allusion to these fiery colors i propose that this species be named: ~_ptychohyla ignicolor_~ new species (plate 25, fig. 1) _holotype._--university of michigan museum of zoology no. 119603, from a stream 6 kilometers south of vista hermosa, oaxaca, méxico (1865 meters); obtained on march 31, 1959, by thomas e. moore. original number wed 14159. _paratype._--ummz 119602 from vista hermosa, oaxaca (1500 meters); obtained on march 30, 1959, by william e. duellman. _diagnosis._--a species of the _schmidtorum_-group of _ptychohyla_ differing from other known members of the group in having the diameter of the tympanum less than one-half the diameter of the eye, no white spot below the eye, no lateral light stripe, bright green dorsum in life and red flash-colors on groin and thighs. _description of holotype._--adult male having a snout-vent length of 30.0 mm.; tibia length, 14.6 mm.; tibia length/snout-vent length, 48.7 per cent; foot length (measured from proximal edge of inner metatarsal tubercle to tip of longest toe), 12.3 mm.; head length, 9.2 mm.; head length/snout-vent length, 32.3 per cent; head width, 9.3 mm.; head width/snout-vent length, 31.0 per cent; diameter of eye, 3.2 mm.; diameter of tympanum, 1.3 mm.; tympanum/eye, 40.6 per cent. snout in lateral profile square, in dorsal profile obtusely rounded; canthus pronounced; loreal region slightly concave; lips moderately flaring; top of head flat; nostrils protuberant; internarial distance, 2.8 mm.; interorbital distance, 3.3 mm., much broader than width of eyelid, 2.8 mm. a heavy dermal fold from posterior corner of eye above tympanum to insertion of forelimb, covering upper edge of tympanum; tympanum elliptical, its greatest diameter equal to its distance from eye. forearm robust with a distinct fold on wrist; pollex moderately enlarged without nuptial spines; second and fourth fingers equal in length; subarticular tubercles round; none is bifid; disc of third finger slightly larger than tympanum; no web between first and second fingers; vestige of web between other fingers. heels overlap when hind limbs adpressed; tibiotarsal articulation extends to anterior corner of eye; no tarsal fold; inner metatarsal tubercle large, flat, and elliptical; outer metatarsal tubercle near inner one and triangular; subarticular tubercles round; length of digits from shortest to longest 1-2-5-3-4; toes about one-half webbed; discs smaller on toes than on fingers. anal opening directed posteriorly at upper level of thighs; no anal flap; pair of large tubercles below anal opening; small tubercles ventral and lateral to these. skin of dorsum and ventral surfaces of limbs smooth, that of throat and belly granular. ventrolateral glands noticeably thickened, extending from axilla nearly to groin and only narrowly separated medially on chest. skin of anterior part of chin thickened and glandular. tongue cordiform, shallowly notched behind and only slightly free posteriorly; vomerine teeth 0-3, situated on rounded elevations between somewhat larger, round inner nares; openings to vocal sac large, one situated along posterior margin of each mandibular ramus. plate 25 [illustration: fig. 1. paratype of _ptychohyla ignicolor_ (ummz 119602). × 3.] [illustration: fig. 2. holotype of _ptychohyla chamulae_ (ku 58063). × 3.] color (in alcohol) dull brown above with irregular dark brown blotches; dorsal surfaces of limbs brown with narrow darker brown transverse bars; posterior surfaces of thighs cream-color with brown spots and mottling; groin and dorsal surfaces of first and second toes white; belly cream-colored; glandular areas orange-brown; chest and chin having black spots. ventral surfaces of hind limbs and first toes cream-colored; undersides of other toes and soles of feet brown. color (in life) uniform bright green above; venter pale creamy yellow; anterior and posterior surfaces of thighs, ventral surfaces of shanks, anterior surfaces of tarsi and upper proximal surfaces of first three toes red; iris pale golden color. the paratype is an adult male, having a snout-vent length of 26.3 mm., and agrees with the holotype in proportions. the ventrolateral glands are less extensive and the chin less spotted than in the holotype. _comparisons_: both _ptychohyla schmidtorum_ and the species described below differ from _p. ignicolor_ in lacking red flash-colors and in having a white spot below the eye. _ptychohyla ignicolor_ also differs in having a small tympanum. as stated above, these species can be distinguished from the rest of the genus by the absence of a tarsal fold and absence of horny nuptial spines in breeding males. _remarks_: the holotype was found on a moss-covered log over a stream in dense cloud forest by day. the paratype was calling at night from a low herb at the edge of a small stream in the cloud forest. nearby a _ptychohyla leonhard-schultzei_ was calling. along two cascading mountain streams in cloud forest on the northern slopes of the mesa central in central chiapas numerous specimens of a distinctive species of _ptychohyla_ were found in association with two species of _hyla_ and two of _plectrohyla_. the first specimen of this new species of _ptychohyla_ was discovered by dale l. hoyt, who found the frog on a rock at midday. at night on august 5, 1960, numerous individuals were found calling from leaves of plants growing on the slopes of the ravine by the streams. none was more than two meters above the ground. tadpoles were found in the fast-flowing stream, where they were holding onto rocks with their mouths. little is known of the herpetofauna of these mountains that are the home of the chamula indians. since the little frog described here comes from the land of the chamulas, i propose that it be named: ~_ptychohyla chamulae_~ new species (plate 25, fig. 2) _holotype._--university of kansas museum of natural history no. 58063, from a stream above (6.2 kilometers by road south) rayón mescalapa, chiapas, méxico (1690 meters); one of a series collected on august 5, 1960, by william e. duellman, dale l. hoyt, and john wellman. original no. wed 17327. _paratypes._--ku nos. 58064-58073 collected with the holotype. _diagnosis._--a species of the _schmidtorum_-group of _ptychohyla_ differing from other known members of the group in having the following combination of characters: diameter of tympanum not noticeably less than half that of eye; white spot below eye; white lateral stripe on body anteriorly; dorsum bright green in life; thighs yellowish brown. _description of holotype._--adult male having snout-vent length of 27.3 mm.; tibia length, 12.8 mm.; tibia length/snout-vent length, 48.7 per cent; foot length (measured from proximal edge of inner metatarsal tubercle to tip of longest toe), 10.8 mm.; head length, 9.2 mm.; head length/snout-vent length, 33.7 per cent; head width, 9.0 mm.; head width/snout-vent length, 30.9 per cent; diameter of eye, 2.8 mm.; diameter of tympanum, 1.4 mm.; tympanum/eye, 50.0 per cent. snout in lateral profile nearly square, slightly rounded above; in dorsal profile bluntly squared; canthus pronounced; loreal region concave; lips thick, rounded, and flaring; nostrils protuberant; internarial distance, 2.3 mm.; top of head flat; interorbital distance, 3.3 mm.; much broader than width of eyelid, 2.4 mm. a thin dermal fold from posterior corner of eye above tympanum to insertion of forelimb, covering upper edge of tympanum; tympanum nearly round, its diameter equal to its distance from eye. forearm slender lacking distinct fold on wrist; a row of low, rounded tubercles on ventrolateral surface of forearm; pollex moderately enlarged without nuptial spines; second and fourth fingers equal in length; subarticular tubercles round, none bifid; discs small, that of third finger noticeably smaller than tympanum; no web between first and second fingers; vestige of web between other fingers. heels overlap when hind limbs adpressed; tibiotarsal articulation reaches to middle of eye; no tarsal fold; inner metatarsal tubercle large, flat, and elliptical; outer metatarsal tubercle slightly more distal than inner, small, and elliptical; subarticular tubercles round; length of digits from shortest to longest 1-2-5-3-4; third and fifth toes webbed to base of disc; fourth toe webbed to base of penultimate phalanx; discs smaller on toes than on fingers. anal opening directed posteriorly at upper level of thighs; no anal flap; pair of large tubercles below anal opening and a slightly smaller pair farther below. skin of dorsum and ventral surfaces of forelimbs and shanks smooth; that of throat, belly, and ventral surfaces of thighs granular. ventrolateral glands well developed, not reaching axilla or groin and broadly separated midventrally. skin of anterior part of chin glandular. tongue cordiform, shallowly notched behind and only slightly free posteriorly; vomerine teeth 2-2, situated on small triangular elevations between large, ovoid inner nares; openings to vocal sac large, one situated along inner posterior edge of each mandibular ramus. color (in alcohol) dark purplish brown on dorsal surfaces of head, body, and shanks; thighs brown above and yellowish tan posteriorly; white stripe extending from below eye above forearm to mid-flank. ventral surfaces creamy white; ventrolateral glands orange-tan flecked with dark brown; edge of lower lip with dark brown spots; narrow white line on upper lip; palms white and soles brown. color (in life) uniform dark bright green above with creamy white bar below eye; lateral stripe silvery white; ventral surfaces deep yellow; posterior surfaces of thighs yellow brown; iris reddish bronze. _variation._--sixteen adult males are available; these have snout-vent lengths of 26.3 to 28.5 mm. (average, 27.6 mm.). the tympanum/eye ratio is 48.2 to 58.6 per cent (average, 53.2 per cent). the number of vomerine teeth varies from four to six. the extent of the ventrolateral glands is variable. in five specimens the glands nearly meet midventrally; in two others the glands include the axillary region; in none do the glands extend into the groin. in other structural details there is no noticeable variation. the greatest variation in color pattern is found in the lateral stripe. the pale spot or bar below the eye is present in all specimens; in one individual there is no lateral stripe; in three the stripe extends posteriorly only to above the forearm, in two to the mid-flank, and in the others to the groin. although all of the males were bright uniform green above when collected at night as they were calling, some changed color later. in these individuals the dorsum became a somewhat paler green with faint irregular yellowish tan blotches. the one available female (ummz 121399) has a snout-vent length of 30.3 mm. and a tympanum/eye ratio of 52.8 per cent, and is colored like the males. the tubercles by the anal opening are placed irregularly and do not consist of two pairs below the opening. there are no ventrolateral glands, glandular area on the chin, or enlarged prepollex. _comparisons._--_ptychohyla chamulae_ resembles _p. schmidtorum_ in color pattern and body proportions, but the ground color of _schmidtorum_ is chocolate brown and not green as in _chamulae_. also, in _schmidtorum_ the webbing and posterior surfaces of the thighs are pale cream-color in preserved specimens as contrasted with tan in _chamulae_. in living _schmidtorum_ the iris is bright red, not reddish bronze as in _chamulae_. the ventrolateral glands in _schmidtorum_ more closely approximate one another midventrally than in _chamulae_. it is conceivable that these populations are subspecifically related; _schmidtorum_ occurs in the same kind of habitat as does _chamulae_, but is known only from the pacific slopes of southeastern chiapas and southwestern guatemala, whereas _chamulae_ is known only from the atlantic slopes of the mesa central in north-central chiapas. both of these species differ from _ptychohyla ignicolor_ in having a relatively larger tympanum, more webbing on the foot, different arrangement of anal tubercles, and different coloration. _description of tadpole._--six tadpoles having fully developed mouth parts have body lengths of 5.5 to 11.9 mm. and total lengths of 17.3 to 44.0 mm. the following description is based on a tadpole (ku 58199) having small hind limbs, a body length of 10.5 mm., and a total length of 39.0 mm. body ovoid, only slightly flattened dorsally and ventrally (fig. 1); body only slightly deeper than wide; eyes directed dorsolaterally and slightly protuberant; nostrils small. tail long and slender; greatest depth of tail-musculature two-thirds greatest depth of tail-fin; tail-musculature extending nearly to tip of tail-fin. mouth directed anteroventrally; thin fleshy lips greatly expanded and forming large suckerlike disc; width of mouth greater than width of snout and nearly as wide as body. outer edge of lips having small papillae; inner surface of mouth smooth; scattered large papillae, seemingly in rows, around teeth and beak (fig. 2). tooth rows 3/3; the upper rows subequal in length; upper rows one and three interrupted medially; lower rows one and two about equal in length to upper rows; third lower row short. upper beak heavy and horn-covered. [illustration: fig. 1. tadpole of _ptychohyla chamulae_ (ku 58199). × 2.5.] [illustration: fig. 2. mouthparts of tadpole of _ptychohyla chamulae_ (ku 58199). × 16.] color (in alcohol) dark brown over entire body and tail-musculature; a white area near base of tail, and a dark streak on anterior one-fourth of tail; tail-fin transparent having brown blotches. _remarks_.--five metamorphosing tadpoles and juveniles (ku 58074, 58234-8) were found at night on vegetation by streams. of two completely metamorphosed young each has a snout-vent length of 15.7 mm. another having a snout-vent length of 16.2 mm. has a tail stub 2 mm. long and a completely metamorphosed mouth. two others have snout-vent lengths of 13.6 and 14.1 mm. and tail lengths of 11.5 and 8.1 mm. respectively; in these the mouth parts are incompletely metamorphosed. the single female available (ummz 121399) contains approximately 60 ovarian eggs, the largest of which are about 2.5 mm. in diameter. _referred specimens.--chiapas_: 6.2 km. s of rayón mescalapa, ku 58063-74, 58199 (1 tadpole), 58234-8; 5.6 km. s of rayón mescalapa, ku 58062, 58200 (5 tadpoles); 11.4 mi. n of pueblo nuevo solistahuacán, ummz 121395-9. the specimens listed last were collected along a stream between pueblo nuevo solistahuacán and rayón mescalapa, which, according to floyd l. downs, is probably the same stream listed above as 5.6 km. s of rayón mescalapa. acknowledgments i take this opportunity to thank dale l. hoyt, thomas e. moore, and john wellman, who ably assisted in collecting and studying these frogs in the field. i am indebted to floyd l. downs for permission to include specimens collected by him and john winklemann. my studies on hylid frogs are supported by the national science foundation (grant g 9827). literature cited duellman, w. e. 1960. synonymy, variation, and distribution of _ptychohyla leonhard-schultzei_ ahl. studies of american hylid frogs. iv. herpetologica, 16:191-197, september 23. stuart, l. c. 1954. descriptions of some new amphibians and reptiles from guatemala. proc. biol. soc. washington, 67:159-178, august 5. taylor, e. h. 1944. a new genus and species of mexican hylid frogs. univ. kansas sci. bull., 30 (pt. 1):41-45, june 12. _transmitted january 23, 1961._ 28-6442 * * * * * transcriber's notes italicized text is shown within _underscores_. italicized bold text is shown within ~_tildes and underscores_~. page 352: rejoined last paragraph, originally split by plate 25. page 355: changed typo "comaprisons" to "comparisons." university of kansas publications museum of natural history volume 17, no. 11, pp. 503-515, 5 figs. march 20, 1968 genera of leptodactylid frogs in méxico by john d. lynch university of kansas lawrence 1968 university of kansas publications, museum of natural history editors: e. raymond hall, chairman, henry s. fitch, frank b. cross volume 17, no. 11, pp. 503-515, 5 figs. published march 20, 1968 university of kansas lawrence, kansas printed by robert r. (bob) sanders, state printer topeka, kansas 1968 31-9418 genera of leptodactylid frogs in méxico by john d. lynch introduction according to the most recent review of the mexican amphibian fauna (smith and taylor, 1948), six genera of leptodactylid frogs occur in méxico. one other genus, _pleurodema_, occurs in lower central america. smith and taylor recognized one species of _engystomops_, 28 of _eleutherodactylus_, three of _leptodactylus_, eight of _microbatrachylus_, 12 of _syrrhophus_, and five of _tomodactylus_. subsequent to the publication of their checklist of the mexican amphibia (1948), numerous taxonomic changes have been proposed. many species of _eleutherodactylus_ have been added to the fauna, either through the extension of their recorded ranges into méxico from guatemala or by the recognition of species unknown in 1948, whereas some nominal species have been synonymized. _microbatrachylus_ has been regarded as synonymous with _eleutherodactylus_ (lynch, 1965); four species of _microbatrachylus_ currently are regarded as valid (duellman, 1961, lynch, 1965). _syrrhophus_ was revised in part by duellman (1958) and firschein (1954), and a species of _tomodactylus_ transferred to _syrrhophus_ by dixon (1957), who redefined _tomodactylus_ and added more species to the genus. since beginning my studies of the mexican leptodactylids in 1962, i have become acutely aware of difficulties involved in defining the genera. a revision of _eleutherodactylus_ and a review of _syrrhophus_ are nearing completion, but prior to their publication it is desirable to redefine the genera of the mexican leptodactylids, and in so doing recognize an heretofore unnamed genus. the definitions of _eleutherodactylus_ and _leptodactylus_ may need to be altered in the future, since both are widespread in south america and occur in the west indies. their definitions as given here are as precise as present knowledge permits. _syrrhophus_ and _tomodactylus_ are small assemblages that occur only in southwestern united states, méxico, and guatemala. taylor (1952) synonymized _engystomops_ with _eupemphix_ which, although related, should be regarded as generically distinct (gallardo, 1965). perhaps the most conservative classification is that of myers (1962) who, without published evidence, combined _eleutherodactylus_, _syrrhophus_, and the south american _lithodytes_ in a single genus. the major problem for students working with the mexican leptodactylids has not been the separation of _engystomops_ or _leptodactylus_ from other genera but the separation and definition of the eleutherodactyline frogs currently placed in three genera, _eleutherodactylus_, _syrrhophus_, and _tomodactylus_. as will be shown in this paper, these are more conveniently placed in four genera. once a fourth genus is recognized, certain phylogenetic problems disappear and a reasonable zoogeographic interpretation is possible for middle american leptodactylid distribution. analysis of characters in méxico and northern central america approximately 55 species of eleutherodactyline frogs (_eleutherodactylus_, _syrrhophus_, and _tomodactylus_) are known. four genera can be recognized on the basis of the nature of inguinal glands, morphology of the hands and feet, and certain osteological features. [illustration: fig. 1. _tomodactylus angustidigitorum_ (ummz 114305, × 4.5) illustrating the lumbo-inguinal gland typical of members of the genus. from a kodachrome by wm. e. duellman.] glands leptodactylids have a variety of glands that have been used as generic characters. smith and taylor (1948) regarded the so-called inguinal gland as a generic character in mexican eleutherodaycty-lines. lynch (1965) showed that _eleutherodactylus_ and _microbatrachylus_ cannot be separated by the nature of the gland or the condition of the prevomers (dentate or not). _syrrhophus_ and _tomodactylus_, as defined by smith and taylor (1948), are not generically distinct because of overlap in the condition of the prevomers and in the development of the gland. firschein (1954) stated that _syrrhophus_ differed from _tomodactylus_ by having an axillary gland, but it is now known that one species of _syrrhophus_ lacks the gland. the inguinal glands of _eleutherodactylus_ and _syrrhophus_, if present, are diffuse, irregular in outline, and generally not prominent; in _tomodactylus_ the gland is higher on the body (a lumbo-inguinal gland), compact, oval in outline, and prominent (fig. 1). axillary glands occur in most _syrrhophus_ but are not known in _tomodactylus_ or _eleutherodactylus_. hands and feet the tips of the digits are laterally expanded in most _eleutherodactylus_, _syrrhophus_, and _tomodactylus_. two species of _eleutherodactylus_ (_augusti_ and _tarahumarensis_) and two _tomodactylus_ (_angustidigitorum_ and _grandis_) lack any expansion of the digital tips. all but two of the species of eleutherodactyline frogs (_e. augusti_ and _e. tarahumarensis_) have a transverse groove across the tips of the digits (fig. 2). [illustration: fig. 2. palmar views of the hands and lateral views of the tip of the third digits of _eleutherodactylus alfredi_ (left, ku 93994, × 5) and _hylactophryne augusti_ (right, ku 102594, × 3).] supernumerary tubercles rarely are present on the feet of _eleutherodactylus_, but are present and numerous in every species of _syrrhophus_, _tomodactylus_, and in the members of the _augusti_ group of _eleutherodactylus_ (fig. 3). the tubercles are small and numerous in _syrrhophus_ and larger in _tomodactylus_ and the _eleutherodactylus augusti_ group. most species of _eleutherodactylus_ have no plantar supernumerary tubercles; a few species have such tubercles, which never extend between the metatarsal tubercles as in _syrrhophus_ and _tomodactylus_. [illustration: fig. 3. plantar views of feet of _eleutherodactylus alfredi_ (left, ku 93994, × 4.5), _syrrhophus pipilans nebulosus_ (middle, ku 58900, × 7.5), and _hylactophryne augusti_ (right, ku 102594, × 3) showing differences in size and arrangement of supernumerary tubercles.] tarsal folds and tubercles are lacking in _syrrhophus_, _tomodactylus_, and the _augusti_ group of _eleutherodactylus_. several species of _eleutherodactylus_ lack tarsal folds and tubercles, but in nearly every species group, one or more species possess either an inner tarsal fold, inner tarsal tubercle(s), or outer tarsal tubercles. the terminal phalanges of _syrrhophus_, _tomodactylus_, and all _eleutherodactylus_ (except the frogs of the _augusti_ group) are distinctly t-shaped. in the latter, the bones are knob-shaped distally (fig. 4). t-shaped terminal phalanges also are present in _lithodytes_ and _trachyphrynus_ but not in other leptodactylid genera. at least one species of _eupsophus_ (_e. quixensis_) has terminal phalanges that resemble those of the _eleutherodactylus augusti_ group. several species of _eleutherodactylus_, _syrrhophus_, and _tomodactylus_ with slender fingers have t-shaped terminal phalanges although the terminal dilations proportionately are only scarcely wider than the finger tips in the _eleutherodactylus augusti_ group. the presence of a terminal groove at the tip of the finger is an external indicator of the t-shaped terminal phalanges. [illustration: fig. 4. terminal phalanges of four leptodactylid frogs (all × 13.5). (a) _eleutherodactylus mexicanus_, ku 55593; (b) _eupsophus roseus_, ku 84731; (c) _eupsophus quixensis_, uimnh 59643; and (d) _hylactophryne augusti_, ku 56192.] skull all mexican eleutherodactyline frogs have quadratojugal-maxillary articulations, completely roofed skulls in adults, median contact of the nasals, separated occipital condyles, and large prevomers. the premaxillae of all species are visible when the skulls are viewed from directly above. the pterygoid lacks a medioventral flange and does not meet the palatine. in no species is the anterior arm of the squamosal in contact with the maxillary. of the numerous species examined (30 _eleutherodactylus_, four _syrrhophus_, and four _tomodactylus_), the species in the _eleutherodactylus augusti_ group are unique in having a sphenethmoid with a blunt anterior edge. pectoral girdle all species have large cartilaginous plates in the pectoral girdles; none possesses a bony style. no divergent modifications of the clavicle and coracoid bones are known in the family. generic accounts genus ~eleutherodactylus~ dumeril and bibron, 1841 _type-species._--_hylodes martinicensis_ tschudi, 1838 _diagnosis and definition._--small to large frogs (12 to 110 mm. snout-vent length) having slightly to widely expanded digital pads, each pad bearing a terminal transverse groove; lumbo-inguinal, inguinal, and axillary glands absent, or if present, diffuse, irregular in outline, not compact; plantar supernumerary tubercles absent, or if present, six or fewer, restricted to distal area of plantar surface, and not extending between metatarsal tubercles; tarsus bearing inner or outer tubercles or folds or not; toes free to one-half webbed; terminal phalanges t-shaped; sternum cartilaginous, lacking bony style; sphenethmoid not truncate anteriorly; nasals in contact medially; maxillary and quadratojugal in contact; anterior arm of squamosal not in contact with maxillary; dermal cranial elements not involved in integumentary-cranial co-ossification; prevomers large, dentigerous processes present or not, dentate or not; maxillary and premaxillary bones dentate; occipital condyles separated; development direct. _composition._--about 420 names have been applied to frogs of this genus; many of these names are synonyms, and many other species remain undescribed and unnamed. perhaps the genus contains 350 species. thirty-one species occur in méxico and northern central america. _distribution._--from tamaulipas and sinaloa, méxico, exclusive of the mexican plateau, to at least peru and southernmost brazil and throughout the west indies. introduced into florida. _etymology._--greek (_eleuthero_ + _dactylus_) meaning free-toed. genus ~engystomops~ jiménez de la espada, 1872 _type species._--_engystomops petersi_ jiménez de la espada, 1872 _diagnosis and definition._--small frogs (20 to 40 mm. snout-vent length) having undilated digital tips lacking transverse grooves; lumbo-inguinal or inguinal glands absent; plantar supernumerary tubercles present, extending between metatarsal tubercles; tarsus bearing spinelike tubercle on inner edge; toes free; terminal phalanges pointed; sternum bearing bony style; spenethmoid not truncate anteriorly; nasals in contact medially; maxillary and quadratojugal in articular contact; anterior arm of squamosal not in contact with maxillary; dermal cranial elements not involved in integumentary-cranial co-ossification; prevomers moderate in size, lacking teeth; maxillary and premaxillary bones edentate; occipital condyles separated; tadpole free living. _composition._--four nominal species (_e. petersi_, _e. pustulatus_, _e. pustulosus_ and _e. schereri_). _distribution._--central veracruz and eastern oaxaca, méxico, to trinidad, bolivia, and peru, east of the andes. _etymology._--greek (_engys_ + _stoma_) meaning narrow-mouthed. genus ~hylactophryne~ new genus _type-species._--_hylodes augusti_ dugés, 1879 _diagnosis and definition._--medium to large frogs (37 to 94 mm. snout-vent length) having undilated digital tips lacking terminal grooves; lumbo-inguinal or inguinal glands absent; plantar supernumerary tubercles present, prominent, extending to but not between metatarsal tubercles; tarsus lacking tubercles or folds; toes free of webbing; terminal phalanges knob-shaped, lacking elongate lateral expansions; sternum cartilaginous, lacking bony style; sphenethmoid truncate anteriorly; nasals in contact medially; maxillary and quadratojugal in articular contact; anterior arm of squamosal not in contact with maxillary; dermal cranial elements not involved in integumentary-cranial co-ossification; prevomers large, bearing dentigerous processes; maxillary and premaxillary bones dentate; occipital condyles separated; development direct. _composition._--two species, _h. augusti_ and _h. tarahumarensis_, the former composed of four subspecies (zweifel, 1956). _distribution._--from arizona, new mexico, and texas to guerrero and puebla, méxico, and a relict population on cerro quingola (just west of the isthmus of tehuantepec, méxico). _etymology._--greek (_hylactor_ + _phryne_) meaning barking toad; in reference to the voice and common name. genus ~leptodactylus~ fitzinger, 1826 _type-species._--_leptodactylus typhonia_ fitzinger, 1826 _diagnosis and definition._--small to large frogs (30 to about 200 m., snout-vent length) having undilated to slightly expanded digital tips bearing pads, no transverse groove at tips of digits; lumbo-inguinal, axillary, and/or ventral glands present or not, low, diffuse; plantar supernumerary tubercles generally absent, if present not extending between metatarsal tubercles; tarsus bearing tarsal folds or not; toes free of webbing, extensive lateral fringes present in some species; terminal phalanges pointed, not t-shaped; sternum bearing bony style; sphenethmoid not truncate anteriorly; nasals in contact medially; maxillary and quadratojugal in articular contact; anterior arm of squamosal not in contact with maxillary; dermal cranial elements not involved in integumentary-cranial co-ossification; prevomers large, bearing dentigerous processes; maxillary and premaxillary bones dentate; occipital condyles separated; tadpole free living. _composition._--sixty species according to smith and taylor (1948); 54 according to gorham (1963); argentinian authors have described several more in recent years. _distribution._--southern sonora, méxico, and southern texas throughout the central and south american lowlands to argentina. also known from hispaniola and puerto rico in the greater antilles and a few islands in the lesser antilles. _entymology._--greek (_leptos_ + _dactylus_) meaning slender toes. genus ~syrrhophus~ cope, 1878 _type-species._--_syrrhophus marnockii_ cope, 1878 _diagnosis and definition._--small to medium sized frogs (18 to 40 mm. snout-vent) having slight to prominent digital expansions with transverse groove at tip of each digit; lumbo-inguinal and inguinal gland flattened, irregular in outline, not compact and oval; axillary glands present or not; plantar supernumerary tubercles numerous, more than eight, usually extending between metatarsal tubercles; tarsus lacking tubercles or folds; toes free or basally webbed; terminal phalanges t-shaped; sternum cartilaginous, lacking bony style; sphenethmoid not truncate anteriorly; nasals in contact medially; maxillary and quadratojugal in articular contact; anterior arm of squamosal not in contact with maxillary; dermal cranial elements not involved in integumentary-cranial co-ossification; prevomers large, usually lacking dentigerous processes and teeth; maxillary and premaxillary bones dentate; occipital condyles separated; development direct. _composition._--thirteen species; the species described as, or later referred to, _syrrhophus_ from lower central america and south america are _eleutherodactylus_ or _eupsophus_. _distribution._--low to moderate elevations from sinaloa, méxico, to guatemala on the pacific versant; from the edwards and stockton plateaus of texas to british honduras on the caribbean versant. _etymology._--greek, emendation of _syrrhaptos_, meaning sewn together in reference to the united outer metatarsals. genus ~tomodactylus~ günther, 1900 _type-species._--_tomodactylus amulae_ günther, 1900. _diagnosis and definition._--small frogs (20 to 35 mm. snout-vent length) having digital expansions or not, with transverse groove across tip of each digit; lumbo-inguinal gland prominently elevated, compact, oval, often patterned; axillary glands absent; plantar supernumerary tubercles numerous, more than eight, usually extending between metatarsal tubercles; tarsus lacking tubercles or folds; toes free; terminal phalanges t-shaped; sternum cartilaginous, lacking bony style; sphenethmoid not truncate anteriorly; nasals in contact medially; maxillary and quadratojugal in articular contact; anterior arm of squamosal not in contact with maxillary; dermal cranial elements not involved in integumentary-cranial co-ossification; prevomers large, usually bearing dentigerous processes; maxillary and premaxillary bones dentate; occipital condyles separated; development direct. _composition._--ten species. _distribution._--the southern edge of the mexican plateau from sinaloa to veracruuz and onto the oaxaca highlands and sierra madre del sur. _etymology._--greek (_tomis_ + _dactylus_) meaning knife toe; in reference to either the sharp subarticular tubercles or the unwebbed toes. discussion the preceding definitions only slightly alter the present generic limits of mexican leptodactylids. two species, previously regarded as _eleutherodactylus_, are transferred to the new genus _hylactophryne_. the arrangement of the species of _syrrhophus_ and _tomodactylus_ remains the same as concluded by dixon (1957), duellman (1958), and firschein (1954) in their reviews of the genera. lumbo-inguinal glands are most prominent in the genera _pleurodema_ and _tomodactylus_. various nondescript glands are present in many genera, but none is so well developed as those of _pleurodema_ and _tomodactylus_. at least nine leptodactylid genera are either known or thought to be terrestrial breeders lacking a free-living tadpole stage (_eleutherodactylus_, _euparkerella_, _hylactophryne_, _niceforonia_, _noblella_, _sminthillus_, _syrrhophus_, _tomodactylus_ and _trachyphrynus_). _niceforonia_ and _trachyphrynus_, and probably _hylactophryne_, are not closely related to the other genera. direct development probably is an adaptation to adverse environmental conditions since many of the species occur in semi-arid or cold (andean páramos) areas. _eleutherodactylus_ is generally thought to be the stock from which _euparkerella_, _noblella_, and _sminthillus_ evolved (griffiths, 1959) and from which _syrrhophus_ and _tomodactylus_ are derived (firschein, 1954). the present distribution of _hylactophryne_ (isolated on the mexican plateau) and its digital form (like that of papuan and many primitive south american leptodactylids) suggest that the genus was isolated in méxico throughout the tertiary, whereas the other central american genera are either post-pliocene derivatives of _eleutherodactylus_ or invaders of central america from south america since the mid-pliocene land bridge was formed (lloyd, 1963). piatt (1934) presented arguments against assigning _eleutherodactylus latrans_ to the genus _lithodytes_ and concluded that it was a "true" _eleutherodactylus_. contrary to his arguments, _latrans_ (= _augusti_ of zweifel) and _e. tarahumarensis_ taylor differ from all other _eleutherodactylus_ (and _syrrhophus_ and _tomodactylus_) in the nature of the tips of the digits (external and skeletal). the digits of _hylactophryne_ are like those of _eupsophus_. my study of nearly all genera of leptodactylids indicates that noble (1925) was correct in suggesting that _borborocoetes_ (= _eupsophus_) is a close relative of _eleutherodactylus latrans_, although noble's arguments were based in part upon false evidence concerning the breeding habits of _e. latrans_, then thought to have a free-living tadpole. kellogg (1932) and piatt (1934) argued that the terminal phalanges of _e. latrans_ were typically eleutherodactyline. the variation of this character in _eupsophus_ (see fig. 4) ranges from knobbed to bifurcate or y-shaped (t-shaped in _eleutherodactylus_, _syrrhophus_ and _tomodactylus_) and encompasses the nature of the character represented in _hylactophryne_. _eupsophus_ differs from _hylactophryne_ in possessing a frontoparietal fontanelle, in generally having a maxillary-quadratojugal gap, and in having a free swimming tadpole stage. [illustration: fig. 5. outline drawings of _leptodactylus melanonotus_ (left, ku 65704, × 2) and _eleutherodactylus alfredi_ (right, ku 93994, × 2).] key to mexican leptodactylid genera 1. small (20-40 mm.), pustular, toadlike frogs; maxillary and premaxillary bones not bearing teeth _engystomops_ large (20-110 mm.), smooth skinned and non-toadlike frogs; maxillary and premaxillary bones bearing teeth 2 2. no conspicuous waist (fig. 5); sternum bearing bony style, _leptodactylus_ constrictions at waist (fig. 5); sternum cartilaginous, no bony style 3 3. few (less than six), if any, supernumerary tubercles on plantar surface _eleutherodactylus_ many (more than 8) supernumerary tubercles on plantar surfaces 4 4. terminal, transverse groove across tip of digits, especially outer two fingers, digits expanded or not; small frogs (18 to 40 mm.) 5 tips of digits lacking transverse groove; digits unexpanded; medium-sized to large frogs (37 to 94 mm.) _hylactophryne_ 5. lumbo-inguinal gland compact, oval _tomodactylus_ lumbo-inguinal or inguinal gland absent or diffuse and irregular in outline _syrrhophus_ literature cited dixon, j. r. 1957. geographic variation and distribution of the genus _tomodactylus_ in mexico. texas jour. sci., 9:379-409, december. duellman, w. e. 1958. a review of the frogs of the genus syrrhophus in western mexico. occas. papers mus. zool. univ. michigan, 594:1-15, june 6. 1961. the amphibians and reptiles of michoacan, mexico. univ. kansas publ. mus. nat. hist., 15:1-148, december 20. firschein, i. l. 1954. definition of some little understood members of the leptodactylid genus _syrrhophus_, with a description of a new species. copeia, 1:48-58, february 19. gallardo, j. m. 1965. a proposito de los leptodactylidae (amphibia anura). papeis avulsos, 17:77-87, january 30. gorham, s. w. 1963. the comparative number of species of amphibians in canada and other countries. iii. summary of species of anurans. canadian field-nat., 77:13-48, march. griffiths, i. 1959. the phylogeny of sminthillus limbatus and the status of the brachycephalidae (amphibia salientia). proc. zool. soc. london, 132:457-87, may. kellogg, r. 1932. mexican tailless amphibians in the united states national museum. bull. u. s. natl. mus., 160:224 pp., march 31. lloyd, j. j. 1963. tectonic history of the south central-american orogen, _in_ childs and beebe eds., backbone of the americas. amer. assoc. petroleum geol., pp. 88-100. lynch, j. d. 1965. a review of the eleutherodactylid frog genus _microbatrachylus_ (leptodactylidae). nat. hist. misc., 182:1-12, december 15. myers, g. s. 1962. the american leptodactylid frog genus _eleutherodactylus_, _hylodes_ (= _elosia_), and _caudiverbera_ (= _calytocephalus_). copeia, 1:195-202, april 11. noble, g. k. 1925. an outline of the relation of the ontogeny to phylogeny within the amphibia. i. amer. mus. nov., 165:1-17, april 16. piatt, j. 1934. the systematic status of eleutherodactylus latrans (cope). amer. midl. nat., 15:89-91, february 15. smith, h. m. and taylor, e. h. 1948. an annotated checklist and key to the amphibia of mexico. bull. u. s. natl. mus., 194:1-118. june 7. taylor, e. h. 1952. a review of the frogs and toads of costa rica. univ. kansas sci. bull., 35:577-942, july 1. zweifel, r. g. 1956. a survey of the frogs of the _augusti_ group, genus _eleutherodactylus_. amer. mus. novitates, 1813:1-35, december 23. _transmitted july 11, 1967._ 31-9418 * * * * * transcriber's notes bold text is shown within ~tildes~. italicized text is shown within _underscores_. illustrations have been moved to avoid breaking up paragraphs of text. page 507: changed know to known (are not know in tomodactylus). page 507: added closing parenthesis in fig. 2 caption after × 3. page 509: changed compeltely to completely (compeltely roofed skulls). page 512: veracruuz may be a typo for veracruz (sinaloa to veracruuz). page 514: changed two occurrences of hylatophryne to hylactophryne. experiments on _the nervous system_, with opium and metalline substances; made chiefly with the view of determining the _nature and effects_ of animal electricity. by alexander monro, m. d. professor of medicine, anatomy and surgery in the university of edinburgh; fellow of the royal college of physicians, and of the royal society of edinburgh, and of the royal academy of surgery in paris. edinburgh: printed by adam neill and company, for bell & bradfute, and t. duncan; and j. johnson, london. m.dcc.xciii. contents. _page_ introduction, 5 observations on the circulating and nervous systems of frogs, 6 experiments with opium, 9 corollaries from the above facts and experiments, 12 summary of experiments made on animals with metalline substances, 17 summary of facts proved by the foregoing experiments, 35 resemblance of the fluid put in motion by the foregoing experiments to the electrical fluid, 38 the nervous fluid or energy not the same with the electrical, nor with the fluid put in motion by the foregoing experiments, 40 general conclusions, 42 introduction. when, in november last, i began to make experiments on animal electricity, of which i read some account to the royal society on the 3d of december; i was not only much hurried with business, but could not procure a sufficient number of frogs for the purpose. during the last winter and spring, i prosecuted the subject more fully and with greater attention; and, on the third day of june, i read a second paper to the royal society, to which i have, since that time, made additions. i shall now state a summary of the chief circumstances i have observed, with a few remarks. observations on the circulating and nervous systems of frogs. as my experiments with opium, as well as those on animal electricity, have been performed on frogs chiefly; i shall premise some observations on their circulating and nervous systems. their heart consists of one auricle and one ventricle only, their aorta supplying their air vesicles or lungs, as well as all their other organs; and, of course, their venæ cavæ return the blood from all parts to the heart. the ventricle of their heart contracts about sixty times in a minute; and the purple colour of the blood which is seen within it, disappears after each contraction, or the blood is entirely expelled by its contraction. for upwards of an hour after cutting out its heart, a frog can crawl or jump; and, for upwards of half an hour longer, it contracts its legs when the toes are hurt, though not with sufficient force to more its body from the place where it is laid. their encephalon consists of brain and cerebellum, each of which, on its upper part, is divided into two hemispheres; and, below, they are conjoined by thick crura, which form the medulla oblongata and spinal marrow, both of which are proportionally larger than in man, and more evidently consist of two cords. there are nine true vertebræ; and at the sixth of these, the spinal marrow terminates in the cauda equina. the sciatic nerves are formed by three pairs of nerves, sent out below the seventh, eighth and ninth vertebræ, and by one pair from the os sacrum. a nerve, resembling our great sympathetic nerve, passes downwards from the abdomen into the pelvis. two days after cutting off the head of a frog at its joining with the first vertebra, i found it sitting with its legs drawn up, in their usual posture; and when its toes were hurt, it jumped with very considerable force. its heart likewise continued to beat about forty times in a minute, and so strongly as to empty itself and circulate the blood. in several frogs, after cutting off the back part of the six undermost true vertebræ, i took out all that part of the spinal marrow with the cauda equina which they cover. the lower extremities were rendered insensible to common injuries, and lay motionless, yet the frogs lived several months thereafter, and the wounded parts of their backs cicatrised; and the bones of their legs, which i fractured, were re-united, the blood circulating freely in their vessels. it is universally known, that if, after amputating the limb of a warm blooded animal, we repeatedly irritate the nerves which terminate in muscles, repeated convulsions of the muscles are for some time produced; and that in frogs, and other cold blooded animals, the nerves retain this power still longer. but it has been commonly supposed, that, after irritating the nerve a given number of times, the effect ceases, authors conceiving that there is lodged in the nerve some fluid, or other energy which is exhausted by repeated explosions. instead of this, i have found that the time the nerves preserve their power is the same, whether we irritate them or not; or that their energy is not exhausted by irritation, unless the irritation be such as sensibly alters their texture. experiments with opium. i cut one hole in the fore and upper part of the cranium and dura mater of a frog, and another in the back part of the lowermost vertebræ, and then injected, from the one hole to the other, a small syringe full of water, in five ounces of which one ounce of opium had been infused for three days. the infusion, by this means brought into contact with the whole surface of the encephalon and spinal marrow, produced almost instantly universal convulsions; and, in less than two minutes thereafter, the animal was incapable of moving its body from the place where it was laid. a quarter of an hour thereafter, i found the heart beating twenty-five times only in the minute; and so feebly, that it could not entirely expel the blood. when, half an hour thereafter, the sciatic nerves were pinched, a light tremor only was excited in the muscles of the leg; and animal electricity produced but feeble twitchings of the muscles. the infusion of opium, injected in the same manner in rabbits and in a pig, produced similar effects. i had long ago[1] observed, that an infusion of opium, poured into the cavity of the abdomen of a frog, after cutting out its heart, occasioned, in a few minutes, convulsions of its hind legs. i have since found, that, after cutting off the head, and cutting out the heart of a frog, its hind legs are considerably weakened by pouring an infusion of opium into the cavity of its abdomen. although an infusion of opium poured into the auricle and ventricle of the heart of a frog, instantly renders that organ incapable of contraction, and, even after the aorta has been previously cut, occasions convulsions of the legs, yet i have not found that by opium applied to the brain, the spinal marrow, the heart, or abdominal viscera, the muscles of the legs were so entirely killed as not to perform some motion when their nerves were pinched, or when they were acted on by animal electricity. after taking out the lower half of the spinal marrow, and likewise cutting transversely all the parts at the pelvis, except the crural arteries and veins and lymphatics, which probably accompany them, i found that an infusion of opium, applied to the skin and muscles of the legs, affected the superior parts of the body[2]: more probably, in my opinion, by absorption, than through any minute remanent branches of the nerves, especially as i do not find, on laying the vessels so prepared over a gold probe, and touching with it zinc laid under the spine, that convulsions of the legs can be excited. at the same time, the quantity of opium absorbed is so small, that i could not distinguish its smell or taste in the blood; nor did i find these distinguishable, in other experiments, in which the frogs were violently convulsed after applying the infusion to the surface of their skin. animal electricity or different metals applied to the head of a frog, or to any part of its spine above its sixth vertebra, do not occasion convulsions of its hind legs. corollaries from the above facts and experiments. from the above facts and experiments, it appears, 1. that the frog, after its head is cut off, feels pain, and, in consequence of feeling, moves its body and limbs. 2. as the nerves of the hind legs are not affected by animal electricity, unless it be applied lower than the fifth vertebra, these nerves do not seem to be derived solely or chiefly from the brain or cerebellum. 3. as opium, after the circulation ceases, affects organs distant from those to which it is applied, it is beyond doubt, that the latter suffer in consequence of sympathy of nerves. 4. it appears that, in this animal, there is sympathy of nerves after the head is cut off; or that sympathy of nerves does not, in this animal, depend entirely on the connection of nerves within the head. 5. as, after cutting off the head, this animal is susceptible of pain, and, in consequence of that, performs voluntary motion, it appears that, in it, the brain is not the sole seat of the _sensorium commune_. 6. several weeks after i had taken out the lowermost half of the spinal marrow, and with it the cauda equina, i daily applied, for four days running, animal electricity to the sciatic nerves, by passing a gold probe between them and the os sacrum, and excited several hundreds of convulsions of the thighs and legs, and yet found that, on laying bare the femoral nerves, and pinching them, the muscles were slightly convulsed. hence, i apprehend, additional force is given to an opinion i ventured many years ago to propose[3], that the nerves do not receive their energy wholly from the head and spinal marrow, but that the texture of every branch of a nerve is such as to furnish it, or that the structure of each nerve is similar to that of the brain. 7. from the above experiments, it appears probable, in the highest degree, that opium may be absorbed in such quantity as to produce fatal symptoms. 8. the following circumstances concur in rendering inadmissible an opinion lately proposed by m. fontana, that poisons operate by changes they produce on the mass of blood, or on some unknown principle connected with the blood. a. if his opinion was just, poison introduced into a vein of the extremities, so as to be in contact with this unknown principle, should operate as quickly, and in the same manner as when the poison is mixed with the blood near the heart, which he admits is not the case[4]. b. cutting the spinal marrow in frogs, before applying the poison of the viper to their legs, prevents it from killing them[5]; which should not happen, if the poison acted on the blood alone. c. he acknowledges that an animal bit in its leg by a viper, instantaneously feels acute pain[6]; and it, in like manner, feels instantly great uneasiness when the poison is mixed with its blood[7]. we know for certain, that, through the medium of the nerves, we are instantly rendered sensible of injury done to the most distant parts of our bodies. are we not, therefore, in the last mentioned experiment, to conclude, that the uneasiness was produced because the poison acted upon the nerves of the vessels? d. in like manner, animals were convulsed as soon as they were wounded, or received the poison into a blood-vessel; and long before the blood could have reached the muscles in action[8]. e. as soon as the distilled water of lauro-cerasus was poured into the stomach of a pigeon, it was convulsed, died instantly[9], that is, before the poison could have entered the mass of blood. f. many years ago, i found, after cutting the venæ cavæ and aorta of a frog, that a watery solution of opium poured into the heart, occasioned, in a few minutes, convulsions in its legs; and, after cutting out the heart, that the opium poured into the cavity of the abdomen affected the legs in like manner; although, in these experiments, the circulation was not only interrupted, but the greater part of the blood evacuated. i therefore then concluded[10], and now conclude, that opium and other poisons, even after they are mixed with the mass of blood, produce their fatal effects, chiefly and almost solely, by acting on the nerves of the heart and vascular system, and, through these, affecting the whole of the nervous system. summary of experiments made on animals with metalline substances. i shall now proceed to state the several circumstances i have observed, in my experiments, which more directly lead us to judge of the nature and cause of animal electricity. 1. when two plates of different metalline substances, particularly of zinc and gold, between which a living frog is placed, are brought into contact with each other, those muscles, which are farther from the brain and spinal marrow than the metals, are convulsed: and this effect follows, although the animal and metals are placed on an inverted glass jar, and that a stick of sealing wax is interposed between the hand of the operator and the metals; that is, although the animal, with the metals, be insulated. i have further observed, that the metals, disposed as above described, excite convulsions in the legs, after all the parts of the frog have been divided transversely at the pelvis, providing only that they are, thereafter, laid in contact with each other. 2. when all the parts of a living frog, except the large nerves called sciatic, are cut transversely at the pelvis, and the fore part of the animal is laid on a plate of zinc, supported by glass, and the hind legs on glass; if a gold probe be applied so as to touch the zinc and one of the legs; or a piece of metal put under one of the legs; the muscles of both legs will be convulsed. the event is the same, after the body of the frog has been cut transversely about the middle of the spine: or when the legs are laid on the zinc and the spine on glass. if a piece of perforated dry paper is placed between the gold probe and the muscles, there will be no convulsions; but wet paper interposed does not prevent the convulsions. on separating the gold probe from the muscles there are no convulsions. 3. if, after the animal and metals are placed as above described, the joining of the two legs at the ossa pubis is cut, that leg only will be convulsed with which the gold is in contact. 4. the spine of the frog with the zinc being placed on one glass, and the legs on another glass, if the gold, supported by one hand, which we shall call the right hand, be applied to the zinc alone, and not to the legs, these are not convulsed. but if the operator applies his left hand to the legs, or if a bystander, communicating with the operator by the medium of the floor only, touches them, they are convulsed. if a stick of sealing-wax be interposed between his right hand and the gold, or between his left hand and the legs; or, if the bystander, touching the legs, is insulated, by standing on a stool supported by glass feet, the legs will not be convulsed. if the insulated bystander touches the legs with one hand, and the operator with his other hand, the legs are immediately convulsed. 5. after cutting the spine transversely under the fifth vertebra, and all the parts of the pelvis, except the sciatic nerves, and laying the spine on zinc supported by glass, and the legs on glass; if gold be applied to the zinc, and then to one of the sciatic nerves, both legs, if they have not been separated from each other at the ossa pubis, will be convulsed[11]. and this happens although a stick of sealing-wax be interposed between the hand of the operator and the gold probe, and although no metalline substance touches the legs. this experiment succeeds after denuding the sciatic nerves for the length of an inch, and wiping them dry; and it continues to succeed for an hour or more, and till the nerves are evidently discoloured and shrunk in their size. and, after that, although we wet the nerves, their powers are not restored; shewing that the influence had been conveyed not by wetness on the surface of the nerves, but by the particular matter of which nerves are composed. the event is the same, when the upper ends of the sciatic nerves are cut away from the spine, and laid on the zinc. 6. after preparing the frog and placing the metals as in last experiment, if a piece of thin dry paper, pierced with a number of small holes, be interposed between the gold probe and the sciatic nerves, the legs will not be convulsed. but, if the paper be wetted, although it is not perforated, the legs will be convulsed. after preparing a frog, as in last experiment, and laying the spine on one glass, and the legs on another, if the zinc be laid on a third glass, and the gold probe applied to it and to the sciatic nerves, the legs will not be convulsed. 8. if the spine and hind legs, connected by the sciatic nerves, are all laid on the same plate of zinc, supported by glass, the legs are not convulsed on touching the zinc with the gold probe held in the right hand, although the left hand is applied to the legs. 9. if several frogs, prepared as above described, are laid upon glass, in a straight line touching each other, and that the first frog is supported on zinc, and the last upon gold; if one end of a brass wire is applied to the zinc, and the other end of it to the gold; the muscles of all the frogs will be convulsed. the event is the same, although a stick of sealing-wax be interposed between the hand of the operator and the brass wire: that is, although the frog with the metals be insulated. 10. when frogs are prepared as in last experiment, and the spine of the first of them laid on zinc, and the last supported by the left hand of the operator, if with a gold probe, held in his right hand, he touches the zinc, the muscles of all the frogs will be convulsed. but if the hind legs, as well as the spine, of the first frog be laid on the zinc, the muscles of that frog will not be convulsed. 11. after a frog was prepared as before described, i cut the sciatic nerves where they are about to enter the thighs, and laid their cut ends in contact with the muscles, and then touched the zinc and nerves with a gold probe, without exciting convulsions in the thighs or legs. 12. after cutting the sciatic nerves, i tied together their divided parts, and then touched the zinc and nerves above the ligature, with the gold, without finding that the legs were convulsed, when the zinc supporting the spine was laid on one glass and the legs on another: but when the metals and parts of the frog were laid on a wet table, the muscles of the leg were convulsed. 13. when the sciatic nerves have been cut and rejoined by ligature, if while the gold is, with one hand, applied to the zinc and nerves, above the ligature, the other hand touches the feet, the legs are convulsed. 14. if the two hind legs of a frog are separated from each other, and their sciatic nerves afterwards tied to each other; if one of the legs be laid on zinc supported by glass, and the other leg on glass, when, with one hand, the toes of one of the legs are touched, whilst with the other hand a gold probe is applied to the zinc and nerve of the leg which it supports, this leg only will be convulsed. but if the gold probe touching the zinc be applied to the nerve of the most distant leg, both legs will be convulsed. 15. i found it was not necessary, in order to excite convulsions, that either of the metals should be in contact with the living nerve or living flesh of the frog; for if, after separating from each other the hind legs of a frog, and cutting transversely the upper part of their sciatic nerves, i laid a piece of putrid or boiled beef between their sciatic nerves, and two other pieces of putrid or boiled beef between their toes and a plate of zinc; if, with the point of a gold probe, the side of which was applied to the piece of beef placed between the sciatic nerves, i touched the zinc, both legs were convulsed. 16. in like manner, when i placed alternately, in a straight line, a number of dead and living frogs touching each other, and in the living frogs cut, at their pelvis, all the parts but the sciatic nerves; if, with my left hand i touched a dead frog at one end of the line, and with a gold probe, held in my right hand, i touched a plate of zinc, on which a dead frog was laid at the other end of the line or chain of frogs, the muscles of all the living frogs were convulsed. 17. when a chain of living and dead frogs was formed, as in the two last experiments, but without cutting at their pelvis all the parts but the nerves; on applying the gold to the zinc, convulsions of the muscles were not excited. 18. it has been found, that, if a plate of zinc is applied to the upper part of the point of the tongue, and a plate of silver to its under part, on bringing the two metals into contact with each other, a pungent disagreeable feeling, which it is difficult to describe, is produced in the point of the tongue. and if a plate of zinc is placed between the upper lip and the gums, and a plate of gold applied to the upper or under part of the tongue, on bringing these two metals into contact with each other, the person imagines that he sees a flash of lightning, which, however, a bystander in a dark room does not perceive; and the person performing the experiment perceives the flash, though he is hoodwinked. it has been alleged, that the flash happens before the two metals touch each other, and is repeated on separating them; but these facts appear to me very doubtful, as i do not find that a flash is produced when a piece of cambric-paper, in which a number of holes is pierced with a pin, is interposed between the zinc and silver, although the paper does not in thickness exceed 1/1500 part of an inch. after performing this experiment repeatedly, i constantly felt a pain in my upper jaw at the place to which the zinc had been applied, which continued for an hour or more: and in one experiment after i had applied a blunt probe of zinc to the septum narium, and repeatedly touched with it a crown piece of silver applied to the tongue, and thereby produced the appearance of a flash, several drops of blood fell from that nostril; and dr fowler, after making such an experiment on his ears, observed a similar effect[12]. i have farther observed, that although the previous application of a second plate of silver to one half of the plate of zinc, does not prevent the flash when the other half of the plate of zinc, touching the tongue, is brought into contact with the first piece of silver placed between the lip and the gum; yet if the zinc and silver are in the first place applied to each other, then placed between the lip and gum, and, after this, touched with the tongue, there is no appearance of a flash, although some degree of pungency and a disagreeable sensation is perceived by the tongue: and a mixed mass, composed of one part of zinc and two parts of quicksilver, or a mass composed of three parts of zinc and one of silver, incorporated in a furnace, have not the effect, when they are applied to nerves, of exciting convulsions of the muscles in which the nerves terminate. i have also found, that two thick pieces of raw or boiled flesh, one between the zinc and tongue, and the other between the silver and tongue, do not prevent the disagreeable pungent sensation when the two metals touch: and, in like manner, that the interpolation of two pieces of flesh between the zinc and tongue, and between the silver and the upper lip, does not prevent the appearance of a flash, on bringing the two metals into contact. 19. i put a very thick plate of zinc into a vessel with water, and placed, near to it, in the water, the under part of the spine and the hind legs of a frog, after cutting all the parts at the pelvis except the sciatic nerves. i then touched the zinc with a gold probe, and found, that, when i touched that part of the zinc which was above the water, the legs of the frog were not affected; but when i touched that part of the zinc which was below the surface of the water, the legs of the frog were convulsed[13]. i next put into the water one of the hind legs of a dead frog, and its other leg into an adjoining vessel with water. into the opposite side of the second vessel, i put one of the hind legs of a living frog, in which all the parts at the pelvis, except the sciatic nerves, were cut; and into a third glass vessel with water, i put its other leg. when i now touched that part of the zinc, which was below the surface of the water with a gold probe, the legs were not convulsed; but, if i, at the same time, dipped the finger of my other hand into the water contained in the third vessel, they were convulsed: when, instead of my finger, i dipped into the water a stick of sealing-wax, held in my other hand, the legs were not convulsed. i found, by the three following experiments, that the muscles are convulsed, whether the influence, produced by the application of the metals, passes upwards or downwards along the nerves. 20. i cut four living frogs transversely at the middle part of their spines, and threw away the fore parts of their bodies and their abdominal viscera. i next cut, at their _pelves_ all the parts but the sciatic nerves; and at their knees, i cut all the parts but the crural nerves; and, in all of them, i cut asunder the joining of the two hind legs at their ossa pubis. i then laid the legs of all of them in a straight line, supported on different glass vessels inverted, in such a manner that the foot of one frog touched the foot of the next to it. having then placed a plate of zinc under the foot of the first frog, and holding in my left hand the foot of the fourth or last frog, i touched the zinc with a gold probe which i held in my right hand; and found that all the muscles of the loins, thighs and legs of the four frogs were convulsed. 21. when i placed the two frogs in the middle, with their spines contiguous to each other, and the feet of both touching the spines of the other two frogs forming the extremities of the chain, and of course the feet of one of these resting on the zinc, and the feet of the other supported by my left hand: on touching the zinc with the gold probe held in my right hand, all the muscles of the frogs were, as before, convulsed. 22. when i now turned aside the right legs of all the frogs, so that they did not form a chain by touching the next frogs; the right legs were not convulsed. it is evident, that in whatever direction we suppose the influence to have passed in its circle, it must, in experiment 20th, have passed up one leg and down the other in the same frog: and, in experiment 21st, if it passed from one end of the chain to the other end of it, it must have passed upwards in two of the frogs, and downwards in the other two; or if the influence passed from the two ends of the chain towards its middle, where the spines of the two middlemost frogs were contiguous, it must have passed upwards in all of them. 23. when after cutting four living frogs transversely at the middle of their spines, but without cutting at their pelves all the parts but the sciatic nerves, i placed the hind parts of them in a chain, as in experiments 20th, 21st and 22d, the muscles were not convulsed on applying the gold to the zinc. i next found, that after placing in contact with each other the several muscles which had been cut transversely in experiments 20th, 21st and 22d, allowing the nerves to remain undivided, the muscles were not convulsed when i touched the plate of zinc with the gold probe held in my right hand, although i touched the other end of the chain of frogs with my left hand. the reason why the muscles were convulsed in experiments 20th, 21st and 22d, and not in experiment 23d, evidently is, that in the former, the influence was concentrated in the nerve, in the latter the influence was diffused; that is, was in part conveyed by other organs, as well as by the trunks of the nerves. 24. after finding that i could readily excite convulsions in the hind legs of a frog, without cutting it, by laying its back on a plate of zinc, and introducing a gold probe within its intestinum rectum and touching the zinc with the side of the probe, i produced two or three hundred convulsions, succeeding each other quickly, and observed that its legs were, by these means, so much weakened, that it could not jump, and crawled with difficulty, but in a few minutes it recovered nearly the full force of its muscles. in other frogs i passed a gold wire between their sciatic nerves and os sacrum, and twisted together the two ends of the wire over the backs of the animals. i then put them into a zinc vessel filled with water, or into a glass vessel filled with water, in the bottom of which i laid a large plate of zinc: so that every time the animals by moving separated the gold from the zinc, and again brought them into contact, their hind legs were convulsed. i allowed them to remain three or four days in this situation, and found that their limbs were weakened considerably, but not exhausted of their power of motion; and, after removing the gold wire, the limbs by degrees recovered their strength. i made the same experiment on those frogs in which i had, six weeks before, cut out, from behind, all that part of their spinal marrow which is covered by the six undermost vertebræ, and found, several days after the frogs had been subjected to the experiment, that, by pinching their sciatic and femoral nerves, and still more readily by the application of the gold and zinc, weak convulsions of the muscles were excited. 25. after frogs were prepared as above described, by cutting their spines transversely, and then all the parts of their pelves, except their sciatic nerves, i found that slight electrical shocks, or a leyden phial discharged directly through the limbs of a frog, or indirectly by the medium of water, produced convulsions in their muscles, exactly resembling those excited by the metals. and when, after moderate electrical shocks had been passed repeatedly through their legs, the metals were applied to their nerves, in the manner before mentioned, the muscles were convulsed. i found, likewise, that after cutting the nerves transversely, and tying them together, electrical shocks were conducted by the nerves, and occasioned convulsions of the muscles. when i had killed frogs, by discharging through them, from their foreheads to their hind feet, large leyden phials highly charged, i found their nerves or muscles, or both, so much deranged, that feeble convulsions only could be excited by pinching the nerves, or by applying the metals to them. summary of facts proved by the foregoing experiments. on reviewing the foregoing experiments, we shall find the following facts fully proved. 1. on forming a circle by means of the parts of a living animal and of two different metallic bodies, especially gold and zinc, in contact with each other, if a nerve makes part of the circle, the muscles in which the nerve terminates are convulsed. 2. although the nerve making part of such a circle has been cut transversely, yet, if the divided parts of the nerve are laid in contact with each other, or tied together, the muscles, in which it naturally terminates, are convulsed. 3. if the metals, composing parts of the circle, are kept steadily in contact with each other, the convulsions of the muscles cease. but, if they are separated from each other and again rejoined, the convulsions are repeated. 4. the effects are the same, although the dead parts of an animal or pure water make parts of the circle. 5. although the dead parts of an animal, making part of such a circle, are in contact with the metals, the effects are the same. 6. a muscle making part of such a circle may be convulsed whilst the matter put in motion is passing in the direction from the muscle to the nerve. 7. the muscle may be convulsed although it makes no part of the circle in which the matter put in motion passes, as appears from comparing experiment 5th with experiments 13th and 14th. from experiment 13th, it appears, that the fluid put in motion by the metals passes readily along a nerve, after it has been cut, providing the divided parts of it are brought into contact with each other. yet in experiment 14th, in which the left hand of the operator was not applied to the foot of the frog, the muscles in which the nerve, lower than the ligature, terminated, were not convulsed, because the fluid put in motion did not descend lower than the place at which the gold probe touched the nerve above the ligature. we may therefore presume that when a nerve which has not been cut, as in experiment 5th, is touched with the gold probe, the fluid put in motion does not pass lower in the nerve than the place of the probe. hence we perceive the error of those who suppose that the moisture on the surface of the nerve conduces the fluid put in motion to the muscles, and that their action is in consequence of the direct operation of this fluid upon their fibres. 8. the effects are the same when the animal and the metals are insulated, by being placed on glass, whilst sealing-wax is interposed between the hand of the operator and the metals. 9. if any part of the circle is composed of sealing-wax or glass, the muscles are not convulsed. 10. convulsions are not excited unless the metals are in contact with each other; and unless both metals are also in contact with the animal substances or the water making part of the circle. resemblance of the fluid put in motion by the foregoing experiments to the electrical fluid. the fluid set in motion by the application of the metals to each other, and to animal bodies or to water, agrees with or resembles the electrical fluid in the following respects. like the electrical fluid, it communicates the sense of pungency to the tongue. like the electrical fluid, it is conveyed readily by water, blood, the bodies of animals, the metals; and is arrested in its course by glass, sealing-wax, _&c._ it passes, with similar rapidity, through the bodies of animals. like the electrical fluid, it excites the activity of the vessels of a living animal, as the pain it gives and hemorrhagy it produces seem to prove. hence perhaps it might be employed with advantage in amenorrhoea. it excites convulsions of the muscles in the same manner, and with the same effects as electricity. when the metals and animal are kept steadily in contact with each other, the convulsions cease, or an equilibrium seems to be produced, as after discharging a leyden phial. the nervous fluid or energy not the same with the electrical nor with the fluid put in motion by the foregoing experiments. that the nervous fluid is the same with the electrical, or with the fluid which is put in motion by the foregoing experiments, is, i apprehend, disproved by the following circumstances. 1. without stating the difficulty there is in conceiving how the electrical fluid can be accumulated by or confined within our nervous system, we may observe that where the electrical fluid, or fluid resembling that put in motion by the foregoing experiments, is accumulated by an animal, such as the torpedo or gymnotus, a proper apparatus is given to the animal, by means of which it is enabled to collect and to discharge this fluid. 2. the nervous power is excited by chemical or by mechanical stimuli; and, on the other hand, is destroyed by opium and other poisons, which cannot be imagined to act on the electrical fluid. 3. i have, i apprehend, refuted the theory of doctors galvani, valli and others, which supposes that the nerve is electrified _plus_ and the muscle _minus_, resembling the leyden phial, by shewing that the muscles are convulsed where there is no communication between them and the metals, but by the medium of the nerve; or when the metals are applied to different parts of the nerve alone, without touching the muscles which are convulsed, and when the muscle which is convulsed makes no part of the circle in which the matter that is put in motion passes. 4. i have proved, that the muscles are convulsed whilst the current of the electrical matter is passing from them and from the smaller branches of the nerves into their trunks; and as a muscle is never thrown into action by the nervous energy, except when this passes from the trunk of the nerve into its branches, and from these into the muscle, it appears that when, in these experiments, the muscles were convulsed, the nervous and the electrical fluids were moving in opposite directions; from which we may infer, that, in their nature, they differ essentially from each other. 5. the nervous energy is stopped by a tight ligature or by the transverse incision of a nerve, although its divided parts are thereafter placed in contact with each other; whereas the electrical fluid or the fluid excited by the metals, passes readily, downwards or upwards, along a nerve which has been tied or cut. 6. after the limb of a living animal has been amputated, frequent convulsions of the same muscles may be excited by applying mechanical or chemical stimuli to its nerves; whereas electrical matter discharges itself suddenly. hence i conclude, 1. that the fluid, which, on the application of metalline bodies to animals, occasions convulsions of their muscles, is electrical, or resembles greatly the electrical fluid. 2. that this fluid does not operate directly on the muscular fibres, but merely by the medium of their nerves. 3. that this fluid and the nervous fluid or energy are not the same, but differ essentially in their nature. 4. that this fluid acts merely as a stimulus to the nervous fluid or energy. 5. that these experiments have merely shown a new mode of exciting the nervous fluid or energy, without throwing any farther or direct light on the nature of this fluid or energy. finis. footnotes: [1] see edin. phys. ess. vol. iii. [2] see edin. phys. ess. vol. iii. [3] see observations on the nervous system, 1783, chap. x. and xi. [4] see fontana sur les poisons, 1781, p. 267. [5] see fontana, p. 293. [6] fontana, p. 244. [7] fontana, p. 259. [8] fontana, p. 112. p. 259. [9] fontana, p. 142. [10] edin. phys. ess. published in 1771, p. 363. [11] very small portions of different metals, applied as above described, have astonishing effects; and although i have found that large portions of the metals produced convulsions, when smaller had failed, or that they produced stronger convulsions; yet the effects are by no means proportioned to the weight of the metals employed, nor to the extent of their surfaces which are suddenly brought into contact. in most of my experiments, i employed a plate of zinc, about five inches long, three inches broad, and about one-third of an inch thick; and a gold probe, somewhat thicker and longer than the probes surgeons commonly use. [12] see dr fowler's book, p. 85. [13] after reading to the royal society, on the 3d of june, an account of this experiment, which i had made in the beginning of may, i found, from an ingenious publication of my pupil dr fowler, which i received that evening, that the same experiment had been performed by him. transcriber's note: --obvious print and punctuation errors were corrected. university of kansas publications museum of natural history volume 17, no. 13, pp. 559-578, 3 pls. 17-19 date, april 5, 1968 descriptions of new hylid frogs from méxico and central america by william e. duellman university of kansas lawrence 1968 university of kansas publications, museum of natural history editors: e. raymond hall, chairman, henry s. fitch, frank b. cross volume 17, no. 13, pp. 559-578, 3 pls. 17-19 published april 5, 1968 university of kansas lawrence, kansas printed by robert r. (bob) sanders, state printer topeka, kansas 1968 31-9420 descriptions of new hylid frogs from méxico and central america by william e. duellman biological exploration of méxico and central america has revealed the presence of a diverse fauna, elements of which have undergone speciation in separate areas within the relatively small region. some genera of amphibians, especially _eleutherodactylus_ and _hyla_, are represented by many species having small geographic ranges in méxico and central america. most of the species of _hyla_ inhabiting the lowlands have been known to science for many years, and most of the novelties today are found in the less accessible highlands. no fewer than 19 new species of hylid frogs have been discovered and named from méxico and central america in the past decade. in the spring and summer of 1966 i studied hylid frogs in many parts of southern méxico and central america; the field work was designed to obtain specimens and data that would resolve certain systematic problems. to a certain extent the studies were successful, but in the course of the work five previously unknown hylids were discovered; these are named and described in this paper. the only species described herein that i do not know in life is one of _plectrohyla_ that has been represented in museum collections for several years but was not obtained in my own field work. in this paper i am presenting diagnoses, descriptions, and brief comments on the relationships of five new species and one subspecies. more exhaustive accounts will be included in a monograph, now in preparation, on the middle american hylids. for use of comparative material used in the preparation of this paper, i am indebted to richard j. baldauf, texas cooperative wildlife collection (tcwc); charles m. bogert, american museum of natural history (amnh); james a. peters, united states national museum (usnm); hobart m. smith, university of illinois museum of natural history (uimnh); charles f. walker, university of michigan museum of zoology (ummz); and ernest e. williams, museum of comparative zoology (mcz). ku refers to the university of kansas museum of natural history. i am especially grateful for help in obtaining specimens and data to linda trueb, who accompanied me throughout méxico and central america, where we were joined by john d. lynch in costa rica and charles w. myers in panamá. linda trueb offered helpful suggestions in the course of preparing the manuscript, and david m. dennis skillfully prepared the illustrations which more accurately depict the frogs than my written descriptions; both of these persons have my thanks for their contributions. ratibor hartmann of finca santa clara, chiriquí, panamá, made possible our travels to the río changena on the atlantic slopes of bocas del toro. field work in costa rica was facilitated by the organization of tropical studies through the courtesy of stephen b. preston and norman scott. rodolfo hernandez corzo of the dirección general de la fauna silvestre provided the necessary permits to collect in méxico. i thank each of these persons for his helpfulness and cooperation. field work in méxico and central america and the associated laboratory studies on middle american hylid frogs are supported by grants from the national science foundation (gb-1441 and gb-5818). the field work in panamá was part of a survey of the herpetofauna of that country carried out in cooperation with the gorgas memorial laboratory and supported by the national institutes of health (gm-12020). hyla xanthosticta new species plate 17 _holotype._--adult female, ku 103772, from the south fork of the río las vueltas on the south slope of volcán barba, near the northwest base of cerro chompipe, heredia province, costa rica, elevation 2100 meters; obtained on june 26, 1966, by john d. lynch. _diagnosis._--a member of the _hyla pictipes_ group (starrett, 1966), characterized by having dorsum uniform green, canthal stripe bronze-color, flanks and anterior and posterior surfaces of thighs dark brown with bright yellow spots, throat and belly yellow, and hands having only vestigial web. _description of holotype._--female having a snout-vent length of 29.3 mm.; tibia length 16.2 mm., 55.3 per cent of snout-vent length; foot length (measured from proximal edge of inner metatarsal tubercle to tip of longest toe) 14.1 mm., 48.1 per cent of snout-vent length; head length 10.0 mm., 34.1 per cent of snout-vent length; head width 10.3 mm., 35.2 per cent of snout-vent length. snout in lateral profile truncate, slightly inclined posteroventrally, in dorsal profile narrow but truncate; canthus angular; loreal region barely concave; lips thick, barely flared. snout long; distance from anterior corner of eye to nostril equal to diameter of eye; nostrils slightly protuberant, directed laterally; internarial distance, 2.6 mm.; internarial area slightly depressed; top of head slightly convex; interorbital distance 3.2 mm., 31.1 per cent of width of head; width at eyelid 2.6 mm.; 24.3 per cent of width of head. diameter of eye 3.0 mm.; thin dermal fold extending posteriorly from posterior corner of eye, above tympanum, to point above insertion of arm. tympanum distinct, its diameter half that of eye. axillary membrane absent; arms slender; thin scalloped dermal fold on ventrolateral edge of forearm; thin dermal fold on wrist; fingers long, tapering; length of fingers from shortest to longest, 1-2-4-3; discs small, only slightly wider than digits; subarticular tubercles large; distal tubercle on third finger broad, flat; distal tubercle on fourth finger strongly bifid; supernumerary tubercles large, round, closely spaced irregularly on proximal segments of digits; prepollex moderately enlarged. web lacking between first and second fingers, vestigial between second and third fingers, extending from middle of antepenultimate phalanx of third to base of penultimate phalanx of fourth. heels overlap by about one-third length of shank when hind limbs adpressed; tibiotarsal articulation extends to anterior edge of eye; thin transverse dermal fold on heel; scalloped dermal fold along outer edge of tarsus; inner metatarsal tubercle large, flat, elliptical, visible from above; toes long, slender; length of toes from longest to shortest, 1-2-5-3-4; discs small, barely wider than digits; subarticular tubercles large, round, subconical; supernumerary tubercles few, scattered on proximal segments of digits; toes about two-thirds webbed; webbing extending from middle of penultimate phalanx of first toe to middle of penultimate phalanx of second, from distal end of penultimate phalanx of second to base of penultimate of third, from distal end of penultimate phalanx of third to middle of antepenultimate of fourth to middle of penultimate of fifth toe. anal opening directed posteroventrally at level of mid-thigh, bordered below by large tubercles; anal sheath lacking. skin smooth on dorsum except for small scattered tubercles, granular on belly and posteroventral surfaces of thighs. tongue round, emarginate, barely free behind. prevomerine teeth 5-5, on large ovoid elevations at level of posterior edges of small round choanae. color (in preservative): dark purplish brown above, brown on limbs; first three fingers and first three toes creamy yellow; other digits brown; flanks dark brown with white spots; anterior and posterior surfaces of thighs and inner surfaces of shanks brown with cream-colored spots. white stripes on edge of upper lip, ventrolateral edge of forearm, outer edge of tarsus, and above anus. chin and throat white; belly and ventral surfaces of limbs cream-color. color (in life): dorsum green, palest on sides of head; dorsal surfaces of thighs tan; canthal stripe bronze-tan (reddish copper at night); flanks, anterior and posterior surfaces of thighs, and inner surfaces of tarsi brown with bright yellow spots. throat and belly pale yellow; ventral surfaces of limbs dull, dark yellow; large, bright yellow spot on anteroventral surface of thigh; bright yellow tubercles on median part of ventral surface of thigh. anal area dark brown with white stripe above and yellow stripe below; white stripe on outer edge of forearm, outer edge of tarsus, and edge of upper lip. iris gold-color with fine black reticulations and faint reddish suffusion medially; palpebral membrane clear. _comparisons._--_hyla xanthosticta_ is a member of the _hyla pictipes_ group that contains _debilis_, _pictipes_, _rivularis_, and _tica_. from all of these, _xanthosticta_ differs by having large yellow spots on the flanks and thighs, a white labial stripe, and a large yellow spot on the proximal ventral surface of each thigh. females of _hyla pictipes_ have small creamy yellow spots on the flanks and thighs but have dark spots on the venter; furthermore, _pictipes_ lacks white stripes on the upper lip and above the anus, lacks a canthal stripe, and has larger discs and less webbing on the hand. _hyla tica_ differs from _xanthosticta_ by having white mottling on the flanks, dark transverse bands on the limbs, and larger discs, and lacks yellow spots on the thighs, and white stripes on the upper lip, limbs, and above the anus. _hyla rivularis_ is notably different in having a tan dorsum and creamy yellow venter with black flecks; moreover, _rivularis_ lacks spots on the flanks and thighs and white stripes on the upper lip, limbs, and above the anus. of all of the species in the _pictipes_ group, _xanthosticta_ most closely resembles _debilis_. this species has a dull green dorsum, usually flecked with brown or black, and a creamy white venter. the flanks of _debilis_ are creamy white with small brown flecks, and the anterior and posterior surfaces of the thighs are bright yellow. _hyla debilis_ has a dull tan canthal stripe and white spots on the upper lip; the webbing on the hand is slightly more extensive, and the discs are slightly larger, in _debilis_ than in _xanthosticta_. the presence of the large yellow spots on the flanks and thighs in combination with the uniformly green dorsum and yellow venter immediately distinguishes _hyla xanthosticta_ from all other known species of middle american hylids. _remarks._--the only known specimen of _hyla xanthosticta_ was perched at night on a leaf about one meter above the ground. the frog was found in humid upper montane forest characterized by large oaks supporting many bromeliads and heavy growths of mosses. two other members of the _hyla pictipes_ group--_pictipes_ and _rivularis_--were abundant along a stream in the oak forest. the specific name is derived from the greek _xanthos_ meaning yellow and the greek _stiktos_ meaning spotted, and alludes to the diagnostic yellow spots on the flanks and thighs. hyla pseudopuma infucata new subspecies plate 17 _holotype._--adult male, ku 101770 from the río changena, bocas del toro province, panamá, elevation 830 meters; obtained may 18, 1966, by william e. duellman. _paratypes._--ku 101771-80; mcz 55251-2, and ummz 126811-12, same locality; collected may 18-22, 1966, by william e. duellman. _diagnosis._--a subspecies of _hyla pseudopuma_ characterized by having dark red, instead of yellow, in groin and on anterior and posterior surfaces of thighs; white stripe above anal opening; and blunt snout. _description of holotype._--adult male having a snout-vent length of 37.8 mm.; tibia length 20.6 mm., 54.4 per cent of snout-vent length; foot length (measured from proximal edge of inner metatarsal tubercle) 18.6 mm., 49.2 per cent of snout-vent length; head length 11.9 mm., 31.4 per cent of snout-vent length; head width 11.4 mm., 30.1 per cent of snout-vent length. snout in lateral profile bluntly rounded, in dorsal profile truncate; canthus rounded; loreal region barely concave; lips thick, moderately flared. snout short, distance from anterior corner of eye to nostril equal to about three-fourths diameter of eye; nostrils slightly protuberant, directed dorsolaterally; internarial distance 2.6 mm.; internarial area not depressed; top of head flat; interorbital distance 3.8 mm., 33.3 per cent of width of head; width of eyelid 3.2 mm., 28.2 per cent of width of head. eye large, protuberant, diameter 4.3 mm.; thin dermal fold extending posteriorly from posterior corner of eye, obscuring upper edge of tympanum, curving downward to point above insertion of arm. tympanum distinct except dorsally, its diameter 51.1 per cent that of eye, separated from eye by distance equal to diameter of tympanum. axillary membrane absent; arms moderately robust; dermal fold on outer edge of forearm indistinct, interrupted; transverse fold on wrist weak; fingers short, stocky; length of fingers from shortest to longest, 1-2-4-3; discs large, width of that on third finger 2.6 mm., larger than tympanum; subarticular tubercles moderately small, flat, none distinctly bifid; supernumerary tubercles conical, present on proximal segments; prepollex enlarged, bearing nuptial excrescence composed of many minute horny spinules; webbing absent between first and second fingers, extending from middle of antepenultimate phalanx of second to base of antepenultimate phalanx of third and beyond to base of penultimate phalanx of fourth finger. heels overlap by about one-third length of tarsus when hind limbs adpressed; tibiotarsal articulation extends to anterior corner of eye; transverse dermal fold on heel; tarsal fold absent; inner metatarsal tubercle long, elliptical, flat, barely visible from above; outer metatarsal tubercle small, conical; toes moderately long, stout; length of toes from shortest to longest, 1-2-3-5-4; discs nearly as large as those on fingers; subarticular tubercles small, flat; supernumerary tubercles large, conical, pigmented, in single row on proximal segments of each toe; toes about two-thirds webbed; webbing extending from distal end of penultimate phalanx of first toe to base of penultimate phalanx of second, from distal end of penultimate phalanx of second to middle of antepenultimate of third, from distal end of penultimate phalanx of third to base of penultimate of fourth to distal end of penultimate of fifth toe. anal opening directed posteriorly at level of upper surfaces of thighs, bordered below by vertical flesh folds; anal sheath absent. skin of belly, ventral surfaces of arms, and proximal posteroventral surfaces of thighs granular, elsewhere smooth. tongue ovoid, about twice as long as wide, shallowly notched posteriorly, barely free behind. prevomerine teeth 5-6, situated on transverse ridges between posterior borders of small round choanae. vocal slit extending from midlateral edge of tongue to angle of jaw. color (in preservative): dorsum grayish tan with large brown blotch extending from eyelids to middle of back, limbs marked with brown transverse bars, 2 on each forearm, 3 on each thigh, shank, and foot. flanks dark gray with white spots; groin, anterior and posterior surfaces of thighs, ventral surfaces of shanks, and inner edges of feet orange-tan; anal region dark brown, bordered above by white stripe; belly and chin creamy white, latter with grayish brown flecks. color (in life): dorsum yellowish tan with olive-brown markings by night and uniform pale yellowish tan by day; axilla, inner surface of elbow, groin, anterior and posterior surfaces of thighs, ventral surfaces of thighs and shanks, inner surfaces of feet, and dorsal surfaces of first three toes tomato red; flanks dark blue with yellow spots and reticulations. throat, chest, and anterior part of belly creamy white; posterior part of belly orange, becoming red in extreme posterior region; throat flecked with brown; iris pale bronze with black reticulations; palpebral membrane clear above, yellowish tan below; nuptial excrescenses dark brown. _variation._--the discussion of variation is based on the type series plus 31 specimens (ku 101781-811) from the río claro near its junction with the río changena, at an elevation of 910 meters. females are slightly larger than males, but do not differ significantly in proportions (table 1). all specimens have the diagnostic red legs and blue flanks with yellow spots or mottling, but the dorsal pattern is highly variable. in most individuals the dark markings on the dorsum are a solid color, but in some the borders of the marks are dark, and the interior of each mark is nearly the same color as the rest of the dorsum. a triangular dark mark with the anterolateral corners on the eyelids is present in all specimens. in some individuals the posteriorly directed apex of this triangular mark is connected to the apex of another triangular mark on the back; in other individuals the marks are narrowly separated, whereas in a few specimens the marks are broadly connected. a dark blotch usually is present on the posterior end of the body. one specimen (ku 101771) has many small white spots on the dorsum. the white stripe above the anus is invariably present, and the transverse bars on the limbs are present in all specimens, although they are indistinct in some individuals. the pattern on the flanks varies from three or four large spots to many (30-44) small spots. all males have dark flecks or reticulations on the throat; in some individuals the chest and belly are heavily flecked. although the amount of flecking is much less in most females, one individual is as heavily flecked on the throat and belly as any male. the change in coloration in this frog is noteworthy. the following description of metachrosis in seven specimens from the río claro illustrates the change. at night the frogs were yellowish tan above with slightly darker dorsal markings. the axilla, groin, anterior and posterior surfaces of the thighs, ventral surfaces of the hind limbs, and webbing on the hands and feet were tomato red. by day, some individuals became creamy yellow, others ashy white, and others grayish tan. the flanks were dark blue with yellow spots. _comparisons._--the population of frogs described here closely resembles _hyla pseudopuma_ günther in the highlands of costa rica. both have the same kind of, and variation in, dorsal markings; conical, pigmented supernumerary tubercles on the toes; bilobate vocal sac; and large prepollex bearing horny nuptial spinules. although at present no evidence for intergradation exists, the population described here is considered to be a subspecies of _hyla pseudopuma_. the two subspecies exhibit few differences in size and proportions, except that the tympanum is larger in _pseudopuma_ (table 1). _hyla p. pseudopuma_ has dark brown or yellowish tan thighs and brown flanks with a few creamy white spots; the groin in some specimens is pale blue. the red color on the limbs characteristic of _infucata_ is lacking in _pseudopuma_, which also lacks the white stripe above the anus characteristic of _infucata_. the only noticeable morphological difference between the subspecies, except in the size of the tympanum, is the shape of the snout. in _infucata_ the snout is bluntly rounded in lateral profile and truncate in dorsal profile, whereas in _pseudopuma_ the snout is more acutely rounded in lateral profile and acuminate in dorsal profile (fig. 1). this external difference is correlated with the nature of the underlying premaxillaries. in _infucata_ the premaxillaries lie in a transverse plane and have short, nearly vertical alary processes, whereas in _pseudopuma_ the premaxillaries lie at a slight angle and have longer alary processes that are inclined posteriorly. table 1.--variation in certain measurements and proportions in the subspecies of hyla pseudopuma. (means are given in parentheses below the observed range.) ==================+========+==+==========+==========+==========+========== | | |snout-vent| tibia | foot | tympanum/ subspecies | sex | n| length | length/ | length/ | eye | | | | s-v l | s-v l | ------------------+--------+--+----------+----------+----------+---------_h. p. pseudopuma_|[male] |31| 37.6-41.4| 48.5-55.0| 44.7-52.9| 47.6-66.7 |[symbol]| | (39.7) | (51.5) | (48.2) | (57.6) | | | | | | _h. p. infucata_ |[male] |31| 37.6-42.9| 50.7-56.9| 43.3-49.5| 41.8-55.3 |[symbol]| | (39.5) | (53.4) | (47.4) | (49.2) | | | | | | _h. p. pseudopuma_|[female]|11| 41.6-47.6| 49.1-53.8| 45.4-49.5| 57.9-71.4 |[symbol]| | (44.3) | (51.7) | (47.6) | (60.2) | | | | | | _h. p. infucata_ |[female]|11| 41.1-45.6| 50.0-55.4| 43.5-47.4| 47.6-56.1 |[symbol]| | (42.6) | (52.6) | (45.8) | (50.1) ------------------+--------+--+----------+----------+----------+---------[illustration: fig. 1. lateral views of the heads of _hyla pseudopuma pseudopuma_ (left, ku 64884) and _h. p. infucata_ (right, ku 101784). × 4.] the only other frog in central america having red webs and anterior and posterior surfaces of the thighs is _hyla loquax_, which has a broad head, extensive axillary membrane, single median vocal sac, and uniformly creamy yellow flanks; furthermore, _loquax_ lacks conical, pigmented supernumerary tubercles on the toes and a large prepollex with horny nuptial spinules. _hyla rufitela_ has red webbing, but in no other diagnostic feature resembles _infucata_, for _rufitela_ is green above, white below, and has angular prevomerine dentigerous ridges. _remarks._--most specimens of _hyla pseudopuma infucata_ were found on bushes and low trees at night. three males and one clasping pair were on the ground. the habitat is humid lower montane forest where the amount of rainfall is high. although no breeding was observed nor calls heard, it is presumed that this subspecies breeds in shallow, temporary pools, like those utilized by the nominate subspecies. the two localities where _hyla pseudopuma infucata_ is known are in the maze of ridges north of cerro pando on the panamanian-costa rican border. the río claro is a tributary of the río changena, in turn a tributary of the río changuinola, which receives many streams and rivers draining the northern slopes of the highlands in bocas del toro province before flowing into the caribbean. we reached the río claro and río changena by walking from finca santa clara on the pacific slopes, over the continental divide, and down the north slope of cerro pando. the subspecific name is derived from the latin _infucatus_, meaning painted, in allusion to the red colors on the limbs and webs. hyla pellita new species plate 18 _holotype._--adult male, ku 100970 from 33 kilometers north of san gabriel mixtepec, oaxaca, méxico, elevation 1675 meters; obtained on february 20, 1966, by william e. duellman and linda trueb. _paratypes._--ku 100971-2 collected with the holotype and ku 100974-5 from 30 kilometers north of san gabriel mixtepec, oaxaca, méxico, elevation 1530 meters; same date and collectors. _diagnosis._--a small yellowish tan _hyla_ characterized by tympanum concealed, anal opening not bordered below by large tubercles, brown bands on shanks, and dark flecks on roof of mouth anteriorly. _description of holotype._--adult male having snout-vent length of 27.3 mm.; tibia length 13.7 mm., 50.2 per cent of snout-vent length; foot length (measured from proximal edge of inner metatarsal tubercle to tip of longest toe) 11.7 mm., 42.9 per cent of snout-vent length; head length 8.7 mm., 31.9 per cent of snout-vent length. snout in lateral profile truncate, rounded above, in dorsal profile rounded; canthus angular; loreal region slightly concave; lips thin, flared. snout moderately long, distance from anterior corner of eye to nostril slightly less than diameter of eye; nostrils slightly protuberant, directed anterolaterally; internarial area barely depressed; distance between nostrils 2.6 mm.; top of head flat; interorbital distance 3.2 mm., 34.4 per cent of width of head. diameter of eye 2.7 mm.; thin dermal fold extending from posterior corner of eye to point above insertion of arm; tympanum absent, not visible through skin. axillary membrane absent; forearms moderately slender, having indistinct tuberculate fold on ventrolateral edge, lacking distinct transverse fold on wrist; fingers short; length of fingers from shortest to longest, 1-4-2-3, fourth nearly as long as second; discs small, about half again as wide as digits; subarticular tubercules large, round, flattened, distal ones on third and fourth fingers bifid; supernumerary tubercles large, round, present only on proximal segments; prepollex barely enlarged, lacking nuptial excrecence. web lacking between first and second fingers, extending from base of penultimate phalanx of second to base of antepenultimate phalanx of third, from middle of antepenultimate phalanx of third to distal end of antepenultimate of fourth finger. heels overlap by about one-fifth length of shank when hind limbs adpressed; tibiotarsal articulation extending to middle of eye, tarsal fold present, extending full length of tarsus; inner metatarsal tubercle flat, ovoid, partly visible from above; outer metatarsal tubercle absent; toes short; length of toes from shortest to longest, 1-2-3-5-4; discs small, about two-thirds width of those on fingers; subarticular tubercles small, round; supernumerary tubercles small, flattened, irregularly arranged on proximal segments. toes three-fourth webbed; web extending from base of disc of first to middle of penultimate phalanx of second, from base of disc of second to middle of penultimate phalanx of third, from base of disc of third to base of penultimate phalanx of fourth and to base of disc of fifth toe. anal opening directed posteriorly at level of dorsal surfaces of thighs, bordered below by vertical dermal folds and few small tubercles; anal sheath absent. skin heavily granular on throat, chest, belly and ventral surfaces of thighs, smooth elsewhere. tongue cordiform, deeply notched posteriorly, barely free behind. prevomerine teeth 3-4, situated on short elevations between small round choanae; vocal slits absent. color (in preservative): pale tan above with dark brown mark in occipital region and large irregular brown mark extending from scapular region to sacral region; anterior and posterior surfaces of thighs and flanks lacking pigment; dorsal surfaces of arms, shanks, and feet tan with brown transverse bars (two on each forearm, two on left shank, one on right shank, and one on each foot); entire dorsal surfaces, except hands and first four toes, peppered with black; venter creamy white; roof of mouth between, and anterior to, choanae speckled with minute black flecks. color (in life): yellowish tan above with reddish brown flecks (later changed to pale brown with dull olive-green interorbital bar, blotch on back, and flecks on dorsum); hands, feet, and anterior and posterior surfaces of thighs dull yellow; belly white; creamy white stripes on outer edge of forearm, foot, and above anus; iris pale silver-bronze. _variation._--three adult males (including holotype) have snout-vent lengths of 25.2-27.3 (mean 26.5) mm., and two females have 28.6 and 31.6 (mean 30.1) mm. one juvenile has a snout-vent length of 21.5 mm. no significant variation occurs in the proportions. males have 6-8, and females have 8 and 9, prevomerine teeth. the tympanum is completely concealed in all specimens. all specimens have distinct transverse bars on the limbs; the number of bars on the shank varies from one to four. two individuals are dark brown dorsally; in these the small black flecks either are not visible or are absent; flecks are present on the dorsal surfaces of four specimens that are tan or pale brown above with darker brown irregular markings. the coloration in life consisted of olive-green or olive-brown markings on the body and olive-green or brown bars on the limbs. the dorsal ground color was yellowish tan or pale brown in all individuals. _comparisons._--_hyla pellita_ differs from all known middle american _hyla_, except _mixe_, _mixomaculata_, _nubicola_, and _pinorum_, by having a concealed tympanum. the first three of these differ from _pellita_ in greater size and by having many bands on the hind limbs. superficially _h. pellita_ resembles _hyla pinorum_, which likewise has a tan dorsum with irregular markings and limbs with transverse bars. _hyla pinorum_ differs from _pellita_ by having a proportionately larger head, no transverse bands on the thighs, and large tubercles below the anus. furthermore, in _pinorum_ the quadratojugal articulates with the maxillary, whereas in _pellita_ the quadratojugal is reduced to a small spur and does not articulate with the maxillary. _remarks._--all individuals were found on low vegetation along streams in cloud forest at night. no specimens were found when the type locality was revisited in august, 1966. duellman (1960) placed _hyla pinorum_ taylor in the synonymy of _ptychohyla leonhardschultzei_ ahl. at that time only the holotype, a female, of _h. pinorum_ was known. in 1964 kraig adler and i independently collected frogs and associated tadpoles in guerrero that subsequently proved to be _hyla pinorum_ and provided evidence that _hyla pinorum_ is not conspecific with _ptychohyla leonhardschultzei_. the specific name _pellita_ is latin, meaning covered with skin, and is here used in reference to the complete concealment of the tympanum beneath the skin. hyla siopela new species plate 18 _holotype._--adult male, ku 100981, from a small stream on the west slope of cofre de perote, veracruz, méxico, elevation 2500-2550 meters; obtained on july 30, 1966, by william e. duellman. _paratypes._--ku 100976-80, 100982-5, same locality, date, and collector; ku 105628-9, same locality, obtained on june 18, 1966, by howard l. freeman; uimnh 57687-57701, same locality, obtained on july 30-31, 1964, by macreay j. landy and john d. lynch. _diagnosis._--a member of the _hyla bistincta_ group characterized by truncate snout with short rostral keel; fingers having little webbing and bearing large discs; axillary membrane absent; thoracic fold weak; prepollex large, flat, bearing small nuptial spines; vocal slits absent; dorsum green or tan with small irregular dark spots; flanks mottled. _description of holotype._--adult male having a snout-vent length of 44.3 mm.; tibia length 21.1 mm., 47.6 per cent of snout-vent length; foot length (measured from proximal edge of inner metatarsal tubercle to tip of longest toe) 20.4 mm., 47.2 per cent of snout-vent length; head length 13.4 mm., 30.2 per cent of snout-vent length; head width 13.7 mm., 30.9 per cent of snout-vent length. snout in lateral profile truncate, in dorsal profile truncate with weak vertical rostral keel; canthus angular; loreal region slightly concave; lips thick, not flaring; snout short; nostrils barely protuberant, directed dorsolaterally, situated about four-fifths distance from anterior corner of eye to tip of snout; internarial distance 3.6 mm.; internarial area not depressed; top of head slightly convex; interorbital distance 4.9 mm., 35.8 per cent of width of head; width of eyelid 3.5 mm., 26.0 per cent of width of head. diameter of eye 4.7 mm.; heavy dermal fold curving posteroventrally from posterior corner of eye, covering upper one-third of tympanum, to insertion of arm; tympanum barely distinct, its diameter 2.2 mm., 46.8 per cent that of eye, separated from eye by distance equal to half again diameter of tympanum. axillary membrane absent; thoracic fold weak; arms moderately robust; fold on wrist heavy; fingers long, slender; length of fingers from shortest to longest, 1-2-4-3; discs large, that on third finger as large as tympanum; subarticular tubercles moderately small, round, none bifid; supernumerary tubercles small, some barely distinguishable, in single row on proximal segment of each digit; prepollex greatly enlarged, flat ventrally, bearing nuptial excrescence composed of minute horny spinules; webbing between first two fingers vestigial; web connecting other fingers at bases of penultimate phalanges of second and fourth, and base of antepenultimate phalanx of third fingers. heels overlap by about one-third length of shank when hind limbs adpressed; tibiotarsal articulation extends to posterior edge of orbit; transverse dermal fold on heel; tarsal fold thin, distinct, extending length of tarsus; inner metatarsal tubercle large, elongate, flat, visible from above; outer metatarsal tubercle absent; toes moderately long, slender; length of toes from shortest to longest, 1-2-3-5-4; discs slightly smaller than those on fingers; subarticular tubercles moderately small, round; supernumerary tubercles small, in single row on proximal segment of each digit; toes about two-thirds webbed; webbing extends from middle of penultimate phalanx of first toe to base of penultimate phalanx of second, from middle of penultimate of second to middle of antepenultimate of third, from middle of penultimate of third to middle of antepenultimate of fourth to middle of penultimate phalanx of fifth toe. anal opening directed posteriorly at level of mid-thigh; anal sheath short. skin granular on chin, belly, and posteroventral surfaces of thighs, smooth elsewhere. tongue broadly cordiform, notched posteriorly, barely free behind. prevomerine teeth 4-4, situated on posteromedially inclined elevations between small ovoid choanae. vocal slits absent. color (in preservative): dull grayish brown above with small, irregularly-shaped black spots on head, back, and limbs; flanks gray mottled with creamy tan; anterior and posterior surfaces of thighs tan; belly dull creamy tan; throat marked with gray blotches; anal region and posterodorsal surfaces of thighs marked with small white spots. color (in life): dorsum pale green with black spots and reticulations; flanks mottled dark brown and creamy white; outer edges of feet silvery white with brown spots; anterior and posterior surfaces of thighs dull brown; webbing and first three toes dull yellowish tan; belly creamy gray; throat silvery white, mottled with gray; iris dull bronze-color with black reticulations; palpebral membrane clear. _variation._--the snout-vent length in seven adult males is 47.2-50.0 mm., and in five females, 45.1-52.5 mm. in neither sex do the average proportions differ noticeably from those of the holotype, except that the tympanum is relatively larger in females. the ratio of the diameter of the tympanum to that of the eye is 0.363-0.468 (mean 0.438) in males and 0.500-0.545 (mean 0.516) in females. the average number of prevomerine teeth in males is 7.9, in females 8.4. in life dorsal coloration varied from pale green to olive-green with darker green or black flecks or reticulations, or pinkish tan to brown with dark brown flecks or reticulations. some preserved specimens have relatively few dark flecks, whereas in most specimens the dorsum is heavily marked. all specimens have some white markings above the anus and on the posterodorsal surfaces of the thighs, but in some individuals the white flecks are expanded and interconnected forming an irregular white line. juveniles have a notably different coloration in life. the dorsum is uniform pale green; the anterior and posterior surfaces of the thighs, fingers, first three toes, and webbing are deep yellow. the anal stripe is creamy white, and the flanks are pale gray with black flecks. the upper lip, supratympanic fold, and canthal stripe are a bronze color. the belly is pale yellow with a silvery cast on the throat. juveniles having snout-vent lengths from 24.5 to 36.6 mm. are so colored in life, and uniform dark bluish gray dorsally in preservative. _comparisons._--the absence of a quadratojugal and the presence of a greatly enlarged, non-projecting prepollex place _hyla siopela_ in the _hyla bistincta_ group (see duellman, 1964, and adler, 1965). the presence of a rostral keel separates _hyla siopela_ from other members of the _hyla bistincta_ group, which is composed of two species having long anal sheaths (_bistincta_ and _pentheter_), two small species having axillary membranes and lacking nuptial excrescences in breeding males (_charadricola_ and _chryses_), and three species (_crassa_, _pachyderma_, and _robertsorum_) having short heads, round snouts, short anal sheaths, and nuptial excrescences in breeding males. _hyla siopela_ differs from the last three species in the shape of the snout and from each in certain structural features; _h. crassa_ has fully webbed feet; _h. pachyderma_ has large nuptial spines, and _h. robertsorum_ has more webbing and a shorter tarsal fold. furthermore, the venter in _h. robertsorum_ is brown with creamy white flecks. in structure and coloration _h. arborescandens_ resembles _siopela_, but the former is smaller, and males of _arborescandens_ have vocal slits. _remarks._--this description brings to eight the number of species now recognized in the _hyla bistincta_ group. _hyla siopela_ is most closely related to _hyla robertsorum_ from the high mountains of the sierra madre oriental in northern puebla and eastern hidalgo. possibly the four species now recognized in the _crassa_ subgroup (_crassa_, _pachyderma_, _robertsorum_, and _siopela_) are only subspecies of a single species, but differences in the amount of webbing in _crassa_ and the nature of the nuptial excrescenses in _pachyderma_ indicate that they are distinct species. the type locality of _hyla siopela_ is a small stream cascading down the western slope of cofre de perote; the lower reaches of the stream can be reached by a dirt road leading east from the village of perote for about 2 kilometers to a small park. the frogs were found in the stream at elevations of 50 to 100 meters higher than the park. the stream flows through a ravine supporting open, dry pine forest. although the stream was searched thoroughly in february, 1966, no frogs were found. in july, 1966, adults and juveniles were found in crevices and under overhanging rocks behind small cascades and waterfalls by day and sitting on rocks and branches in the spray of cascades at night. the specific name _siopela_ is derived from the greek _siopelos_, meaning silent, and alludes to the absence of a voice in this species. hyla altipotens new species plate 19 _holotype._--adult male, ku 101001, from 37 kilometers (by road) north of san gabriel mixtepec (kilometer post 183 on road from oaxaca to puerto escondido), oaxaca, méxico, elevation 1860 meters; obtained on february 19, 1966, by william e. duellman. _paratypes._--ku 101002-6 collected at the same locality on february 19 and 20, 1966, by william e. duellman, and ku 101008 from 33 kilometers (by road) north of san gabriel mixtepec, oaxaca, méxico, elevation 1675 meters; obtained on february 20, 1966, by linda trueb. _diagnosis._--a member of the _hyla taeniopus_ group characterized by a yellow venter, yellow flecks on posterior surfaces of thighs, bronze-colored stripe from snout, along canthus and edge of upper eyelid to point above arm, pointed snout, smooth dorsum, and no sexual dimorphism in shape of snout. _description of holotype._--adult male having a snout-vent length of 68.8 mm.; tibia length 36.2 mm., 52.6 per cent of snout-vent length; foot length (measured from proximal edge of inner metatarsal tubercle) 31.1 mm., 45.2 per cent of snout-vent length; head length 20.2 mm., 29.4 per cent of snout-vent length; head width 20.0 mm., 29.1 per cent of snout-vent length. snout in lateral profile acutely rounded, protruding beyond tip of lower jaw, in dorsal profile pointed; canthus angular; loreal region flat; lips thick, barely flared. snout long; nostrils slightly protuberant, directed dorsolaterally, situated about two-thirds distance from anterior corner of eye to tip of snout; internarial distance 5.6 mm.; internarial area slightly depressed; top of head flat; interorbital distance 6.0 mm., 30.0 per cent of width of head; width of eyelid 4.7 mm., 23.5 per cent of width of head. diameter of eye 6.2 mm.; heavy dermal fold extending from posterior corner of eye, over upper edge of tympanum to point above insertion of arm; tympanum distinct, its diameter 3.4 mm., 54.8 per cent of that of the eye, separated from eye by distance equal to diameter of tympanum. axillary membrane absent; arms moderately robust, lacking dermal fold on lateral edge of forearm, having transverse fold on wrist; fingers moderately short, broad; length of fingers from shortest to longest, 1-2-4-3; discs large, that on third finger one-fourth larger than tympanum; subarticular tubercles large, round, none bifid; supernumerary tubercles large, granule-like, present only on proximal segments; prepollex enlarged, not bearing nuptial excrescence. fingers about one-half webbed; webbing connects first and second fingers at level of distal end of antepenultimate phalanx, extending from middle of penultimate phalanx of second finger to middle of antepenultimate phalanx of third, and between bases of penultimate phalanges of third and fourth fingers. heels overlap by about one-half length of shank when hind limbs adpressed; tibiotarsal articulation extends to point between eye and nostril; thin transverse dermal fold on heel; tarsal fold strong, extending full length of tarsus; inner metatarsal tubercle small, flat, elongate, barely visible from above; outer metatarsal tubercle small, conical; toes moderately long, stout; length of toes from shortest to longest, 1-2-3-5-4; discs slightly smaller than those on fingers; subarticular tubercles large, round, subconical; supernumerary tubercles large, conical, in single row on proximal segment of each digit; toes about four-fifths webbed; webbing extending from base of disc on first to base of disc on second to base of penultimate phalanx of third toe, from base of disc on third to base of penultimate phalanx of fourth to base of disc of fifth toe. anal opening directed posteroventrally at mid-level of thighs; anal sheath long, tubular. skin smooth on dorsum and on ventral surfaces of shanks, granular on throat, belly, and ventral surfaces of arms and thighs. tongue ovoid, widest posteriorly, neither notched nor free behind. prevomerine teeth 6-7, situated on robust transverse ridges between small, ovoid choanae. vocal slits absent. testes large, ovoid, granular; length of left testis 11.0 mm. color (in preservative): brown above with many darker brown spots and narrow middorsal stripe on back; six or seven dark brown transverse bars on each segment of hind limbs and four bars on each forearm; flanks white with dark brown spots; anterior surfaces of thighs creamy white with brown reticulations; posterior surfaces of thighs dark brown with creamy yellow flecks; stripe on snout, canthus, edge of upper eyelid, and supratympanic fold tan; ventral surfaces of feet brown; rest of venter creamy white; stripe above anus white. color (in life): green above with slightly darker green spots; dorsal surfaces of upper arms and thighs tan with green transverse bars; upper surfaces of forearms and shanks green with darker green transverse bars; feet, fourth and fifth toes, and third and fourth fingers tan with brown transverse bars; other fingers and toes tan with brown flecks. ventral surfaces creamy yellow, brightest on throat and chest; flanks and anterior surfaces of thighs bright creamy yellow with dark brown reticulations and spots; posterior surfaces of thighs and ventral surfaces of feet dark brown with yellow flecks; ventral surfaces of hands and webbing on hands and feet yellowish tan. labial stripe tan; stripes on outer edge of forearm, along outer edge of foot, and above anus cream-color; stripe on canthus, edge of upper eyelid, and on supratympanic fold bronze-color. iris pale bronze with black reticulations and faint median, horizontal copper-colored streak; pupil horizontally elliptical with ventral notch; palpebral membrane clear above, pale bluish green with brown reticulations below. _variation._--in life all individuals had creamy yellow venters and yellow flanks and anterior surfaces of thighs with brown or black spots and mottling. most of the adults were colored like the holotype, but one was a much darker olive-green, and one was uniform brown above with a dark brown middorsal stripe. most subadults (snout-vent lengths 31.6-50.1 mm.) were pale reddish tan above with darker reddish brown bars on the limbs and blotches on the back. the side of the head was dark brown and the stripe along the canthus, edge of upper eyelid, and supratympanic fold was yellowish tan. some individuals had a dark brown middorsal stripe. the posterior surfaces of the thighs were dull yellowish tan; yellow flecks were present in the larger individuals. table 2.--variation in measurements and proportions in hyla altipotens. (means are given in parentheses below the observed range.) =========+===+=============+=============+=============+============= | | snout | tibia | foot | tympanum/ sex | n | vent | length/ | length/ | eye | | length | s-v l | s-v l | ---------+---+-------------+-------------+-------------+------------ males | 5 | 68.8-75.1 | 52.6-55.8 | 28.1-30.0 | 41.4-55.2 | | (70.7) | (53.7) | (29.2) | (50.6) | | | | | females | 2 | 69.4-75.3 | 55.8-56.2 | 28.8-31.1 | 53.3-63.0 | | (72.4) | (56.0) | (29.9) | (58.8) ---------+---+-------------+-------------+-------------+------------the number of transverse bars on each thigh and shank varies from five to eight. the white stripe above the anus and the stripe from the snout along the side of the head are invariably present. in some of the largest individuals the brown reticulations on the anterior surface of the thigh extend onto the ventral surface; in these specimens brown flecks are present on the ventral surfaces on the shanks. the tympanum is proportionately larger in females than in males; the variation in size and proportions is given in table 2. the total number of prevomerine teeth varies from 13 to 18 (mean, 15) in five adult males and from 10 to 12 (mean 11) in two females. the testes in all adult males are granular, ovoid in shape, and greatly enlarged. the lengths of the left testis in each of the five males are 11.0 to 23.5 (mean 14.6) mm. _comparisons._--on the basis of external appearance and certain cranial characters (large frontoparietal fontanelle, broad sphenethmoid, large nasals broadly separated medially having thin lateral processes articulating with the palatines, short squamosal not articulating with the maxillary, and quadratojugal present and articulating with the maxillary), _hyla altipotens_ can be associated with the _hyla taeniopus_ group (duellman, 1965, lynch and smith, 1966). _hyla altipotens_ can be distinguished from all of the other members of the group by its narrow head, pointed snout in both sexes, and uniformly yellow throat and belly. small brown individuals of _hyla altipotens_ superficially resemble adult _hyla pinorum_. the latter species has a covered tympanum, less webbing on the hands, and a short, blunt snout. _remarks._--this stream-breeding frog is like _hyla taeniopus_ in having greatly enlarged testes, which possibly through the production of vast quantities of sperm are an adaptation for successful breeding in torrential streams (duellman, 1965:164). all individuals were found in trees and bushes near streams in cloud forest at night in february. the type locality is the same as that of _hyla pentheter_ and _hyla thorectes_, discovered by kraig adler in june, 1964. our field work there in february, 1966, resulted in finding _hyla altipotens_, _h. pellita_, and _ptychohyla leonhardschultzei_, but no individuals of the species found by adler. a visit to the same locality in august, 1966, revealed no individuals of either _h. altipotens_ or _pellita_; instead _pentheter_ and _thorectes_ were found along the stream. duellman (1965:166) listed a specimen (tcwc 16184) of _hyla chaneque_ from los fustes, 3 kilometers east of san sebastian, oaxaca. reëxamination of this specimen reveals that it is _hyla altipotens_. the frog was obtained by dilford carter on april 29, 1960; it was under a rock at the edge of a stream in an oak-pine-cypress association at an elevation of 1800 meters. the specific name _altipotens_ is latin, meaning mighty, here used in allusion to the supposed potentiality of fertilization by the production of vast quantities of sperm in the large testes. plectrohyla hartwegi new species plate 19 _holotype._--adult male, ummz 94428, from barrejonel (19 kilometers west of chicomuselo), chiapas, méxico, elevation 1000 meters, obtained on june 12, 1941, by eizi matuda. _paratypes._--two subadult males, ku 58873 from parajé el triunfo, north of mapastepec, chiapas, méxico, elevation 2050 meters, obtained on may 12, 1960, by miguel alvarez del toro, and uimnh 40837 from cerro azul oaxaca, méxico, obtained on march 7, 1956, by thomas macdougall. _diagnosis._--a _plectrohyla_ having a bifid prepollex, bold mottling on flanks and ventral surfaces of shanks, and vertical dark bars on anterior and posterior surfaces of thighs, and lacking vocal slits and outer tarsal fold. _description of holotype._--adult male having a snout-vent length of 63.8 mm.; tibia length 34.9 mm., 54.7 per cent of snout-vent length; foot length (measured from proximal edge of inner metatarsal tubercle to tip of longest toe) 31.1 mm., 48.7 per cent of snout-vent length; head length 19.7 mm., 30.9 per cent of snout-vent length; head width 22.6 mm., 35.4 per cent of snout-vent length. snout short, distance from level of anterior edge of orbit to tip of snout 70.6 per cent of length of eye; snout in lateral profile angular, sloping abruptly from nostrils to jaw, in dorsal profile bluntly rounded, lacking rostral keel; canthal ridge thickened; loreal region deeply concave; lips thick, barely flared. nostrils small, barely protuberant, directed anterolaterally, situated about two-thirds distance from eye to tip of snout; internarial distance 5.6 mm. internarial area barely depressed near convergence of canthal ridges; top of head flat; interorbital distance 6.4 mm., 28.3 per cent of head width; diameter of eye 6.8 mm.; width of eyelid 5.5 mm., 24.3 per cent of head width. heavy dermal fold extending posteriorly from posterior edge of orbit, covering upper edge of tympanum; two thinner folds extending ventrally from longitudinal heavy fold covering posterior edge of tympanum; anterior and ventral edges of tympanum distinct; length of tympanum 2.9 mm., 42.6 per cent of diameter of eye. axillary membrane absent; arms robust, forearm not noticeably heavier than upper arm; distinct transverse fold on wrist. fingers long, moderately slender; length of fingers from shortest to longest, 1-2-4-3; discs moderately large, that on third finger larger than tympanum; webbing vestigial; subarticular tubercles small, conical; terminal tubercle on fourth finger somewhat flattened; supernumerary tubercles small, in one row on proximal segment of fourth finger and in two rows on proximal segments of other fingers; prepollex greatly enlarged, barely bifurcate; spines not protruding through skin; distal spine much longer than proximal one (fig. 2). heels overlap by about one-third length of shank when hind limbs adpressed; tibiotarsal articulation extends slightly beyond snout; heavy transverse dermal fold on heel; inner tarsal fold heavy, extending full length of tarsus; outer tarsal fold absent; inner metatarsal tubercle high, elliptical, visible from above, outer metatarsal tubercle absent. toes long, slender; length of toes from shortest to longest, 1-2-5-3-4; fifth toe nearly as long as third; discs small; subarticular tubercles small, round; supernumerary tubercles small, in single row on proximal segment of each digit; toes about three-fourths webbed; webbing extending from base of disc of first toe to base of penultimate phalanx of third, from base of disc of third to base of penultimate phalanx of fourth to base of disc of fifth toe. [illustration: fig. 2. palmar view of prepollical spine of right hand of _plectrohyla hartwegi_ (ummz 94428). × 5.] anal opening directed posteroventrally at level of mid-thigh; anal sheath long with membranous connection to posterior surfaces of thighs. skin on dorsal surfaces finely tuberculate; that on throat, chest, belly, and ventral surfaces of thighs granular, that on ventral surfaces of arms and shanks smooth. tongue nearly round, free posteriorly for about one-fourth its length, barely notched behind. upper jaw shallowly notched medially. maxillary-premaxillary teeth 38-35; prevomerine teeth 5-5, situated on small elliptical elevations between quadrangular choanae; vocal slits absent. color (in preservative): uniform dull brown above and creamy yellow below; flanks brown with creamy yellow mottling and dark brown spots in groin; anterior surfaces of thighs creamy yellow with two broad, vertical, dark brown bars proximally and two narrower, dull brown bars distally; posterior surfaces of thighs brown with dark brown vertical bars, interspaces cream-colored or brown. ventral surfaces of shanks creamy yellow with bold brown reticulations. _variation._--the paratypes are smaller, having snout-vent lengths of 48.3 and 41.8 mm. in these specimens the ratio of the length of the tibia to the snout-vent length is 55.9 and 57.9 per cent, and the ratio of the diameter of the tympanum to the diameter of the eye is 47.3 and 43.6 per cent, respectively. both specimens have 4-4 prevomerine teeth; one specimen has 37-36, and the other has 40-37, maxillary-premaxillary teeth. in these small specimens the supratympanic fold is thin, and the arms are not so robust as in the holotype. in one specimen (ku 58873) the tongue is not notched posteriorly. the terminal subarticular tubercle on each fourth finger is broad and flattened in uimnh 40837, but conical in ku 58873. both specimens have bold creamy-yellow and dark brown mottling on the flanks and dark brown reticulations on the ventral surfaces of the shanks. there are two dark brown vertical bars on the anterior and posterior surfaces of each thigh in ku 58873 and three bars on each surface in uimnh 40837. _comparisons._--_plectrohyla hartwegi_ differs from all known species in the genus by having boldly mottled flanks, dark reticulations on the ventral surfaces of the shanks, and dark vertical bars on the shanks. in all of the other species the anterior and posterior surfaces of the thighs are unmarked, and the flanks are either plain or marked with small spots or flecks. structurally, _p. hartwegi_ belongs in the _guatemalensis_ group of the genus, containing _avia_, _glandulosa_, _guatemalensis_, and _pycnochila_. the species in this group lack vocal slits and have either large, rectangular, or bifid prepollices. _plectrohyla hartwegi_ differs from all of these species, except _pycnochila_, by having a tuberculate, instead of a smooth, dorsum, and _hartwegi_ differs from _pycnochila_ by having a bifid, instead of a rectangular, prepollex. _remarks._--the known distribution of _plectrohyla hartwegi_ includes three localities at elevations of 1000 to 2050 meters in the sierra madre of chiapas and extreme eastern oaxaca. the specimen from parajé el triunfo was found in a rocky stream in cloud forest at an elevation of 2050 meters. one _plectrohyla sagorum_ was obtained from the same stream. eizi matuda sent the holotype to the late dr. norman hartweg, who recognized that the specimen was unique but was reluctant to name the species on the basis of a single specimen. now that two additional specimens are available, it seems appropriate to associate hartweg's name with this new species of _plectrohyla_, a genus that hartweg first adequately defined. literature cited adler, k. 1965. three new frogs of the genus _hyla_ from the sierra madre del sur of méxico. occas. papers mus. zool. univ. michigan, 642:1-18, pl. 1, december 16. duellman, w. e. 1960. synonymy, variation, and distribution of _ptychohyla leonhardschultzei_ ahl. studies of american hylid frogs, iv. herpetologica, 16:191-197, september 23. 1964. a review of the frogs of the _hyla bistincta_ group. univ. kansas publ. mus. nat. hist., 15:469-491, march 2. 1965. frogs of the _hyla taeniopus_ group. copeia, 2:159-168, june 25. starrett, p. 1966. rediscovery of _hyla pictipes_ cope, with description of a new montane stream _hyla_ from costa rica. bull. south. california acad. sci., 65 (1):17-28, march. _transmitted july 11, 1967._ [illustration: plate 17 upper figure, _hyla xanthosticta_ (ku 103772); lower figure, _hyla pseudopuma infucata_ (ku 101770). × 2.] [illustration: plate 18 upper figure, _hyla pellita_ (ku 100973); middle figure, _hyla pellita_ (ku 100970); lower figure, _hyla siopela_ (ku 100977). × 2.] [illustration: plate 19 upper figure, _hyla altipotens_ (ku 101001); lower figure, _plectrohyla hartwegi_ (ummz 94428). × 1.] 31-9420 transcriber's notes: obvious punctuation errors corrected. spelling corrections: 'mexico' to 'méxico' on title page for consistency. 'granulelike' to 'granule-like' ...large, granule-like, present... 'midlevel' to 'mid-level' ... at mid-level of thighs... 'posterio' to 'posterior' ...anterior and posterior surfaces of thighs... 'flank' to 'flanks' ...mottling on flanks... transcriber's notes in this plain text version of the book, italic typeface is represented with _underscores_, and small capital typeface is represented in upper case. [=e] represents a macron (horizontal line) over an e. [female] represents the symbol for female. [male] represents the symbol for male. a small number of inconsistencies and typographical errors have been changed in the text. these are listed at the end of this book. the title page and verso are in error in stating that the pages run 275 to 306. this should read 276-307. * * * * * university of kansas publications museum of natural history volume 8, no. 4, pp. 275-306, 9 figs. in text february 10, 1956 a field study of the kansas ant-eating frog, gastrophryne olivacea by henry s. fitch university of kansas lawrence 1956 university of kansas publications, museum of natural history editors: e. raymond hall, chairman, a. byron leonard, robert w. wilson volume 8, no. 4, pp. 275-306, 9 figs. in text published february 10, 1956 university of kansas lawrence, kansas printed by ferd voiland. jr., state printer topeka, kansas 1956 25-7819 a field study of the kansas ant-eating frog, gastrophryne olivacea by henry s. fitch introduction the ant-eating frog is one of the smallest species of vertebrates on the university of kansas natural history reservation, but individually it is one of the most numerous. the species is important in the over-all ecology; its biomass often exceeds that of larger species of vertebrates. because of secretive and subterranean habits, however, its abundance and effects on community associates are largely obscured. the reservation, where my field study was made, is the most northeastern section in douglas county, kansas, and is approximately 5½ miles north and 2½ miles east of the university campus at lawrence. the locality represents one of the northernmost occurrences of the species, genus, and family. the family microhylidae is a large one, and most of its representatives are specialized for a subterranean existence and a diet of termites or ants. the many subfamilies of microhylids all have distributions centering in the regions bordering the indian ocean, from south africa and madagascar to the east indies, new guinea, and australia (parker, 1934). only one subfamily, the microhylinae, is represented in the new world, where it has some 17 genera (de carvalho, 1954) nearly all of which are tropical. _g. olivacea_, extending north into extreme southern nebraska (loomis, 1945: 211), ranges farther north than any other american species. in the old world only _kaloula borealis_ has a comparable northward distribution. occurring in the vicinity of peiping (pope, 1931: 587), it reaches approximately the same latitude as does _gastrophryne_ in nebraska. the great majority of microhylid genera and species are confined to the tropics. nearly all ant-eating frogs seen on the reservation have been caught and examined and individually marked. by november 1, 1954, 1215 individuals had been recorded with a total of 1472 captures. in the summer of 1950, richard freiburg studied this frog on the reservation and his findings (1951) led to a better understanding of its natural history. the numbers of frogs studied by him however, were relatively small and the field work was limited to the one summer. the data now at hand, representing six consecutive years, 1949 through 1954, serve to supplement those obtained by freiburg, corroborating and extending his conclusions in most instances, and also indicating that certain of his tentative conclusions need to be revised. while the present report was in preparation, anderson (1954) published an excellent account of the ecology of the eastern species _g. carolinensis_ in southern louisiana. anderson's findings concerning this closely related species in a much different environment have been especially valuable as a basis for comparison. the two species are basically similar in their habits and ecology but many minor differences are indicated. some of these differences result from the differing environments where anderson's study and my own were made and others certainly result from innate genetic differences between the species. the frog with which this report is concerned is the _microhyla carolinensis olivacea_ of the check list (schmidt, 1953: 77) and recent authors. de carvalho (1954: 12) resurrected the generic name, _gastrophryne_, for the american species formerly included in _microhyla_, and presented seemingly valid morphological evidence for this plausible generic separation. _g. olivacea_ is obviously closely related to _g. carolinensis_; the differences are not greater than those to be expected between well marked subspecies. nevertheless, in eastern oklahoma and eastern texas, where the ranges meet, the two kinds have been found to maintain their distinctness, differing in coloration, behavior, calls, and time of breeding. hecht and matalas (1946: 2) found seeming intergrades from the area of overlapping in eastern texas, but some specimens from this same area were typical of each form. their study was limited to preserved material, in which some characters probably were obscured. more field work throughout the zone of contact is needed. the evidence of intergradation obtained so far seems to be somewhat equivocal. besides _g. olivacea_ and typical _g. carolinensis_ there are several named forms in the genus, including some of doubtful status. the name _mazatlanensis_ has been applied to a southwestern population, which seems to be a well marked subspecies of _olivacea_, but as yet _mazatlanensis_ has been collected at few localities and the evidence of intergradation is meager. the names _areolata_ and _texensis_ have been applied to populations in texas. hecht and matalas (1946: 3) consider _areolata_ to be a synonym of _olivacea_, applied to a population showing intergradation with _carolinensis_, but wright and wright (1949: 568) consider _areolata_ to be a distinct subspecies. _g. texensis_ generally has been considered to be a synonym of _olivacea_. other species of the genus include the tropical _g. usta_, _g. elegans_ and _g. pictiventris_. of the vernacular names hitherto applied to _g. olivacea_ none seems appropriate; i propose to call the species the kansas ant-eating frog because of its range extending over most of the state, and because of its specialized food habits. the type locality, originally stated to be "kansas and nebraska" (hallowell, 1856: 252) has been restricted to fort riley, kansas (smith and taylor, 1950: 358). members of the genus have most often been referred to as toads rather than frogs because of their more toadlike appearance and habits. however, this family belongs to the firmisternial or froglike division of the salientia and the terms "frog" and "toad," originally applied to _rana_ and _bufo_ respectively, have been extended to include assemblages of related genera or families. members of the genus and family usually have been called "narrow-mouthed" toads from the old generic name _engystoma_, a synonym of _gastrophryne_. _g. olivacea_ usually has been referred to as the texas narrow-mouthed toad, or western narrow-mouthed toad. the latter name is inappropriate because the geographic range is between that of a more western representative (_mazatlanensis_) and a more eastern one (_carolinensis_). the names _texensis_, _areolata_ and _carolinensis_ have all been applied to populations in texas, and it is questionable whether typical _olivacea_ even extends into texas. habitat in the northeastern part of kansas at least, rocky slopes in open woods seem to provide optimum habitat conditions. this type of habitat has been described by several earlier workers in this same area, dice (1923: 46), smith (1934: 503) and freiburg (1951: 375). smith (1950: 113) stated that in kansas this frog is found in wooded areas, and that rocks are the usual cover, but he mentioned that outside of kansas it is often found in mesquite flats that are devoid of rocks. freiburg's field work was done almost entirely on the reservation and was concentrated in "skink woods" and vicinity, where much of my own field work, both before and afterward, was concentrated. on the reservation and in nearby counties of kansas, the habitat preferences of the ant-eating frog and the five-lined skink largely coincide. in an account of the five-lined skink on the reservation, i have described several study areas in some detail (fitch, 1954: 37-41). it was on these same study areas (quarry, skink woods, rat woods) that most of the frogs were obtained. although _g. olivacea_ thrives in an open-woodland habitat in this part of its range, it seems to be essentially a grassland species, and it occurs throughout approximately the southern half of the great plains region. bragg (1943: 76) emphasized that in oklahoma it is widely distributed over the state, occupying a variety of habitats, with little ecological restriction. bragg noted, however, that the species is rarely, if ever, found on extensive river flood plains. on various occasions i have heard _gastrophryne_ choruses in a slough two miles south of the reservation. this slough is in the kaw river flood plain and is two miles from the bluffs where the habitat of rocky wooded slopes begins that has been considered typical of the species in northeastern kansas. it seems that the frogs using this slough are not drawn from the populations living on the bluffs as mud creek, a kaw river tributary, intervenes. the creek channel at times of heavy rainfall, carries a torrent of swirling water which might present a barrier to migrating frogs as they are not strong swimmers. the frogs could easily find suitable breeding places much nearer to the bluffs. those using the slough are almost certainly permanent inhabitants of the river flood plain. the area in the neighborhood of the slough, where the frogs probably live, include fields of alfalfa and other cultivated crops, weedy fallow fields, and the marshy margins of the slough. in these situations burrows of rodents, notably those of the pocket gopher (_geomys bursarius_), would provide subterranean shelter for the frogs, which are not efficient diggers. the frogs may live in many situations such as this where they have been overlooked. in the absence of flat rocks providing hiding places at the soil surface, the frogs would rarely be found by a collector. the volume and carrying quality of the voice are much less than in other common anurans. large breeding choruses might be overlooked unless the observer happened to come within a few yards of them. most of the recorded habitats and localities of occurrence may be those where the frog happens to be most in evidence to human observers, rather than those that are limiting to it or even typical of it. on september 20, 1954, after heavy rains, juveniles dispersing from breeding ponds were in a wide variety of situations, including most of the habitat types represented on the reservation. along a small dry gully in an eroded field formerly cultivated, and reverted to tall grass prairie (big bluestem, little bluestem, switch grass, indian grass), the frogs were numerous. many of them were flushed by my footsteps from cracks in the soil along the gully banks. in reaching this area the frogs had moved up a wooded slope from the pond, crossed the limestone outcrop area at the hilltop edge, and wandered away from the woods and rocks, out into the prairie habitat. in this prairie habitat there were no rocks providing hiding places at the soil surface, but burrows of the vole (_microtus ochrogaster_) and other small rodents provided an abundance of subterranean shelter. in the summer of 1955 the frogs were seen frequently in this same area, especially when the soil was wet from recent rain. when the surface of the soil was dry, none could be found and presumably all stayed in deep cracks and burrows. anderson (1954: 17) indicated that _g. carolinensis_ in louisiana likewise occurs in diverse habitats, being sufficiently adaptable to satisfy its basic requirements in various ways. behavior ordinarily the ant-eating frog stays beneath the soil surface, in cracks or holes or beneath rocks. probably it obtains its food in such situations, and rarely wanders on the surface. the occasional individuals found moving about above ground are in most instances flushed from their shelters by the vibrations of the observer's footsteps. on numerous occasions i have noticed individuals, startled by nearby footfalls, dart from cracks or under rocks and scuttle away in search of other shelter. such behavior suggests that digging predators may be important natural enemies. the gait is a combination of running and short hops that are usually only an inch or two in length. the flat pointed head seems to be in contact with the ground or very near to it as the animal moves about rapidly and erratically. the frog has a proclivity for squeezing into holes and cracks, or beneath objects on the ground. the burst of activity by one that is startled lasts for only a few seconds. then the frog stops abruptly, usually concealed wholly or in part by some object. having stopped it tends to rely on concealment for protection and may allow close approach before it flushes again. less frequently, undisturbed individuals have been seen wandering on the soil surface. such wandering occurs chiefly at night. diurnal wandering may occur in relatively cool weather when night temperatures are too low for the frogs to be active. wandering above ground is limited to times when the soil and vegetation are wet, mainly during heavy rains and immediately afterward. pitfalls made from gallon cans buried in the ground with tops open and flush with the soil surface were installed in 1949 in several places along hilltop rock outcrops where the frogs were abundant. the number of frogs caught from day to day under varying weather-conditions provided evidence as to the factors controlling surface activity. after nights of unusually heavy rainfall, a dozen frogs, or even several dozen, might be found in each of the more productive pitfalls. a few more might be caught on the following night, and occasional stragglers as long as the soil remained damp with heavy dew. activity is greatest on hot summer nights. below 20° c. there is little surface activity but individuals that had body temperatures as low as 16° c. have been found moving about. frogs uncovered in their hiding places beneath flat rocks often remained motionless depending on concealment for protection, but if further disturbed, they made off with the running and hopping gait already described. although they were not swift, they were elusive because of their sudden changes of direction and the ease with which they found shelter. when actually grasped, a frog would struggle only momentarily, then would become limp with its legs extended. the viscous dermal secretions copiously produced by a frog being handled made the animal so slippery that after a few seconds it might slide from the captor's grasp, and always was quick to escape when such an opportunity was presented. temperature relationships ant-eating frogs are active over a temperature range of at least 16° c. to 37.6° c. they tolerate high temperatures that would be lethal to many other kinds of amphibians, but are more sensitive to low temperatures than any of the other local species, and as a result their seasonal schedule resembles that of the larger lizards and snakes more than those of other local amphibians. the latter become active earlier in the spring. earliest recorded dates when the frogs were found active in the course of the present study from 1950 to 1955 were in april every year; the 20th, 25th, 24th, 2nd, 25th, and 21st. latest dates when the frogs were found in the six years of the study were: october 22, 1949; october 13, 1950; october 7, 1951; august 24, 1952; august 18, 1953; and october 27, 1954 (excluding two late stragglers caught in a pitfall on december 5). severe drought caused unseasonably early retirement in 1952 and 1953. body temperatures of the frogs were taken with a small mercury thermometer of the type described by bogert (1949: 197); the bulb was used to force open the mouth and was thrust down the gullet into the stomach. to prevent conduction of heat from the hand, the frog was held down through several layers of cloth, at the spot where it was discovered, until the temperature reading could be made. this required approximately five seconds. [illustration: fig. 1. temperatures of ant-eating frogs grouped in one-degree intervals; upper figure is of frogs found active in the open, and lower is of those found under shelter. the frogs are active over a temperature range of more than 20 degrees, and show no clear cut preference within this range.] most of the 79 frogs of which temperatures were measured, were found under shelter, chiefly beneath flat rocks. the rocks most utilized were in open situations, exposed to sunshine. most of the frogs were in contact with the warmed undersurfaces of such rocks. forty-three of the frogs, approximately 54.5 percent, were in the eight-degree range between 24° and 31° c. probably the preferred temperatures lie within this range. the highest body temperature recorded, 37.6° c., was in a frog which "froze" and remained motionless in the sunshine for half a minute after the rock sheltering it was overturned. probably its temperature was several degrees lower while it was sheltered by the rock. other unusually high temperatures were recorded in newly metamorphosed frogs found hiding in piles of decaying vegetation near the edge of the pond, on hot afternoons of late august. temperatures ranged from 17.0° to 30.7° in frogs that were found actually moving about. several with relatively low temperatures, 22° to 17°, were juveniles travelling in rain or mist on cool days. these frogs, having relatively low temperature, were sluggish in their movements, as compared with individuals at the upper end of the temperature range. [illustration: fig. 2. body temperatures and nearby air temperatures for frogs found under natural conditions. dots represent frogs found under shelter; circles represent those found in the open.] after the first frost each year the frogs usually could not be found, either in the open or in their usual hiding places beneath rocks. they probably had retired to deep subterranean hibernation sites. the only exception was in 1954, when two immature frogs were found together in a pitfall on the morning of december 5 after a rain of .55 inches ending many weeks of drought. air temperature had been little above 10° c. that night, but had often been below freezing in the preceding five weeks. reactions of these same two individuals to low temperatures were tested in the laboratory. at a body temperature of 11° c. they were extremely sluggish. they were capable of slow, waddling movements, but were reluctant to move and tended to crouch motionless. even when they were prodded, they usually did not move away, but merely flinched slightly. at 6° c. they were even more sluggish, and seemed incapable of locomotion, as they could not be induced to hop or walk by prodding with a fine wire. when placed upside down on a flat surface, they could turn over, but did so slowly, sometimes only after a minute or more had elapsed. respiratory throat movements numbered 46 and 60 per minute. breeding many observers have noted that breeding activity is initiated by heavy rains in summer. in my experience precipitation of at least two inches within a few days is necessary to bring forth large breeding choruses. with smaller amounts of precipitation only stragglers or small aggregations are present at the breeding ponds. tanner (1950: 48) stated that in three years of observation, near lawrence, kansas, the first storms to bring large numbers of males to the breeding ponds occurred on june 20, 1947, june 18, 1948, and may 1, 1949. in 1954 the frogs were recorded first on april 25, but these were under massive boulders, and were still semi-torpid. frogs were found fully active, in numbers, under small flat rocks on may 7. they were found frequently thereafter. on the afternoon of may 13, the third consecutive day with temperature slightly above 21° c., low croaking of a frog was heard among rocks at an old abandoned quarry. throughout the remainder of may, calling was heard frequently at the quarry on warm, sunny afternoons. often several were calling within an area of a few square yards, answering each other and maintaining a regular sequence. in the last week of may rains were frequent, and the precipitation totalled 2.09 inches. on june 1 and 2 also, there were heavy rains totalling 2.26 inches. on the evening of june 2 many frogs were calling at a pond ½ mile south of the reservation, and one was heard at the pond on the reservation. by the evening of june 4, dozens were calling in shallow water along the edge of this pond in dense _polygonum_ and other weeds. there was sporadic calling even in daylight and there was a great chorus each evening for the next few days, but its volume rapidly diminished. in mid-june a system of drift fences and funnel traps was installed 200 yards west of the pond in the dry bottom of an old diversion ditch leading from the pond. the ditch constituted the boundary between bottomland pasture and a wooded slope, and therefore was a natural travelway. the object of the installation was to intercept and catch small animals travelling along the ditch bottom. the drift fence was w-shaped, with a funnel trap at the apex of each cone so that the animals travelling in either direction would be caught. the numbers of frogs caught from time to time during the summer provided information as to their responses to weather in migrating to the pond. table 1. numbers of frogs caught within two days after rain in funnel traps in 1954, from mid-june, to the time of first frost. date precipitation no. of in inches caught frogs july 1 2.02 8 july 10 .11 none july 16 1.26 none july 20-21 .94 3 july 24 .38 2 july 28 .29 none august 1-2 3.22 31 august 6-7-8 2.43 none august 12 .28 none august 16 .29 none august 19-22 .70 none august 27-28 1.05 none september 9 .50 none september 29-30 .38 none october 4 .74 none october 12-14 3.51 none from the positions of the traps and drift fences, it was obvious that all of the frogs that were caught were travelling toward the pond. capture of an equal number moving away from the pond a few days afterward might have been expected but none at all was caught while making a return trip. therefore it seems that the frogs returned by a different route to their home ranges after breeding. of necessity they make the return trip under conditions drier than those that prevail on the pondward trip, which is usually made in a downpour. probably the return travel is slower, more leisurely, and with more tendency to keep to sheltered situations. the call is a bleat, resembling that of a sheep, but higher, of lesser volume, and is not unlike the loud rattling buzz of an angry bee. the call is usually of three to four seconds duration, with an interval several times as long. calling males were floating, almost upright, in the water within a few yards of shore, where there was dense vegetation. the throat pouch when fully expanded is several times as large as the entire head. when a person approached to within a few yards of frogs they usually stopped calling, submerged, and swam to a place of concealment. having heard the call of typical _g. carolinensis_ in louisiana, i have the impression that it is a little shorter, more sheeplike, and less insectlike than that of _g. olivacea_. the call of _gastrophryne_ is of such peculiar quality that it is difficult to describe. different observers have described it in different terms. stebbins (1951: 391) has described the call in greatest detail, and also has quoted from the descriptions of it previously published. these descriptions include the following: "high, shrill buzz"; "buzz, harsh and metallic"; "like an electric buzzer"; "like bees at close range but more like sheep at a distance"; "bleating baa"; "shrill, long-drawn quaw quaw"; "whistled wh[=e][=e] followed by a bleat." stebbins observed breeding choruses (_mazatlanensis_) at peña blanca springs, arizona, and stated that sometimes three or four called more or less together, but that they seldom started simultaneously. occasionally many voices would be heard in unison followed by an interval of silence, but this performance was erratic. at the pond on the reservation i noted this same tendency many times. after a lull the chorus would begin with a few sporadic croaks, then four or five or even more frogs would be calling simultaneously from an area of a few square yards. anderson (_op. cit._: 34) found that in small groups of calling _g. carolinensis_ there was a distinct tendency to maintain a definite pattern in the sequence of the calls. one "dominant" individual would initiate a series of calls, and others each in turn would take up the chorus. pairing takes place soon after the breeding aggregations are formed. on the night of june 4, 1954, a clasping pair was captured and kept in the laboratory in a large jar of water. this pair did not separate, and spawning occurred between noon and 1:30 p. m. on june 5. when the newly laid eggs were discovered at 1:30 p. m. most of them were in a surface film. some were attached to submerged leaves and a few rested on the bottom. the pair was still joined, but the male was actually clasping only part of the time, and as the frogs moved about in the water, it became evident that they were adhering to each other by the areas of skin contact, which were glued together by their dermal secretion. they were unable to separate immediately, even when they struggled to do so. they were observed for approximately 15 minutes before separation occurred, and during this time they were moving about actively. as they separated, the area of adhesion was discernible on the back of the female. it was u-shaped, following the ridges of the ilia and the sacrum. on august 2, 1954, after a rain of 3.22 inches, the previously mentioned funnel trap in the ditch had caught 31 ant-eating frogs. water had collected to a depth of several inches in the depression where the trap was situated. a dozen of the trapped frogs were clasping pairs. these frogs struggled vigorously as they were removed from the traps, handled and marked. as a result most of the clasping males were separated from the females. in handling those of each pair i noticed that they were glued together by dermal secretions, as were those of the pair observed on june 5. the areas of adhesion were of similar shape and location in the different pairs, and included the u-shaped ridge of the female's back and the male's belly, and the inner surfaces of the male's forelegs with the corresponding surfaces of the female's sides where the male clasped. this adhesion of the members of a pair during mating may be a normal occurrence. the copious secretion of the dermal glands is of especially glutinous quality in _gastrophryne_. the adhesion of members of a pair may have survival value. these small frogs are especially shy, and in the breeding ponds they respond to any disturbance with vigorous attempts to escape and hide. under such circumstances the adhesion may prevent separation. also, it may serve to prevent displacement of a clasping male by a rival. anderson (_op. cit._) who observed many details of the mating behavior of _g. carolinensis_, both in the laboratory and under natural conditions, mentioned no such adhesion between members of a pair. anderson (_op. cit._: 31) discussed the possibility that reproductive isolation might arise in sympatric populations, such as those of _g. carolinensis_ in southern louisiana, through inherent differences in time of spawning. however, in _g. olivacea_ at least, such isolation would be prevented by individual males returning to breed at different times in the same season. furthermore, individual differences in choice of breeding time probably result from environmental factors rather than genetic factors in most instances. in _g. olivacea_ in kansas, time of breeding is controlled by the distribution of heavy rainfall creating favorable conditions. onset of the breeding season may be hastened or delayed, or an entire year may be missed because of summer drought. if favorable heavy rains are well distributed throughout the summer, frogs of age classes that are not yet sexually mature in the early part of the breeding season, may comprise the bulk of the breeding population in late summer. development of eggs and larvae eggs laid on june 5 by the pair kept in the laboratory were hatching on june 7, on the average approximately 48 hours from the time of laying. by june 8 all the eggs had hatched and the tadpoles were active. on august 28 and 29 thousands of newly metamorphosed young were in evidence on wet soil at the pond margin; in some the head still was tadpolelike and they had a vestige of the tail stump. these young were remarkably uniform in size, 15 to 16 mm. (the smallest one found was 14½ mm.) and almost all of them had originated from eggs laid after heavy precipitation, totalling 3.22 inches, in the first 36 hours of august. allowing one day for adults to reach the pond and spawn, and two days more for eggs to hatch, the tadpole stage must have lasted approximately 24 days in this crop of young. wright and wright (1949: 582) stated that the tadpoles metamorphosed after 30 to 50 days, and that the newly metamorphosed frogs are 10 to 12 mm. in length. length of time required for larval development probably varies a great deal depending on the interaction of several factors such as temperature and food supply. growth little has been recorded concerning the growth rate of _gastrophryne_ or the time required for it to attain sexual maturity. wright (1932) found that _g. carolinensis_ in the okefinokee swamp region has a mean metamorphosing-size of 10.8 mm. young thought to be those recently emerged from their first hibernation were those in the size group 15.0 to 20.0 mm., while the frogs in the 20 to 27 mm. size class and those in the 27 to 36 mm. class were interpreted as representing two successively older annual age classes. anderson (1954: 41) thought he could recognize four successive annual age classes in the same species in southern louisiana. he found that sexual maturity is attained at a length of 21 to 24 mm. in frogs which he believed to be late in the second year of life. allowing for size differences between the two species, wright's and anderson's conclusions regarding growth in _g. carolinensis_, on the basis of size groups, are largely substantiated by my own data on the growth of marked individuals of _g. olivacea_ living under natural conditions in kansas. in 1954, an opportunity to investigate the early growth was afforded by unusually favorable circumstances. the population of frogs that emerged from hibernation in the late spring of 1954 included few, if any, that were below adult size; drought had prevented successful breeding in 1952 and 1953. heavy rains in the first week of june, 1954, and again in the first week of august, resulted in the production of two successive crops of young so widely spaced that they were easily distinguishable. some young may have been hatched after other minor rains, but certainly these were relatively few. young from the eggs laid in the first week of august were metamorphosing during the last week of august. growth in the frogs of this group can be shown by the average size and the size range of the successive samples collected. table 2. growth in frogs metamorphosed in the last week of august, 1954. ========================================================= |number in| mean size |size range time of sample | sample | in mm. | in mm. --------------------+---------+--------------+---------- august 27 to 31 | 27 | 15.55 ± .079 | 15 to 17 --------------------+---------+--------------+---------- september 11 | 114 | 17.2 ± .033 | 14 to 20 --------------------+---------+--------------+---------- september 15 to 22 | 12 | 18.7 ± .090 | 16 to 20 --------------------+---------+--------------+---------- september 27 to 30 | 37 | 19.3 ± .055 | 17 to 21.5 --------------------+---------+--------------+---------- october 1 to 7 | 62 | 20.8 ± .072 | 17 to 24 --------------------+---------+--------------+---------- october 12 to 17 | 49 | 22.3 ± .092 | 18 to 24 ========================================================= by mid-october, six weeks after metamorphosis, these frogs had increased in over-all length by approximately 50 percent. having grown a little more than 1 mm. per week on the average, they were approximately intermediate in size between small adults and newly metamorphosed young. the frogs hatched in june were present in relatively small numbers compared with those hatched in august, and were not observed metamorphosing. in late august a sample of 33 judged to belong to the june brood averaged 26.2 (22-28) mm. long. a sample of 39 from the first week of october averaged 28.1 (24.5-32) mm. frogs of this group thus were approaching small adult size late in their first growing season. such individuals possibly breed in the summer following their first hibernation, when they are a year old or a little more. because recaptured frogs were not sacrificed to determine the state of their gonads, the minimum time required to attain sexual maturity was not definitely determined. the available evidence indicates that sexual maturity is most often attained late in the second year of life, at an age of approximately two years. the darkened and distensible throat pouch of the adult male probably is the best available indicator of sexual maturity. [illustration: fig. 3. growth shown by successive samples of young ant-eating frogs of two size groups in late summer and early fall of 1954. for each sample the mean, standard deviation, and range are shown. lower series are those metamorphosed in late august, and upper series are those metamorphosed in late june.] [illustration: fig. 4. rapid growth of a young female caught in june, july, and august, 1949. presumably this individual metamorphosed late in the summer of 1948, and at the age of approximately one year it was near small adult size.] frogs that metamorphose in late summer have little time to grow before hibernating, and still are small when they emerge in spring. the smallest one found was 19 mm. long (may 19, 1951), and in each year except 1954 many such young were found that were less than 25 mm. in length in may or early june. none of the frogs marked at or near metamorphosing size has been recaptured, but the trend of early growth is well shown by table 2 and fig. 3. however, many juveniles that were captured and marked within a few weeks of metamorphosis were recaptured as adults. the selected individuals in table 3 are considered typical of growth from "half-grown" to small adult size. growth in many other individuals is shown in figs. 6 and 7. table 3. growth in frogs marked as young and recaptured as small adults. ============================================================== individual | dates | length | probable time and sex | of capture | in mm. |of metamorphosis -----------------+-----------------+---------+--------------- no. 1 [female] | august 28, 1951 | 21.5 |mid-july, 1951 | may 5, 1952 | 23 | | july 3, 1952 | 32 | | august 31, 1952 | 33 | -----------------+-----------------+---------+--------------- no. 2 [female] | june 8, 1950 | 25 |late july, 1949 | may 24, 1951 | 31 | | july 30, 1951 | 34 | | june 24, 1952 | 35 | -----------------+-----------------+---------+--------------- no. 3 [male] | august 31, 1951 | 24 |late june, 1951 | may 23, 1953 | 32 | ============================================================== [illustration: fig. 5. ant-eating frogs, a little less than twice natural size, adult and newly metamorphosed young, showing differences in size and coloration. the young is darker and has a leaflike middorsal mark which fades as growth proceeds.] the trend of growth after attainment of minimum adult size is also well shown by the records of marked individuals recaptured. many of these were marked while they were still small so that their approximate ages are known. for those recaptured in their second year, after one hibernation, length averaged 30.92 mm. some of this group were young metamorphosed late the preceding summer and still far short of adult size (as small as 23 mm.) when recaptured. others were relatively large, up to 33 mm. a group of 22 recaptured frogs known to be in their third year averaged 33.3 mm. (males 31.9, females 35.3, excluding four individuals of undetermined sex). fifteen other recaptured frogs were known to be in their fourth year at least, and some probably were older, as they were already large adults when first examined. these 15 averaged 36.6 mm. (males 34.7, females 37.9 mm.). size was similar in a sample of 58 individuals intercepted en route to the breeding pond in heavy rains of june and august, 1954. the 38 males in this sample ranged in size from 30 mm. to 38 mm., averaging 34.5. the 20 females ranged from 34 mm. to 40 mm., averaging 37.65. the large average and maximum size in this sample of a breeding population may be typical after periods of drought years have prevented successful reproduction. summer drought in 1952 and 1953 prevented breeding in those years, or, at least, it drastically reduced the numbers of young produced. one-year-old and two-year-old frogs may not have been represented at all in the sample of 58. three-year-old frogs presumably made up a substantial part of the sample, since 1951 was a year of successful breeding. [illustration: fig. 6. growth in a group of frogs, each marked while still short of adult size and mostly recaptured after lapse of one or more hibernation periods. each line connects records of an individual frog.] differences in size between species and geographic variation in size in _gastrophryne_ have been given little attention by herpetologists, but if understood, would help to clarify relationships. hecht and matalas stated in their revision (1946: 5) that size is of no importance as a taxonomic character, as typical _carolinensis_, _olivacea_, and _mazatlanensis_ all averaged approximately the same--26 to 28 mm.--females slightly larger than males. however, they arbitrarily classed as adults all individuals 22.5 mm. in length or larger, having found individuals this small that showed the darkened and distensible throat pouches characteristic of adult males. from the trend of my own measurements of _g. olivacea_ in northeastern kansas, i conclude that either many immature individuals were included in their samples, or that the populations sampled included some with individuals that were remarkably small as adults. [illustration: fig. 7. growth in another group of frogs that were marked as young or small adults and recaptured after intervals of more than a year. frogs of this group were, on the average, larger than the individuals shown in fig. 6, and they made less rapid growth.] the population which i studied may be considered typical of _g. olivacea_. they averaged large, including individuals up to 42 mm. in length, well above the maximum sizes for any reported in the literature. at metamorphosis these _olivacea_ are of approximately 50 percent greater length than _g. carolinensis_ as reported by wright and wright (1949: 573) and anderson (1954: 41). yet blair (1950: 152) observed that in eastern oklahoma, where the ranges of _olivacea_ and _carolinensis_ overlap, the latter is larger. on the basis of field and laboratory observations he tentatively concluded that one of the main barriers to interbreeding was the reluctance of the males of _carolinensis_ to clasp the smaller females of _olivacea_. that size differs in different populations, and is still poorly understood, is illustrated by the following discrepant figures from various authors. table 4. size range of adults in various populations of gastrophryne. ===============+=======================+=================+============= species or | geographic population | authority |size range of subspecies | sampled | |adults in mm. ---------------+-----------------------+-----------------+------------ | | | _olivacea_ |douglas co., kansas |present study | 31 to 42 | | | _olivacea_ |entire range |wright and wright| 19 to 38 | | (1949) | | | | _carolinensis_ |entire range |wright and wright| 20 to 36 | | (1949) | | | | _carolinensis_ |southern louisiana |anderson | 22 to 35 | | (1954) | | | | _areolata_ |southeastern texas |wright and wright| 23 to 29 | | (1949) | | | | _mazatlanensis_|arizona and new mexico |wright and wright| 22 to 30 | | (1949) | | | | _mazatlanensis_|santa cruz co., arizona|stebbins | 25.2 to 31.5 | | (1951) | ---------------+-----------------------+-----------------+------------color and pattern the color pattern changes in the course of development, and the shade of color changes in response to environmental conditions. at the time of metamorphosis, young are dark brown with specks of black and with a dark, cuneate, leaflike middorsal mark. the narrow end of this mark arises just behind the head, and the mark extends posteriorly as far as the hind leg insertions. at its widest, the mark covers about half the width of the dorsal surface. the lateral edges of the mark are sharply defined, but at its anterior and posterior ends it blends into the ground color. in most individuals smaller than 20 mm., this dorsal mark is well defined and conspicuous. as growth proceeds, however, it becomes faint. in frogs 19 to 25 mm. long the marks have disappeared. in individuals of this size the brown ground color is markedly paler than in those newly metamorphosed, but is darker than in adults. in large adults the dorsal coloration is a uniform pale tan, paler on the average in females than in males. temperature and moisture both affect the shade of coloration. in frogs that were partly desiccated, the color was unusually pale, with a distinctly greenish tint, and at high temperatures coloration tended to be relatively pale. hecht and matalas (1946) have described and figured color patterns in various populations of _gastrophryne_, demonstrating geographic trends and helping to clarify relationships. their account indicates that the dark dorsal mark present in young of _olivacea_ but not present in adults, is better developed and longer persisting in other forms. specimens of _carolinensis_, presumably adult, are figured which have the dark middorsal area contrasting with paler color of the sides. the dark area is seen to consist of dots or blotches of black pigment which may be in contact producing more or less continuous black areas, or may be separate and distinct producing a spotted pattern. pigmentation is usually most intense along the lateral edges of the dorsal leaflike mark; the central portion may be so much paler that the effect is that of a pair of dorsolateral stripes. this latter type of pattern is best developed in the population of key west, florida. hecht and matalas did not consider these insular frogs to be taxonomically distinct, because only 48 percent of specimens from the florida keys had the "key west" pattern, while 29 per cent resembled _olivacea_ and 23 per cent resembled _carolinensis_. in the southwestern subspecies (or species) _mazatlanensis_, recorded from several localities in sonora and from extreme southern arizona, the dorsal pigmentation similarly tends to be concentrated in dorsolateral bands, but is much reduced or almost absent, and there is corresponding pigmentation dorsally across the middle of the thigh, across the middle of the shank, and on the foot. when the leg is folded, these three dark areas are brought in contact with each other and with the dorsolateral body mark, if it is present, to form a continuous dark area, in a characteristic "ruptive" pattern. hecht and matalas found similar leg bars, less well developed, in certain specimens of _olivacea_ including one from gage county, nebraska, at the northern end of the known geographic range. movements freiburg (_op. cit._: 384) concluded that ant-eating frogs seem to have no individual home ranges, but wander in any direction where suitable habitat is present. however, from records covering a much longer span of time, it became increasingly evident that a frog ordinarily tends to stay within a small area, familiar to it and providing its habitat requirements. nevertheless, in all but a few instances the marked frogs recaptured were in new locations a greater or lesser distance from the site of original capture. the movements made by these frogs were of several distinct types: 1. routine day to day movements from shelter to shelter within the area familiar to the animal, the "home range." 2. shifts from one home range to another; such shifts may have been either long or short, and may have occurred abruptly or by gradual stages. 3. travel by adults to or from a breeding pond. in most or all instances these adults were regularly established in permanent home ranges, and they often moved through areas unsuitable as habitat to reach the ponds. 4. movements of dispersal in the young, recently metamorphosed and not yet settled in a regular home range. usually there was uncertainty as to which types of movements had been made by the recaptured individuals. some may have made two or three different types of movements in the interval between captures. on many occasions individuals were found beneath the same rock on two consecutive days, or occasionally on several successive days. rarely, such continued occupancy of a niche lasted several weeks. in 1949, a frog was found under the same rock on june 4, 6, 26, 27, and july 1, 3 and 11. this was an immature female, presumably metamorphosed late in the summer of 1948. during the five weeks period covered by the records, it grew from 27 mm. to 34 mm. in 1952, another individual was found under its home rock on june 23 and 30, july 2 and 3, and august 14 and 20. in 1952 a juvenile was found under a rock on may 30, june 4, and june 17. these three individuals were exceptional in their continued occupancy of the same niches. among the hundreds of others recorded, none was found more than twice in any one place. despite the fact that field work was concentrated on small areas which were worked intensively, only eight per cent of the frogs recorded were ever recaptured, and most of those were recaptured only once. only 13 individuals yielded series of records, well spaced, in two or more different years. these few individuals recaptured frequently may not be typical of the entire population. the low incidence of recaptures indicates that relatively few of the frogs present on an area at any one time have been taken. because of their secretive and subterranean habits most of the frogs are missed by a collector who searches by turning rocks, or trapping with pitfalls. therefore, even though a marked frog may survive and remain within a radius of a few hundred feet of one point for months or even years, the chances of recapture are poor. one female was caught first as a juvenile on june 8, 1950. on april 24, 1951, when first recaptured, she had grown to small adult size, and was only 18 feet from the original location. on july 30, 1951, however, she was recaptured 750 feet away. at a fourth capture on may 21, 1952, she had shifted 70 feet farther in the same direction. at the final capture on june 24, 1952, she was approximately 140 feet from both the third and fourth locations. the sequence of these records suggests that the frog had already settled in a home range at the time of her first capture in 1950, and that approximately a year later she shifted to a second home range, which was occupied for the following year, at least. [illustration: fig. 8. distances between captures in frogs marked, and recaptured after substantial intervals including one or more hibernations. distances are grouped in 25-foot intervals. for longer distances the trend is toward progressively fewer records, indicating that typical home ranges are small.] in several instances, after recaptures as far as 400 feet from the original location, frogs were again captured near an original location, suggesting that for some individuals, at least, home ranges may be as much as 400 feet in diameter. figure 8 shows that for movements of up to 400 feet, numbers of individuals gradually decrease with greater distance. for distances of more than 400 feet there are comparatively few records. of the 59 individuals recaptured after one or more hibernations, only nine had moved more than 400 feet from the original location. twenty-five were recaptured at distances of 75 feet or less. the mean distance for movement for all individuals recaptured was 72 feet. a typical home range, therefore, seems to average no more than 75 feet in radius. of the 59 individuals recaptured after one or more hibernations, 47 were adults and probably many of these had made round-trip migrations to the breeding pond. this was not actually demonstrated for any one individual, but several were captured in each of three or four different years near the same location. [illustration: fig. 9. distances between captures and elapsed time in months in marked frogs recaptured. few records are for distances more than 400 feet. there is but little tendency to longer movements in those caught after relatively long intervals.] the trend of movements differed in the sexes. males are more vagile. of 21 adult males recaptured, none was less than 40 feet from its original location, whereas six of the 26 adult females were less than 40 feet away from the original point of capture. of seven frogs that had wandered 700 feet or more, five were males. food habits according to smith (1934: 503) stomachs of many specimens, from widely scattered localities in kansas, contained only large numbers of small ants. tanner (1950: 47) described the situation of a frog found on the reservation buried in loose soil beneath a flat rock, beside an ant burrow, where, presumably, the frog could snap up the passing ants without shifting its position. anderson (_op. cit._: 21) examined alimentary tracts of 203 specimens of _carolinensis_ from louisiana, representing a year round sample for several different habitats. he found a variety of small animals including ants, termites, beetles, springtails, bugs, ear-wigs, lepidopterans, spiders, mites, centipedes, and snails. most of these prey animals were represented by few individuals, and ants were much more numerous than any of the other groups. anderson concluded that ants, termites, and small beetles were the principal foods. he noted that some of the beetles were of groups commonly found in ant colonies. tanner reported that in a large number of the frogs which he collected in douglas, riley, pottawatomie, and geary counties, kansas, the digestive tracts and feces contained only ants. wood (1948: 226) reported an individual of _g. carolinensis_ in tennessee found under a flat rock in the center of an ant nest. freiburg (_op. cit._: 383) reported on the stomach contents of 52 ant-eating frogs collected near the reservation. ants constituted nearly all these stomach contents, though remains of a few small beetles were found. the ants eaten were of two kinds, _lasius interjectus_ and _crematogaster_ sp. the latter was by far the more numerous. although i made no further study of stomach contents, the myrmecophagous habits of _gastrophryne_ have come to my attention frequently in the course of routine field work. individuals kept in confinement for a day or more almost invariably voided feces which consisted mainly or entirely of ant remains, chiefly the heads, as these are most resistant to digestion. often upon examining frogs i have found ants (_crematogaster_ sp.) or their severed heads, attached with mandibles embedded in the skin. to have been attacked by ants, the frogs must have been in or beside the ants' burrow systems. frequently the frogs that were uncovered beneath rocks were adjacent to clusters of ants or to their nests or travelways, in a position strategically located to feed upon them, as described by tanner. often the feces of the frogs were found in pitfalls or under flat rocks. although these feces were not analyzed, they seemed to consist mainly or entirely of ant remains. the species of _crematogaster_, which is the chief food of _gastrophryne_ in this region, is largely subterranean in habits, and is extremely abundant. any flat rock in damp soil is likely to harbor a colony beneath it. colonies are situated also in damp soil away from rocks, beneath almost any kind of debris, and in hollow weed stalks and decaying wood. live-traps for small mammals, having nest boxes attached, almost always were occupied by colonies of _crematogaster_, if they were left in the field in warm, humid weather. occasionally the ants attacked and killed small mammals caught in such traps. among the thousands of kinds of insects occurring on the reservation, this ant is one of the most numerous in individuals, one of the most important on the basis of biomass and provides an abundant food source for those predators that are ant eaters. food supply probably is not a limiting factor to populations of _gastrophryne_ on the area. predation young copperheads are known to feed upon ant-eating frogs occasionally (anderson, 1942: 216; freiburg, 1951: 378). other kinds of snakes supposedly eat them also. the common water snake (_natrix sipedon_) and garter snake (_thamnophis sirtalis_) probably take heavy toll of the adults at the time they are concentrated at the breeding pools. larger salientians may be among the more important enemies of the breeding adults, the tadpoles, and the newly metamorphosed young. bullfrogs (_rana catesbeiana_) and leopard frogs (_rana pipiens_) are normally abundant at the pond on the reservation. these large voracious frogs lining the banks are quick to lunge at any moving object, and must take heavy toll of the much smaller ant-eating frogs that have to pass through their ranks to reach the water. the newly metamorphosed young often are forced to remain at a pond's edge for many days, or even for weeks, by drought and they must be subject to especially heavy predation by ranid frogs. even the smallest newly metamorphosed bullfrogs and leopard frogs would be large enough to catch and eat them. as a result of persistent drought conditions in 1952 and 1953, bullfrogs were completely eliminated from the pond by early 1954. re-invasion by a few individuals occurred in the course of the summer; these probably made long overland trips from ponds or streams that had persisted through the drought. leopard frogs reached the pond in somewhat larger numbers, but their population in 1954 was only a small percentage of that present in most other years. notable success in the ant-eating frog's reproduction in 1954 may have been due largely to the scarcity of these large ranids at the breeding ponds. freiburg (_loc. cit._) noted that many of the ant-eating frogs he examined were scarred, and some had digits or limbs amputated. he did not speculate concerning the origin of these injuries. however, it seems likely that many or all of them were inflicted by the short-tailed shrew (_blarina brevicauda_). five-lined skinks living on the same area were likewise found to be scarred by bites which i identified (fitch, 1954: 133) as bites of the short-tailed shrew. this shrew is common on the reservation, especially in woodland. many have been trapped in the pitfalls. on several occasions when a short-tailed shrew was caught in the same pitfall with ant-eating frogs, it was found to have killed and eaten them. like the frogs, the shrews were most often caught in pitfalls just after heavy rains. once in 1954 a shrew was found at the quarry in a pitfall that had been one of those most productive of frogs. the bottom of the pitfall was strewn with the discarded remains (mostly feet and skins) of perhaps a dozen ant-eating frogs. all had been eaten during one night and the following morning, as the trap had been checked on the preceding day. on other occasions shrews caught in pitfalls with several frogs had killed and eaten some and left others unharmed. summary in northeastern kansas the ant-eating frog, _gastrophryne olivacea_, is one of the more common species of amphibians. this area is near the northern limits of the species, genus, and family. the species prefers a dry, rocky upland habitat often in open woods or at woodland edge where other kinds of salientians do not ordinarily occur. it is, however, tolerant of a wide variety of habitat conditions, and may occur in river flood plains or cultivated land. in these situations where surface rocks are absent, cracks and rodent burrows presumably furnish the subterranean shelter that it requires. this frog is secretive and spends most of the time in subterranean shelter, obtaining its food there rather than in the open. only on warm rainy nights is it inclined to venture into the open. then, it moves about rapidly and with a scuttling gait, a combination of running and short hops. however, it may be flushed in daylight from a hiding place by the vibrations from footsteps of a person or an animal, or it may move about in the daytime when temperatures at night are too low for activity. though not swift of foot, the frogs are elusive because of their tendency to keep under cover, their slippery dermal secretion, and the ease with which they find and enter holes, or crevices to escape. breeding occurs at any time from late may through august and is controlled by the distribution of rainfall. heavy precipitation, especially rains of two inches or more, stimulates the frogs to migrate in large numbers to breeding ponds. even though there are several well spaced periods of unusually heavy rainfall in the course of a summer, each one initiates a new cycle of migration, mating and spawning. heavy rainfall is a necessity, not only to ensure a water supply in temporary pools where the frogs breed, but to create the moist conditions they require for an overland migration. an individual male may migrate to a pond and breed at least twice in the same season. whether or not the females do likewise is unknown. amplexus and spawning occur mainly within a day or two after the frogs reach the ponds. the males call chiefly at night, but there may be daytime choruses when breeding activity is at its peak. many males concentrate within a few square yards in the choruses and float upright usually beside or beneath a stem or leaf, or other shelter, rendering them extremely inconspicuous. the call is a bleat of three seconds duration, or a little more. in amplexus the members of a pair sometimes become glued together by their viscous dermal secretions. the eggs hatch in approximately 48 hours. the tadpoles metamorphose in as few as 24 days. newly metamorphosed frogs are 15 to 16 mm. in length, or, rarely as small as 14.5 mm. they are thus much larger than newly metamorphosed _g. carolinensis_, which have been described as 10-12 mm. or even as small as 8.5 mm. the newly metamorphosed frogs disperse from the breeding ponds as soon as there is a heavy rain. the young grow a little more than one mm. in length per week. those metamorphosed in early summer may attain minimum adult size before hibernation which begins in october. it seems that sexual maturity is most often attained in the second season, at an age of one to two years. _gastrophryne_ belongs to a family that is primarily tropical in distribution, and frogs of this genus have much higher temperature thresholds than most other amphibians of northeastern kansas, with a correspondingly short season of activity. for more than half the year, mid-october to early may the frogs are normally in hibernation. body temperatures of active frogs ranged from 17.0° c. to 37.6° c., but more than two-thirds were within the relatively narrow range, 24.0° to 31°. near the date of the first autumn frost the frogs disappear from the soil surface and from their usual shelters near the surface, presumably having retired into hibernation in deep holes and crevices. the natural enemies include young of the copperhead. the bullfrog and leopard frog probably take heavy toll of both the adults and the newly metamorphosed young at the breeding ponds. reproductive success of the ant-eating frogs was much greater in 1954 when these ranids were unusually scarce. the short-tailed shrew is an important enemy. on occasion it took heavy toll of frogs trapped in pitfalls, and many of the larger adults were scarred or mutilated from bites, probably of the shrew. each of several frogs was found consistently under the same rock for periods of weeks. the hundreds of other frogs that were marked were rarely found twice in any one spot. usually an individual recaptured after weeks or months was still near the original site. in many instances the distance involved was only a few yards, but there is some evidence that home ranges may be as long as 400 feet in greatest diameter. of those caught in two or more different years only 15 per cent were shown to have moved more than 400 feet. these few exceptionally long movements, up to 2000 feet, involve shifts in home range or migrations motivated by reproductive urge. literature cited anderson, p. 1942. amphibians and reptiles of jackson county, missouri. bull. chicago acad. sci., 6: 203-220. anderson, p. k. 1954. studies in the ecology of the narrow-mouthed toad, microhyla carolinensis carolinensis. tulane studies in zool., 2: 15-46. blair, a. p. 1950. note on oklahoma microhylid frogs. copeia, 1950: 152. bogert, c. m. 1949. thermoregulation in reptiles, a factor in evolution. evolution, 3: 195-211. bragg, a. n. 1943. observations on the ecology and natural history of anura, xv. the hylids and microhylids in oklahoma. great basin nat., 4: 62-80. de carvalho, a. l. 1954. a preliminary synopsis of the genera of american microhylid frogs. occas. papers mus. zool. univ. michigan, no. 555: 19 pp., 1 pl. dice, l. r. 1923. notes on the communities of vertebrates of riley county, kansas, with especial reference to the amphibians, reptiles and mammals. ecology, 4: 40-53. fitch, h. s. 1954. life history and ecology of the five-lined skink, eumeces fasciatus. univ. kansas publ. mus. nat. hist., 8: 1-156. freiburg, r. e. 1951. an ecological study of the narrow-mouthed toad (microhyla) in northeastern kansas. trans. kansas acad. sci., 54: 374-386. hecht, m. k., and matalas, b. l. 1946. a review of the middle american toads of the genus microhyla. american mus. novitates, no. 1315: 1-21. loomis, r. b. 1945. microhyla olivacea (hallowell) in nebraska. herpetologica, 2: 211-212. mittleman, m. b. 1950. miscellaneous notes on some amphibians and reptiles from the southeastern united states. herpetologica, 6: 20-24. parker, h. w. 1934. a monograph of the frogs of the family microhylidae. british mus. (nat. hist.) london, vii + 208 pp., figs. 1-67. pope, c. h. 1931. notes on amphibians from fukien, hainan, and other parts of china. bull. american mus. nat. hist., 61: 397-611. schmidt, k. p. 1953. a check list of north american amphibians and reptiles. univ. chicago press, viii + 280 pp. smith, h. m. 1934. the amphibians of kansas. american midland nat., 15: 377-528, pls. 12-20, maps 1-24. 1950. handbook of amphibians and reptiles of kansas. univ. kansas publ. mus. nat. hist. misc. publ., 2: 1-336 pp., 233 figs. smith, h. m., and taylor, e. h. 1950. type localities of mexican reptiles and amphibians. univ. kansas sci. bull. 33: 313-380. stebbins, r. c. 1951. amphibians of western north america. univ. california press, xviii + 539 pp. tanner, w. w. 1950. notes on the habits of microhyla carolinensis olivacea (hallowell). herpetologica, 6: 47-48. wood, j. t. 1948. microhyla c. carolinensis in an ant nest. herpetologica, 4: 226. wright, a. h. 1932. life-histories of the frogs of okefinokee swamp, georgia. macmillan co., new york, n. y. wright, a. h., and wright, a. a. 1949. handbook of frogs and toads of the united states and canada. comstock publ. co., ithaca, new york. _transmitted february 28, 1955._ * * * * * transcriber's notes a small number of inconsistencies and typographical errors have been changed in the text as follows: p. 279 "near-by" changed to "nearby" (in nearby counties of kansas) p. 289 "successivly" changed to "successively" (two successively older annual age classes) p. 297 "per cent" changed to "percent" (only 48 percent of specimens from the florida keys) p. 303 "famliy" changed to "family" (the northern limits of the species, genus, and family.) note: project gutenberg also has an html version of this file which includes the original illustrations. see 14375-h.htm or 14375-h.zip: (http://www.gutenberg.net/dirs/1/4/3/7/14375/14375-h/14375-h.htm) or (http://www.gutenberg.net/dirs/1/4/3/7/14375/14375-h.zip) the bedtime story-books the adventures of grandfather frog by thornton w. burgess author of _the adventures of reddy fox_, _old mother west wind_, etc. with illustrations by harrison cady boston little, brown, and company 1920 [illustration: "have a nice nap?" inquired jerry, with a broad grin. (frontispiece)] contents chapter i. billy mink finds little joe otter ii. longlegs the blue heron receives callers iii. longlegs visits the smiling pool iv. the patience of longlegs the blue heron v. grandfather frog jumps just in time vi. longlegs and whitetail quarrel vii. grandfather frog's big mouth gets him in trouble viii. spotty the turtle plays doctor ix. old mr. toad visits grandfather frog x. grandfather frog starts out to see the great world xi. grandfather frog is stubborn xii. grandfather frog keeps on xiii. danny meadow mouse feels responsible xiv. grandfather frog has a strange ride xv. grandfather frog gives up hope xvi. the merry little breezes work hard xvii. striped chipmunk cuts the string xviii. grandfather frog hurries away xix. grandfather frog jumps into more trouble xx. grandfather frog loses heart xxi. the merry little breezes try to comfort grandfather frog xxii. grandfather frog's troubles grow xxiii. the dear old smiling pool once more list of illustrations "have a nice nap?" inquired jerry, with a broad grin "thank you," said longlegs. "i believe i have an errand up that way" as soon as they saw grandfather frog, they began to laugh, too "you won't see much of the great world if you jump like that every time you get a scare," said danny he seized the other end of the string and began to pull "that's just what i'm afraid of!" croaked grandfather frog the adventures of grandfather frog i billy mink finds little joe otter billy mink ran around the edge of the smiling pool and turned down by the laughing brook. his eyes twinkled with mischief, and he hurried as only billy can. as he passed jerry muskrat's house, jerry saw him. "hi, billy mink! where are you going in such a hurry this fine morning?" he called. "to find little joe otter. have you seen anything of him?" replied billy. "no," said jerry. "he's probably down to the big river fishing. i heard him say last night that he was going." "thanks," said billy mink, and without waiting to say more he was off like a little brown flash. jerry watched him out of sight. "hump!" exclaimed jerry. "billy mink is in a terrible hurry this morning. now i wonder what he is so anxious to find little joe otter for. when they get their heads together, it is usually for some mischief." jerry climbed to the top of his house and looked over the smiling pool in the direction from which billy mink had just come. almost at once he saw grandfather frog fast asleep on his big green lily-pad. the legs of a foolish green fly were sticking out of one corner of his big mouth. jerry couldn't help laughing, for grandfather frog certainly did look funny. "he's had a good breakfast this morning, and his full stomach has made him sleepy," thought jerry. "but he's getting careless in his old age. he certainly is getting careless. the idea of going to sleep right out in plain sight like that!" suddenly a new thought popped into his head. "billy mink saw him, and that is why he is so anxious to find little joe otter. he is planning to play some trick on grandfather frog as sure as pollywogs have tails!" exclaimed jerry. then his eyes began to twinkle as he added: "i think i'll have some fun myself." without another word jerry slipped down into the water and swam over to the big green lily-pad of grandfather frog. then he hit the water a smart blow with his tail. grandfather frog's big goggly eyes flew open, and he was just about to make a frightened plunge into the smiling pool when he saw jerry. "have a nice nap?" inquired jerry, with a broad grin. "i wasn't asleep!" protested grandfather frog indignantly. "i was just thinking." "don't you think it a rather dangerous plan to think so long with your eyes closed?" asked jerry. "well, maybe i did just doze off," admitted grandfather frog sheepishly. "maybe you did," replied jerry. "now listen." then jerry whispered in grandfather frog's ear, and both chuckled as if they were enjoying some joke, for they are great friends, you know. afterward jerry swam back to his house, and grandfather frog closed his eyes so as to look just as he did when he was asleep. meanwhile billy mink had hurried down the laughing brook. half-way to the big river he met little joe otter bringing home a big fish, for you know little joe is a great fisherman. billy mink hastened to tell him how grandfather frog had fallen fast asleep on his big green lily-pad. "it's a splendid chance to have some fun with grandfather frog and give him a great scare," concluded billy. little joe otter put his fish down and grinned. he likes to play pranks almost as well as he likes to go fishing. "what can we do?" said he. "i've thought of a plan," replied billy. "do you happen to know where we can find longlegs the blue heron?" "yes," said little joe. "i saw him fishing not five minutes ago." then billy told little joe his plan, and laughing and giggling, the two little scamps hurried off to find longlegs the blue heron. ii longlegs the blue heron receives callers longlegs the blue heron felt decidedly out of sorts. it was a beautiful morning, too beautiful for any one to be feeling that way. indeed, it was the same beautiful morning in which grandfather frog had caught so many foolish green flies. jolly, round, bright mr. sun was smiling his broadest. the merry little breezes of old mother west wind were dancing happily here and there over the green meadows, looking for some good turn to do for others. the little feathered people to whom old mother nature has given the great blessing of music in their throats were pouring out their sweetest songs. so it seemed as if there was no good reason why longlegs should feel out of sorts. the fact is the trouble with longlegs was an empty stomach. yes, sir, that is what ailed longlegs the blue heron that sunshiny morning. you know it is hard work to be hungry and happy at the same time. so longlegs stood on the edge of a shallow little pool in the laughing brook, grumbling to himself. just a little while before, he had seen little joe otter carrying home a big fish, and this had made him hungrier and more out of sorts than ever. in the first place it made him envious, and envy, you know, always stirs up bad feelings. he knew perfectly well that little joe had got that fish by boldly chasing it until he caught it, for little joe can swim even faster than a fish. but longlegs chose to try to make himself think that it was all luck. moreover, he wanted to blame some one for his own lack of success, as most people who fail do. so when little joe had called out: "hi, longlegs, what luck this fine morning?" longlegs just pretended not to hear. but when little joe was out of sight and hearing, he began to grumble to himself. "no wonder i have no luck with that fellow racing up and down the laughing brook," said he. "he isn't content to catch what he wants himself, but frightens the rest of the fish so that an honest fisherman like me has no chance at all. i don't see what old mother nature was thinking of when she gave him a liking for fish. he and billy mink are just two worthless little scamps, born to make trouble for other people." he was still grumbling when these two same little scamps poked their heads out of the grass on the other side of the little pool. "you look happy, longlegs. must be that you have had a good breakfast," said little joe, nudging billy mink. longlegs snapped his great bill angrily. "what are you doing here, spoiling my fishing?" he demanded. "haven't you got the big river and all the rest of the laughing brook to fool around in? this is my pool, and i'll thank you to keep away!" billy mink chuckled so that longlegs heard him, and that didn't improve his temper a bit. but before he could say anything more, little joe otter spoke. "oh," said he, "we beg your pardon. we just happen to know that grandfather frog is sound asleep, and we thought that if you hadn't had good luck this morning, you might like to know about it. as long as you think so ill of us, we'll just run over and tell blackcap the night heron." little joe turned as if to start off in search of blackcap at once. "hold on a minute!" called longlegs, and tried to make his voice sound pleasant, a difficult thing to do, because, you know, his voice is very harsh and disagreeable. "the truth is, i haven't had a mouthful of breakfast and to be hungry is apt to make me cross. where did you say grandfather frog is?" "i didn't say," replied little joe, "but if you really want to know, he is sitting on his big green lily-pad in the smiling pool fast asleep right in plain sight." "thank you," said longlegs. "i believe i have an errand up that way, now i think of it. i believe i'll just go over and have a look at him. i have never seen him asleep." [illustration: "thank you," said longlegs. "i believe i have an errand up that way." _page 10_.] iii longlegs visits the smiling pool longlegs the blue heron watched billy mink and little joe otter disappear down the laughing brook. as long as they were in sight, he sat without moving, his head drawn down between his shoulders just as if he had nothing more important to think about than a morning nap. but if you had been near enough to have seen his keen eyes, you would never have suspected him of even thinking of a nap. just as soon as he felt sure that the two little brown-coated scamps were out of sight, he stretched his long neck up until he was almost twice as tall as he had been a minute before. he looked this way and that way to make sure that no danger was near, spread his great wings, flapped heavily up into the air, and then, with his head once more tucked back between his shoulders and his long legs straight out behind him, he flew out over the green meadows, and making a big circle, headed straight for the smiling pool. all this time billy mink and little joe otter had not been so far away as longlegs supposed. they had been hiding where they could watch him, and the instant he spread his wings, they started back up the laughing brook towards the smiling pool to see what would happen there. you see they knew perfectly well that longlegs was flying up to the smiling pool in the hope that he could catch grandfather frog for his breakfast. they didn't really mean that any harm should come to grandfather frog, but they meant that he should have a great fright. you see, they were like a great many other people, so heedless and thoughtless that they thought it fun to frighten others. "of course we'll waken grandfather frog in time for him to get away with nothing more than a great scare," said little joe otter, as they hurried along. "it will be such fun to see his big goggly eyes pop out when he opens them and sees longlegs just ready to gobble him up! and won't longlegs be hopping mad when we cheat him out of the breakfast he is so sure he is going to have!" they reached the smiling pool before longlegs, who had taken a roundabout way, and they hid among the bulrushes where they could see and not be seen. "there's the old fellow just as i left him, fast asleep," whispered billy mink. sure enough, there on his big green lily-pad sat grandfather frog with his eyes shut. at least, they seemed to be shut. and over on top of his big house sat jerry muskrat. jerry seemed to be too busy opening a fresh-water clam to notice anything else; but the truth is he was watching all that was going on. you see, he had suspected that billy mink was going to play some trick on grandfather frog, so he had warned him. when he had seen longlegs coming towards the smiling pool, he had given grandfather frog another warning, and he knew that now he was only pretending to be asleep. straight up to the smiling pool came longlegs the blue heron, and on the very edge of it, among the bulrushes, he dropped his long legs and stood with his toes in the water, his long neck stretched up so that he could look all over the smiling pool. there, just as little joe otter had said, sat grandfather frog on his big green lily-pad, fast asleep. at least, he seemed to be fast asleep. the eyes of longlegs sparkled with hunger and the thought of what a splendid breakfast grandfather frog would make. very slowly, putting each foot down as carefully as he knew how, longlegs began to walk along the shore so as to get opposite the big green lily-pad where grandfather frog was sitting. and over in the bulrushes on the other side, little joe otter and billy mink nudged each other and clapped their hands over their mouths to keep from laughing aloud. iv the patience of longlegs the blue heron patience often wins the day when over-haste has lost the way. if there is one virtue which longlegs the heron possesses above another it is patience. yes, sir, longlegs certainly has got patience. he believes that if a thing is worth having, it is worth waiting for, and that if he waits long enough, he is sure to get it. perhaps that is because he has been a fisherman all his life, and his father and his grandfather were fishermen. you know a fisherman without patience rarely catches anything. of course billy mink and little joe otter laugh at this and say that it isn't so, but the truth is they sometimes go hungry when they wouldn't if they had a little of the patience of longlegs. now grandfather frog is another who is very, very patient. he can sit still the longest time waiting for something to come to him. indeed, he can sit perfectly still so long, and longlegs can stand perfectly still so long, that jerry muskrat and billy mink and little joe otter have had many long disputes as to which of the two can keep still the longest. "he will make a splendid breakfast," thought longlegs, as very, very carefully he walked along the edge of the smiling pool so as to get right opposite grandfather frog. there he stopped and looked very hard at grandfather frog. yes, he certainly must be asleep, for his eyes were closed. longlegs chuckled to himself right down inside without making a sound, and got ready to wade out so as to get within reach. now all the time grandfather frog was doing some quiet chuckling himself. you see, he wasn't asleep at all. he was just pretending to be asleep, and all the time he was watching longlegs out of a corner of one of his big goggly eyes. very, very slowly and carefully, so as not to make the teeniest, weeniest sound, longlegs lifted one foot to wade out into the smiling pool. grandfather frog pretended to yawn and opened his big goggly eyes. longlegs stood on one foot without moving so much as a feather. grandfather frog yawned again, nodded as if he were too sleepy to keep awake, and half closed his eyes. longlegs waited and waited. then, little by little, so slowly that if you had been there you would hardly have seen him move, he drew his long neck down until his head rested on his shoulders. "i guess i must wait until he falls sound asleep again," said longlegs to himself. but grandfather frog didn't go to sleep. he would nod and nod and then, just when longlegs would make up his mind that this time he really was asleep, open would pop grandfather frog's eyes. so all the long morning longlegs stood on one foot without moving, watching and waiting and growing hungrier and hungrier, and all the long morning grandfather frog sat on his big green lily-pad, pretending that he was oh, so sleepy, and all the time having such a comfortable sun-bath and rest, for very early he had had a good breakfast of foolish green flies. over in the bulrushes on the other side of the smiling pool two little scamps in brown bathing suits waited and watched for the great fright they had planned for grandfather frog, when they had sent longlegs to try to catch him. they were billy mink and little joe otter. at first they laughed to themselves and nudged each other at the thought of the trick they had played. then, as nothing happened, they began to grow tired and uneasy. you see they do not possess patience. finally they gave up in disgust and stole away to find some more exciting sport. grandfather frog saw them go and chuckled harder than ever to himself. v grandfather frog jumps just in time back and forth over the green meadows sailed whitetail the marsh hawk. like longlegs the blue heron, he was hungry. his sharp eyes peered down among the grasses, looking for something to eat, but some good fairy seemed to have warned the very little people who live there that whitetail was out hunting. perhaps it was one of old mother west wind's children, the merry little breezes. you know they are always flitting about trying to do some one a good turn. they love to dance and romp and play from dawn to dusk the livelong day, but more than this they love to find a chance to do some favor kind. anyway, little mr. green snake seemed to know that whitetail was out hunting and managed to keep out of sight. danny meadow mouse wasn't to be found. only a few foolish grasshoppers rewarded his patient search, and these only served to make him feel hungrier than ever. but old whitetail has a great deal of persistence, and in spite of his bad luck, he kept at his hunting, back and forth, back and forth, until he had been all over the green meadows. at last he made up his mind that he was wasting time there. "i'll just have a look over at the smiling pool, and if there is nothing there, i'll take a turn or two along the big river," thought he and straightway started for the smiling pool. long before he reached it, his keen eyes saw longlegs the blue heron standing motionless on the edge of it, and he knew by the looks of longlegs that he was watching something which he hoped to catch. "if it's a fish," thought whitetail, "it will do me no good, for i am no fisherman. but if it's a frog--well, frogs are not as good eating as fat meadow mice, but they are very filling." with that he hurried a little faster, and then he saw what longlegs was watching so intently. it was, as you know, grandfather frog sitting on his big green lily-pad. old whitetail gave a great sigh of satisfaction. grandfather frog certainly would be very filling, very filling, indeed. now longlegs the blue heron was so intently watching grandfather frog that he saw nothing else, and grandfather frog was so busy watching longlegs that he quite forgot that there might be other dangers. besides, his back was toward old whitetail. of course whitetail saw this, and it made him almost chuckle aloud. ever so many times he had tried to catch grandfather frog, but always grandfather frog had seen him long before he could get near him. now, with all his keen sight, old whitetail had failed to see some one else who was sitting right in plain sight. he had failed because his mind was so full of grandfather frog and longlegs that he forgot to look around, as he usually does. just skimming the tops of the bulrushes he sailed swiftly out over the smiling pool and reached down with his great, cruel claws to clutch grandfather frog, who sat there pretending to be asleep, but all the time watching longlegs and deep down inside chuckling to think how he was fooling longlegs. slap! that was the tail of jerry muskrat hitting the water. grandfather frog knew what that meant--danger! he didn't know what the danger was, and he didn't wait to find out. there would be time enough for that later. when jerry muskrat slapped the water with his tail that way, danger was very near indeed. with a frightened "chugarum!" grandfather frog dived head first into the smiling pool, and so close was old whitetail that the water was splashed right in his face. he clutched frantically with his great claws, but all he got was a piece of the big green lily-pad on which grandfather frog had been sitting, and of course this was of no use for an empty stomach. with a scream of disappointment and anger, he whirled in the air and made straight for jerry muskrat. but jerry just laughed in the most provoking way and ducked under water. vi longlegs and whitetail quarrel "you did!" "i didn't! i didn't!" "you did!" such a terrible fuss when grandfather hid! you see longlegs the blue heron had stood very patiently on one foot all the long morning waiting for grandfather frog to go to sleep on his big green lily-pad. he had felt sure he was to have grandfather frog for his breakfast and lunch, for he had had no breakfast, and it was now lunch time. he was so hungry that it seemed to him that the sides of his stomach certainly would fall in because there was nothing to hold them up, and then, without any warning at all, old whitetail the marsh hawk had glided out across the smiling pool with his great claws stretched out to clutch grandfather frog, and grandfather frog had dived into the smiling pool with a great splash just in the very nick of time. now is there anything in the world so hard on the temper as to lose a good meal when you are very, very, very hungry? of course longlegs didn't really have that good meal, but he had thought that he was surely going to have it. so when grandfather frog splashed into the smiling pool, of course longlegs lost his temper altogether. his yellow eyes seemed to grow even more yellow. "you robber! you thief!" he screamed harshly at old whitetail. now old whitetail was just as hungry as longlegs, and he had come even nearer to catching grandfather frog. he is even quicker tempered than longlegs. he had whirled like a flash on jerry muskrat, but jerry had just laughed in the most provoking manner and ducked under water. this had made old whitetail angrier than ever, and then to be called bad names--robber and thief! it was more than any self-respecting hawk could stand. yes, sir, it certainly was! he fairly shook with rage as he turned in the air once more and made straight for longlegs the blue heron. "i'm no more robber and thief than you are!" he shrieked. "you frightened away my frog!" screamed longlegs. "i didn't!" "you did!" "i didn't! it wasn't your frog; it was mine!" "chugarum!" said grandfather frog to jerry muskrat, as they peeped out from under some lily-pads. "i didn't know i belonged to anybody. i really didn't. did you?" "no," replied jerry, his eyes sparkling with excitement as he watched longlegs and whitetail, "it's news to me." "you're too lazy to hunt like honest people!" taunted old whitetail, as he wheeled around longlegs, watching for a chance to strike with his great, cruel claws. "i'm too honest to take the food out of other people's mouths!" retorted longlegs, dancing around so as always to face whitetail, one of his great, broad wings held in front of him like a shield, and his long, strong bill ready to strike. every feather on whitetail's head was standing erect with rage, and he looked very fierce and terrible. at last he saw a chance, or thought he did, and shot down. but all he got was a feather from that great wing which longlegs kept in front of him, and before he could get away, that long bill had struck him twice, so that he screamed with pain. so they fought and fought, till the ground was covered with feathers, and they were too tired to fight any longer. then, slowly and painfully, old whitetail flew away over the green meadows, and with torn and ragged wings, longlegs flew heavily down the laughing brook towards the big river, and both were sore and stiff and still hungry. "dear me! dear me! what a terrible thing and how useless anger is," said grandfather frog, as he climbed back on his big green lily-pad in the warm sunshine. vii grandfather frog's big mouth gets him in trouble grandfather frog has a great big mouth. you know that. everybody does. his friends of the smiling pool, the laughing brook, and the green meadows have teased grandfather frog a great deal about the size of his mouth, but he hasn't minded in the least, not the very least. you see, he learned a long time ago that a big mouth is very handy for catching foolish green flies, especially when two happen to come along together. so he is rather proud of his big mouth, just as he is of his goggly eyes. but once in a while his big mouth gets him into trouble. it's a way big mouths have. it holds so much that it makes him greedy sometimes. he stuffs it full after his stomach already has all that it can hold, and then of course he can't swallow. then grandfather frog looks very foolish and silly and undignified, and everybody calls him a greedy fellow who is old enough to know better and who ought to be ashamed of himself. perhaps he is, but he never says so, and he is almost sure to do the same thing over again the first chance he has. now it happened that one morning when grandfather frog had had a very good breakfast of foolish green flies and really didn't need another single thing to eat, who should come along but little joe otter, who had been down to the big river fishing. he had eaten all he could hold, and he was taking the rest of his catch to a secret hiding-place up the laughing brook. now grandfather frog is very fond of fish for a change, and when he saw those that little joe otter had, his eyes glistened, and in spite of his full stomach his mouth watered. "good morning, grandfather frog! have you had your breakfast yet?" called little joe otter. grandfather frog wanted to say no, but he always tells the truth. "ye-e-s," he replied. "i've had my breakfast, such as it was. why do you ask?" "oh, for no reason in particular. i just thought that if you hadn't, you might like a fish. but as long as you have breakfasted, of course you don't want one," said little joe, his bright eyes beginning to twinkle. he held the fish out so that grandfather frog could see just how plump and nice they were. "chugarum!" exclaimed grandfather frog. "those certainly are very nice fish, very nice fish indeed. it is very nice of you to think of a poor old fellow like me, and i--er--well, i might find room for just a little teeny, weeny one, if you can spare it." little joe otter knows all about grandfather frog's greediness. he looked at grandfather frog's white and yellow waistcoat and saw how it was already stuffed full to bursting. the twinkle in his eyes grew more mischievous than ever as he said: "of course i can. but i wouldn't think of giving such an old friend a teeny, weeny one." with that, little joe picked out the biggest fish he had and tossed it over to grandfather frog. it landed close by his nose with a great splash, and it was almost half as big as grandfather frog himself. it was plump and looked so tempting that grandfather frog forgot all about his full stomach. he even forgot to be polite and thank little joe otter. he just opened his great mouth and seized the fish. yes, sir, that is just what he did. almost before you could wink an eye, the fish had started down grandfather frog's throat head first. now you know grandfather frog has no teeth, and so he cannot bite things in two. he has to swallow them whole. that is just what he started to do with the fish. it went all right until the head reached his stomach. but you can't put anything more into a thing already full, and grandfather frog's stomach was packed as full as it could be of foolish green flies. there the fish stuck, and gulp and swallow as hard as he could, grandfather frog couldn't make that fish go a bit farther. then he tried to get it out again, but it had gone so far down his throat that he couldn't get it back. grandfather frog began to choke. viii spotty the turtle plays doctor greed's a dreadful thing to see, as everybody will agree. at first little joe otter, sitting on the bank of the smiling pool, laughed himself almost sick as he watched grandfather frog trying to swallow a fish almost as big as himself, when his white and yellow waistcoat was already stuffed so full of foolish green flies that there wasn't room for anything more. such greed would have been disgusting, if it hadn't been so very, very funny. at least, it was funny at first, for the fish had stuck, with the tail hanging out of grandfather frog's big mouth. grandfather frog hitched this way and hitched that way on his big green lily-pad, trying his best to swallow. twice he tumbled off with a splash into the smiling pool. each time he scrambled back again and rolled his great goggly eyes in silent appeal to little joe otter to come to his aid. [illustration: as soon as they saw grandfather frog, they began to laugh, too. _page 37._] but little joe was laughing so that he had to hold his sides, and he didn't understand that grandfather frog really was in trouble. billy mink and jerry muskrat came along, and as soon as they saw grandfather frog, they began to laugh, too. they just laughed and laughed and laughed until the tears came. they rolled over and over on the bank and kicked their heels from sheer enjoyment. it was the funniest thing they had seen for a long, long time. "did you ever see such greed?" gasped billy mink. "why don't you pull it out and start over again?" shouted little joe otter. now this is just what grandfather frog was trying to do. at least, he was trying to pull the fish out. he hadn't the least desire in the world to try swallowing it again. in fact, he felt just then as if he never, never wanted to see another fish so long as he lived. but grandfather frog's hands are not made for grasping slippery things, and the tail of a fish is very slippery indeed. he tried first with one hand, then with the other, and at last with both. it was of no use at all. he just couldn't budge that fish. he couldn't cough it up, because it had gone too far down for that. the more he clawed at that waving tail with his hands, the funnier he looked, and the harder little joe otter and billy mink and jerry muskrat laughed. they made such a noise that spotty the turtle, who had been taking a sun-bath on the end of an old log, slipped into the water and started to see what it was all about. now spotty the turtle is very, very slow on land, but he is a good swimmer. he hurried now because he didn't want to miss the fun. at first he didn't see grandfather frog. "what's the joke?" he asked. little joe otter simply pointed to grandfather frog. little joe had laughed so much that he couldn't even speak. spotty looked over to the big green lily-pad and started to laugh too. then he saw great tears rolling down from grandfather frog's eyes and heard little choky sounds. he stopped laughing and started for grandfather frog as fast as he could swim. he climbed right up on the big green lily-pad, and reaching out, grabbed the end of the fish tail in his beak-like mouth. then spotty the turtle settled back and pulled, and grandfather frog settled back and pulled. splash! grandfather frog had fallen backward into the smiling pool on one side of the big green lily-pad. splash! spotty the turtle had fallen backward into the smiling pool on the opposite side of the big green lily-pad. and the fish which had caused all the trouble lay floating on the water. "thank you! thank you!" gasped grandfather frog, as he feebly crawled back on the lily-pad. "a minute more, and i would have choked to death." "don't mention it," replied spotty the turtle. "i never, never will," promised grandfather frog. ix old mr. toad visits grandfather frog grandfather frog and old mr. toad are cousins. of course you know that without being told. everybody does. but not everybody knows that they were born in the same place. they were. yes, sir, they were. they were born in the smiling pool. both had long tails and for a while no legs, and they played and swam together without ever going on shore. in fact, when they were babies, they couldn't live out of the water. and people who saw them didn't know the difference between them and called them by the same names--tadpoles or pollywogs. but when they grew old enough to have legs and get along without tails, they parted company. you see, it was this way: grandfather frog (of course he wasn't grandfather then) loved the smiling pool so well that he couldn't think of leaving it. he heard all about the great world and what a wonderful place it was, but he couldn't and wouldn't believe that there could be any nicer place than the smiling pool, and so he made up his mind that he would live there always. but mr. toad could hardly wait to get rid of his tail before turning his back on the smiling pool and starting out to see the great world. nothing that grandfather frog could say would stop him, and away mr. toad went, when he was so small that he could hide under a clover leaf. grandfather frog didn't expect ever to see him again. but he did, though it wasn't for a long, long time. and when he did come back, he had grown so that grandfather frog hardly knew him at first. and right then and there began a dispute which they have kept up ever since: whether it was best to go out into the great world or remain in the home of childhood. each was sure that what he had done was best, and each is sure of it to this day. so whenever old mr. toad visits grandfather frog, as he does every once in a while, they are sure to argue and argue on this same old subject. it was so on the day that grandfather frog had so nearly choked to death. old mr. toad had heard about it from one of the merry little breezes of old mother west wind and right away had started for the smiling pool to pay his respects to grandfather frog, and to tell him how glad he was that spotty the turtle had come along just in time to pull the fish out of grandfather frog's throat. now all day long grandfather frog had had to listen to unpleasant remarks about his greediness. it was such a splendid chance to tease him that everybody around the smiling pool took advantage of it. grandfather frog took it good-naturedly at first, but after a while it made him cross, and by the time his cousin, old mr. toad, arrived, he was sulky and just grunted when mr. toad told him how glad he was to find grandfather frog quite recovered. old mr. toad pretended not to notice how out of sorts grandfather frog was but kept right on talking. "if you had been out in the great world as much as i have been, you would have known that little joe otter wasn't giving you that fish for nothing," said he. grandfather frog swelled right out with anger. "chugarum!" he exclaimed in his deepest, gruffest voice. "chugarum! go back to your great world and learn to mind your own affairs, mr. toad." right away old mr. toad began to swell with anger too. for a whole minute he glared at grandfather frog, so indignant he couldn't find his tongue. when he did find it, he said some very unpleasant things, and right away they began to dispute. "what good are you to anybody but yourself, never seeing anything of the great world and not knowing anything about what is going on or what other people are doing?" asked old mr. toad. "i'm minding my own affairs and not meddling with things that don't concern me, as seems to be the way out in the great world you are so fond of talking about," retorted grandfather frog. "wise people know enough to be content with what they have. you've been out in the great world ever since you could hop, and what good has it done you? tell me that! you haven't even a decent suit of clothes to your back." grandfather frog patted his white and yellow waistcoat as he spoke and looked admiringly at the reflection of his handsome green coat in the smiling pool. old mr. toad's eyes snapped, for you know his suit is very plain and rough. "people who do honest work for their living have no time to sit about in fine clothes admiring themselves," he replied sharply. "i've learned this much out in the great world, that lazy people come to no good end, and i know enough not to choke myself to death." grandfather frog almost choked again, he was so angry. you see old mr. toad's remarks were very personal, and nobody likes personal remarks when they are unpleasant, especially if they happen to be true. grandfather frog was trying his best to think of something sharp to say in reply, when mr. redwing, sitting in the top of the big hickory-tree, shouted: "here comes farmer brown's boy!" grandfather frog forgot his anger and began to look anxious. he moved about uneasily on his big green lily-pad and got ready to dive into the smiling pool, for he was afraid that farmer brown's boy had a pocketful of stones as he usually did have when he came over to the smiling pool. old mr. toad didn't look troubled the least bit. he didn't even look around for a hiding-place. he just sat still and grinned. "you'd better watch out, or you'll never visit the smiling pool again," called grandfather frog. "oh," replied old mr. toad, "i'm not afraid. farmer brown's boy is a friend of mine. i help him in his garden. how to make friends is one of the things the great world has taught me." "chugarum!" said grandfather frog. "i'd have you to know that--" but what it was that he was to know old mr. toad never found out, for just then grandfather frog caught sight of farmer brown's boy and without waiting even to say good-by he dived into the smiling pool. x grandfather frog starts out to see the great world grandfather frog looked very solemn as he sat on his big green lily-pad in the smiling pool. he looked very much as if he had something on his mind. a foolish green fly actually brushed grandfather frog's nose and he didn't even notice it. the fact is he did have something on his mind. it had been there ever since his cousin, old mr. toad, had called the day before and they had quarreled as usual over the question whether it was best never to leave home or to go out into the great world. right in the midst of their quarrel along had come farmer brown's boy. now grandfather frog is afraid of farmer brown's boy, so when he appeared, grandfather frog stopped arguing with old mr. toad and with a great splash dived into the smiling pool and hid under a lily-pad. there he stayed and watched his cousin, old mr. toad, grinning in the most provoking way, for he wasn't afraid of farmer brown's boy. in fact, he had boasted that they were friends. grandfather frog had thought that this was just an idle boast, but when he saw farmer brown's boy tickle old mr. toad under his chin with a straw, while mr. toad sat perfectly still and seemed to enjoy it, he knew that it was true. grandfather frog had not come out of his hiding-place until after old mr. toad had gone back across the green meadows and farmer brown's boy had gone home for his supper. then grandfather frog had climbed back on his big green lily-pad and had sat there half the night without once leading the chorus of the smiling pool with his great deep bass voice as he usually did. he was thinking, thinking very hard. and now, this bright, sunshiny morning, he was still thinking. the fact is grandfather frog was beginning to wonder if perhaps, after all, mr. toad was right. if the great world had taught him how to make friends with farmer brown's boy, there really must be some things worth learning there. not for the world would grandfather frog have admitted to old mr. toad or to any one else that there was anything for him to learn, for you know he is very old and by his friends is accounted very wise. but right down in his heart he was beginning to think that perhaps there were some things which he couldn't learn in the smiling pool. so he sat and thought and thought. suddenly he made up his mind. "chugarum!" said he. "i'll do it!" "do what?" asked jerry muskrat, who happened to be swimming past. "i'll go out and see for myself what this great world my cousin, old mr. toad, is so fond of talking about is like," replied grandfather frog. "don't you do it," advised jerry muskrat. "don't you do anything so foolish as that. you're too old, much too old, grandfather frog, to go out into the great world." now few old people like to be told that they are too old to do what they please, and grandfather frog is no different from others. "you just mind your own affairs, jerry muskrat," he retorted sharply. "i guess i know what is best for me without being told. if my cousin, old mr. toad, can take care of himself out in the great world, i can. he isn't half so spry as i am. i'm going, and that is all there is about it!" with that grandfather frog dived into the smiling pool, swam across to a place where the bank was low, and without once looking back started across the green meadows to see the great world. xi grandfather frog is stubborn "fee, fi, fe, fum! chug, chug, chugarum!" grandfather actually had started out to see the great world. yes, sir, he had turned his back on the smiling pool, and nothing that jerry muskrat could say made the least bit of difference. grandfather frog had made up his mind, and when he does that, it is just a waste of time and breath for any one to try to make him change it. you see grandfather frog is stubborn. yes, that is just the word--stubborn. he would see for himself what this great world was that his cousin, old mr. toad, talked so much about and said was so much better than the smiling pool where grandfather frog had spent his whole life. "if old mr. toad can take care of himself, i can take care of myself out in the great world," said grandfather frog, to himself as, with great jumps, he started out on to the green meadows. "i guess he isn't any smarter than i am! he isn't half so spry as i am, and i can jump three times as far as he can. i'll see for myself what this great world is like, and then i'll go back to the smiling pool and stay there the rest of my life. chugarum, how warm it is!" it was warm. jolly, round, bright mr. sun was smiling his broadest and pouring his warmest rays down on the green meadows. the merry little breezes of old mother west wind were taking a nap. you see, they had played so hard early in the morning that they were tired. so there was nobody and nothing to cool grandfather frog, and he just grew warmer and warmer with every jump. he began to grow thirsty, and how he did long for a plunge in the dear, cool smiling pool! but he was stubborn. he wouldn't turn back, no matter how uncomfortable he felt. he _would_ see the great world if it killed him. so he kept right on, jump, jump, jump, jump. grandfather frog had been up the laughing brook and down the laughing brook, where he could swim when he grew tired of traveling on the bank, and where he could cool off whenever he became too warm, but never before had he been very far away from water, and he found this a very different matter. at first he had made great jumps, for that is what his long legs were given him for; but the long grass bothered him, and after a little the jumps grew shorter and shorter and shorter, and with every jump he puffed and puffed and presently began to grunt. you see he never before had made more than a few jumps at a time without resting, and his legs grew tired in a very little while. now if grandfather frog had known as much about the green meadows as the little people who live there all the time do, he would have taken the lone little path, where the going was easy. but he didn't. he just started right out without knowing where he was going, and of course the way was hard, very hard indeed. the grass was so tall that he couldn't see over it, and the ground was so rough that it hurt his tender feet, which were used to the soft, mossy bank of the smiling pool. he had gone only a little way before he wished with all his might that he had never thought of seeing the great world. but he had said that he was going to and he would, so he kept right on--jump, jump, rest, jump, jump, jump, rest, jump, and then a long rest. it was during one of these rests that he heard footsteps, and then a dreadful sound that made cold chills run all over him. sniff, sniff, sniff! it was coming nearer. grandfather frog flattened himself down as close to the ground as he could get. but it was of no use, no use at all. the sniffing came nearer and nearer, and then right over him stood bowser the hound! bowser looked just as surprised as he felt. he put out one paw and turned grandfather frog over on his back. grandfather frog struggled to his feet and made two frightened jumps. "bow, wow!" cried bowser and rolled him over again. bowser thought it great fun, but grandfather frog thought that his last day had come. xii grandfather frog keeps on grandfather frog is old and wise, but even age is foolish. i'm sure you'll all agree with me his stubbornness was mulish. that his very last day had come grandfather frog was sure. he didn't have the least doubt about it. here he was at the mercy of bowser the hound out on the green meadows far from the dear, safe smiling pool. every time he moved, bowser flipped him over on his back and danced around him, barking with joy. every minute grandfather frog expected to feel bowser's terrible teeth, and he grew cold at the thought. when he found that he couldn't get away, he just lay still. he was too tired and frightened to do much of anything else, anyway. now when he lay still, he spoiled bowser's fun, for it was seeing him jump and kick his long legs that tickled bowser so. bowser tossed him up in the air two or three times, but grandfather frog simply lay where he fell without moving. "bow, wow, wow!" cried bowser, in his great deep voice. grandfather frog didn't so much as blink his great goggly eyes. bowser sniffed him all over. "i guess i've frightened him to death," said bowser, talking to himself. "i didn't mean to do that. i just wanted to have some fun with him." with that, bowser took one more sniff and then trotted off to try to find something more exciting. you see, he hadn't had the least intention in the world of really hurting grandfather frog. grandfather frog kept perfectly still until he was sure that bowser was nowhere near. then he gave a great sigh of relief and crawled under a big mullein leaf to rest, and think things over. "chugarum, that was a terrible experience; it was, indeed!" said he to himself, shivering at the very thought of what he had been through. "nothing like that ever happened to me in the smiling pool. i've always said that the smiling pool is a better place in which to live than is the great world, and now i know it. the question is, what had i best do now?" now right down in his heart grandfather frog knew the answer. of course the best thing to do was to go straight back to the smiling pool as fast as he could. but grandfather frog is stubborn. yes, sir, he certainly is stubborn. and stubbornness is often just another name for foolishness. he had told jerry muskrat that he was going out to see the great world. now if he went back, jerry would laugh at him. "i won't!" said grandfather frog. "what won't you do?" asked a voice so close to him that grandfather frog made a long jump before he thought. you see, at the smiling pool he always jumped at the least hint of danger, and because one jump always took him into the water, he was always safe. but there was no water here, and that jump took him right out where anybody passing could see him. then he turned around to see who had startled him so. it was danny meadow mouse. "i won't go back to the smiling pool until i have seen the great world," replied grandfather frog gruffly. [illustration: "you won't see much of the great world if you jump like that every time you get a scare," said danny. _page 62._] "you won't see much of the great world if you jump like that every time you get a scare," said danny, shaking his head. "no, sir, you won't see much of the great world, because one of these times you'll jump right into the claws of old whitetail the marsh hawk, or his cousin redtail, or reddy fox. you take my advice, grandfather frog, and go straight back to the smiling pool. you don't know enough about the great world to take care of yourself." but grandfather frog was set in his ways, and nothing that danny meadow mouse could say changed his mind in the least. "i started out to see the great world, and i'm going to keep right on," said he. "all right," said danny at last. "if you will, i suppose you will. i'll go a little way with you just to get you started right." "thank you," replied grandfather frog. "let's start right away." xiii danny meadow mouse feels responsible responsible is a great big word. but it is just as big in its meaning as it is in its looks, and that is the way words should be, i think, don't you? anyway, re-spon-sible is the way danny meadow mouse felt when he found grandfather frog out on the green meadows so far from the smiling pool and so stubborn that he would keep on to see the great world instead of going back to his big green lily-pad in the smiling pool, where he could take care of himself. you remember peter rabbit felt re-spon-sible when he brought little miss fuzzy tail down from the old pasture to the dear old briar-patch. he felt that it was his business to see to it that no harm came to her, and that is just the way danny meadow mouse felt about grandfather frog. you see, danny knew that if grandfather frog was going to jump like that every time he was frightened, he wouldn't get very far in the great world. it might be the right thing to do in the smiling pool, where the friendly water would hide him from his enemies, but it was just the wrong thing to do on the green meadows or in the green forest. danny had learned, when a very tiny fellow, that there the only safe thing to do when danger was near was to sit perfectly still and hardly breathe. now danny meadow mouse is fond of grandfather frog, and he couldn't bear to think that something dreadful might happen to him. so when he found that he couldn't get grandfather frog to go back to the smiling pool, he made up his mind that he just _had_ to go along with grandfather frog to try to keep him out of danger. yes, sir, he just _had_ to do it. he felt re-spon-sible for grandfather frog's safety. so here they were, danny meadow mouse running ahead, anxious and worried and watching sharply for signs of danger, and grandfather frog puffing along behind, bound to see the great world which his cousin, old mr. toad, said was a better place to live in than the smiling pool. now danny has a great many private little paths under the grass all over the green meadows, and along these he can scamper ever so fast without once showing himself to those who may be looking for him. of course he started to take grandfather frog along one of these little paths. but grandfather frog doesn't walk or run; he jumps. there wasn't room in danny's little paths for jumping, as they soon found out. grandfather frog simply couldn't follow danny along those little paths. danny sat down to think, and puckered his brows anxiously. he was more worried than ever. it was very clear that grandfather frog would have to travel out in the open, where there was room for him to jump, and where also he would be right out in plain sight of all who happened along. once more danny urged him to go back to the smiling pool, but he might just as well have talked to a stick or a stone. grandfather frog had started out to see the great world, and he was going to see it. danny sighed. "if you will, you will, i suppose," said he, "and i guess the only place you can travel in any comfort is the lone little path. it is dangerous, very dangerous, but i guess you will have to do it." "chugarum!" replied grandfather frog, "i'm not afraid. you show me the lone little path and then go about your business, danny meadow mouse." so danny led the way to the lone little path, and grandfather frog sighed with relief, for here he could jump without getting all tangled up in long grass and without hurting his tender feet on sharp stubble where the grass had been cut. but danny felt more worried than ever. he wouldn't leave grandfather frog because, you know, he felt re-spon-sible for him, and at the same time he was terribly afraid, for he felt sure that some of their enemies would see them. he wanted to go back, but he kept right on, and that shows just what a brave little fellow danny meadow mouse was. xiv grandfather frog has a strange ride a thousand things may happen to, ten thousand things befall, the traveler who careless is, or thinks he knows it all. grandfather frog, jumping along behind danny meadow mouse up the lone little path, was beginning to think that danny was the most timid and easiest frightened of all the little meadow people of his acquaintance. danny kept as much under the grass that overhung the lone little path as he could. when there were perfectly bare places, danny looked this way and looked that way anxiously and then scampered across as fast as he could make his little legs go. when he was safely across, he would wait for grandfather frog. if a shadow passed over the grass, danny would duck under the nearest leaf and hold his breath. "foolish!" muttered grandfather frog. "foolish, foolish to be so afraid! now, i'm not afraid until i see something to be afraid of. time enough then. what's the good of looking for trouble all the time? now, here i am out in the great world, and i'm not afraid. and here's danny meadow mouse, who has lived here all his life, acting as if he expected something dreadful to happen any minute. pooh! how very, very foolish!" now grandfather frog is old and in the smiling pool he is accounted very, very wise. but the wisest sometimes become foolish when they think that they know all there is to know. it was so with grandfather frog. it was he who was foolish and not danny meadow mouse. you see danny knew all the dangers on the green meadows, and how many sharp eyes were all the time watching for him. he had long ago learned that the only way to feel safe was to feel afraid. you see, then he was watching for danger every minute, and so he wasn't likely to be surprised by his hungry enemies. so while grandfather frog was looking down on danny for being so timid, danny was really doing the wisest thing. more than that, he was really very, very brave. he was showing grandfather frog the way up the lone little path to see the great world, when he himself would never, never have thought of traveling anywhere but along his own secret little paths, just because grandfather frog couldn't jump anywhere excepting where the way was fairly clear, as in the lone little path, and danny was afraid that unless grandfather frog had some one with him to watch out for him, he would surely come to a sad end. the farther they went with nothing happening, the more foolish danny's timid way of running and hiding seemed to grandfather frog, and he was just about to tell danny just what he thought, when danny dived into the long grass and warned grandfather frog to do the same. but grandfather frog didn't. "chugarum!" said he, "i don't see anything to be afraid of, and i'm not going to hide until i do." so he sat still right where he was, in the middle of the lone little path, looking this way and that way, and seeing nothing to be afraid of. and just then around a turn in the lone little path came--who do you think? why farmer brown's boy! he saw grandfather frog and with a whoop of joy he sprang for him. grandfather frog gave a frightened croak and jumped, but he was too late. before he could jump again farmer brown's boy had him by his long hind-legs. "ha, ha!" shouted farmer brown's boy, "i believe this is the very old chap i have tried so often to catch in the smiling pool. these legs of yours will be mighty fine eating, mr. frog. they will, indeed." with that he tied grandfather frog's legs together and went on his way across the green meadows with poor old grandfather frog dangling from the end of a string. it was a strange ride and a most uncomfortable one, and with all his might grandfather frog wished he had never thought of going out into the great world. xv grandfather frog gives up hope with his legs tied together, hanging head down from the end of a string, grandfather frog was being carried he knew not where by farmer brown's boy. it was dreadful. half-way across the green meadows the merry little breezes of old mother west wind came dancing along. at first they didn't see grandfather frog, but presently one of them, rushing up to tease farmer brown's boy by blowing off his hat, caught sight of grandfather frog. now the merry little breezes are great friends of grandfather frog. many, many times they have blown foolish green flies over to him as he sat on his big green lily-pad, and they are very fond of him. so when this one caught sight of him in such a dreadful position, he forgot all about teasing farmer brown's boy. he raced away to tell the other merry little breezes. for a minute they were perfectly still. they forgot all about being merry. "it's awful, just perfectly awful!" cried one. "we must do something to help grandfather frog!" cried another. "of course we must," said a third. "but what can we do?" asked a fourth. nobody replied. they just thought and thought and thought. finally the first one spoke. "we might try to comfort him a little," said he. "of course we will do that!" they shouted all together. "and if we throw dust in the face of farmer brown's boy and steal his hat, perhaps he will put grandfather frog down," continued the merry little breeze. "the very thing!" the others cried, dancing about with excitement. "then we can rush about and tell all grandfather frog's friends what has happened to him and where he is. perhaps some of them can help us," the little breeze continued. they wasted no more time talking, but raced after farmer brown's boy as fast as they could go. one of them, who was faster than the others, ran ahead and whispered in grandfather frog's ear that they were coming to help him. but poor old grandfather frog couldn't be comforted. he couldn't see what there was that the merry little breezes could do. his legs smarted where the string cut into the skin, and his head ached, for you know he was hanging head down. no, sir, grandfather frog couldn't be comforted. he was in a terrible fix, and he couldn't see any way out of it. he hadn't the least bit of hope left. and all the time farmer brown's boy was trudging along, whistling merrily. you see, it didn't occur to him to think how grandfather frog must be suffering and how terribly frightened he must be. he wasn't cruel. no, indeed, farmer brown's boy wasn't cruel. that is, he didn't mean to be cruel. he was just thoughtless, like a great many other boys, and girls too. so he went whistling on his way until he reached the long lane leading from the green meadows up to farmer brown's dooryard. no sooner was he in the long lane than something happened. a great cloud of dust and leaves and tiny sticks was dashed in his face and nearly choked him. dirt got in his eyes. his hat was snatched from his head and went sailing over into the garden. he dropped grandfather frog and felt for his handkerchief to wipe the dirt from his eyes. "phew!" exclaimed farmer brown's boy, as he started after his hat. "it's funny where that wind came from so suddenly!" but you know and i know that it was the merry little breezes working together who made up that sudden wind. and grandfather frog ought to have known it too, but he didn't. you see the dust had got in his nose and eyes just as it had in those of farmer brown's boy, and he was so frightened and confused that he couldn't think. so he lay just where farmer brown's boy dropped him, and he didn't have any more hope than before. xvi the merry little breezes work hard the merry little breezes almost shouted aloud with delight when they saw farmer brown's boy drop grandfather frog to feel for his handkerchief and wipe out the dust which they had thrown in his eyes. then he had to climb the fence and chase his hat through the garden. they would let him almost get his hands on it and then, just as he thought that he surely had it, they would snatch it away. it was great fun for the merry little breezes. but they were not doing it for fun. no, indeed, they were not doing it for fun! they were doing it to lead farmer brown's boy away from grandfather frog. just as soon as they dared, they dropped the hat and then separated and rushed away in all directions across the green meadows, over to the green forest, and down to the smiling pool. what were they going for? why, to hunt for some of grandfather frog's friends and ask their help. you see, the merry little breezes could make farmer brown's boy drop grandfather frog, but they couldn't untie a knot or cut a string, and this is just what had got to be done to set grandfather frog free, for his hind-legs were tied together. so now they were looking for some one with sharp teeth, who thought enough of grandfather frog to come and help him. one thought of striped chipmunk and started for the old stone wall to look for him. another went in search of danny meadow mouse. a third headed for the dear old briar-patch after peter rabbit. a fourth remembered jimmy skunk and how he had once set blacky the crow free from a snare. a fifth remembered what sharp teeth happy jack squirrel has and hurried over to the green forest to look for him. a sixth started straight for the smiling pool to tell jerry muskrat. and every one of them raced as fast as he could. all this time grandfather frog was without hope. yes, sir, poor old grandfather frog was wholly in despair. you see, he didn't know what the merry little breezes were trying to do, and he was so frightened and confused that he couldn't think. when farmer brown's boy dropped him, he lay right where he fell for a few minutes. then, right close at hand, he saw an old board. without really thinking, he tried to get to it, for there looked as if there might be room for him to hide under it. it was hard work, for you know his long hind-legs, which he uses for jumping, were tied together. the best he could do was to crawl and wriggle and pull himself along. just as farmer brown's boy started to climb the fence back into the long lane, his hat in his hand, grandfather frog reached the old board and crawled under it. now when the merry little breezes had thrown the dust in farmer brown's boy's face and snatched his hat, he had dropped grandfather frog in such a hurry that he didn't notice just where he did drop him, so now he didn't know the exact place to look for him. but he knew pretty near, and he hadn't the least doubt but that he would find him. he had just started to look when the dinner horn sounded. farmer brown's boy hesitated. he was hungry. if he was late, he might lose his dinner. he could come back later to look for grandfather frog, for with his legs tied grandfather frog couldn't get far. so, with a last look to make sure of the place, farmer brown's boy started for the house. if the merry little breezes had known this, they would have felt ever so much better. but they didn't. so they hurried as fast as ever they could to find grandfather frog's friends and worked until they were almost too tired to move, for it seemed as if every single one of grandfather frog's friends had taken that particular day to go away from home. so while farmer brown's boy ate his dinner, and grandfather frog lay hiding under the old board in the long lane, the merry little breezes did their best to find help for him. xvii striped chipmunk cuts the string "hippy hop! flippy flop! all on a summer day my mother turned me from the house and sent me out to play!" striped chipmunk knew perfectly well that that was just nonsense, but striped chipmunk learned a long time ago that when you are just bubbling right over with good feeling, there is fun in saying and doing foolish things, and that is just how he was feeling. so he ran along the old rail fence on one side of the long lane, saying foolish things and cutting up foolish capers just because he felt so good, and all the time seeing all that those bright little eyes of his could take in. now striped chipmunk and the merry little breezes of old mother west wind are great friends, very great friends, indeed. almost every morning they have a grand frolic together. but this morning the merry little breezes hadn't come over to the old stone wall where striped chipmunk makes his home. anyway, they hadn't come at the usual time. striped chipmunk had waited a little while and then, because he was feeling so good, he had decided to take a run down the long lane to see if anything new had happened there. that is how it happened that when one of the merry little breezes did go to look for him, and was terribly anxious to ask him to come to the help of grandfather frog, he was nowhere to be found. but striped chipmunk didn't know anything about that. he scampered along the top rails of the old fence, jumped up on top of a post, and sat up to wash his face and hands, for striped chipmunk is very neat and cannot bear to be the least bit dirty. he looked up and winked at ol' mistah buzzard, sailing round and round way, way up in the blue, blue sky. he chased his own tail round and round until he nearly fell off of the post. he made a wry face in the direction of redtail the hawk, whom he could see sitting in the top of a tall tree way over on the green meadows. he scolded bowser the hound, who happened to come trotting up the long lane, and didn't stop scolding until bowser was out of sight. then he kicked up his heels and whisked along the old fence again. half-way across a shaky old rail, he suddenly stopped. his bright eyes had seen something that filled him with curiosity, quite as much curiosity as peter rabbit would have had. it was a piece of string. yes, sir, it was a piece of string. now striped chipmunk often had found pieces of string, so there was nothing particularly interesting in the string itself. what did interest him and make him very curious was the fact that this piece of string kept moving. every few seconds it gave a little jerk. whoever heard of a piece of string moving all by itself? certainly striped chipmunk never had. he couldn't understand it. for a few minutes he watched it from the top rail of the old fence. then he scurried down to the ground and, a few steps at a time, stopping to watch sharply between each little run, he drew nearer and nearer to that queer acting string. it gave him a funny feeling inside to see a string acting like that, so he was very careful not to get too near. he looked at it from one side, then ran around and looked at it from the other side. at last he got where he could see that one end of the string was under an old board, and then he began to understand. of course there was somebody hiding under that old board and jerking the string. [illustration: he seized the other end of the string and began to pull. _page 88._] striped chipmunk sat down and scratched his head thoughtfully. whoever was pulling that string couldn't be very big, or they would never have been able to crawl under that old board, therefore he needn't be afraid. a gleam of mischief twinkled in striped chipmunk's eyes. he seized the other end of the string and began to pull. such a jerking and yanking as began right away! but he held on and pulled harder. then out from under the old board appeared the queer webbed feet of grandfather frog tied together. striped chipmunk was so surprised that he let go of the string and nearly fell over backward. "why, grandfather frog, what under the sun are you doing here?" he shouted. when striped chipmunk let go of the string, grandfather frog promptly drew his feet back under the old board, but when he heard striped chipmunk's voice, he slowly and painfully crawled out. he told how he had been caught and tied by farmer brown's boy and finally dropped near the old board. he told how terribly frightened he was, and how sore his legs were. striped chipmunk didn't wait for him to finish. in a flash he was at work with his sharp teeth and had cut the cruel string before grandfather frog had finished his story. xviii grandfather frog hurries away when striped chipmunk cut the string that bound the long legs of grandfather frog together, grandfather frog was so relieved that he hardly knew what to do. of course he thanked striped chipmunk over and over again. striped chipmunk said that it was nothing, just nothing at all, and that he was very glad indeed to help grandfather frog. "we folks who live out in the great world have to help one another," said striped chipmunk, "because we never know when we may need help ourselves. now you take my advice, grandfather frog, and go back to the smiling pool as fast as you can. the great world is no place for an old fellow like you, because you don't know how to take care of yourself." now when he said that, striped chipmunk made a great mistake. old people never like to be told that they are old or that they do not know all there is to know. grandfather frog straightened up and tried to look very dignified. "chugarum!" said he, "i'd have you to know, striped chipmunk, that people were coming to me for advice before you were born. it was just an accident that farmer brown's boy caught me, and i'd like to see him do it again. yes, sir, i'd like to see him do it again!" dear me, dear me! grandfather frog was boasting. if he had been safe at home in the smiling pool, there might have been some excuse for boasting, but way over here in the long lane, not even knowing the way back to the smiling pool, it was foolish, very foolish indeed. no one knew that better than striped chipmunk, but he has a great deal of respect for grandfather frog, and he knew too that grandfather frog was feeling very much out of sorts and very much mortified to think that he had been caught in such a scrape, so he put a hand over his mouth to hide a smile as he said: "of course he isn't going to catch you again. i know how wise and smart you are, but you look to me very tired, and there are so many dangers out here in the great world that it seems to me that the very best thing you can do is to go back to the smiling pool." but grandfather frog is stubborn, you know. he had started out to see the great world, and he didn't want the little people of the green meadows and the green forest to think that he was afraid. the truth is, grandfather frog was more afraid of being laughed at than he was of the dangers around him, which shows just how foolish wise people can be sometimes. so he shook his head. "chugarum!" said he, "i am going to see the great world first, and then i am going back to the smiling pool. do you happen to know where there is any water? i am very thirsty." now over on the other side of the long lane was a spring where farmer brown's boy filled his jug with clear cold water to take with him to the cornfield when he had to work there. striped chipmunk knew all about that spring, for he had been there for a drink many times. so he told grandfather frog just where the spring was and how to get to it. he even offered to show the way, but grandfather frog said that he would rather go alone. "watch out, grandfather frog, and don't fall in, because you might not be able to get out again," warned striped chipmunk. grandfather frog looked up sharply to see if striped chipmunk was making fun of him. the very idea of any one thinking that he, who had lived in the water all his life, couldn't get out when he pleased! but striped chipmunk looked really in earnest, so grandfather frog swallowed the quick retort on the tip of his tongue, thanked striped chipmunk, and hurried away to look for the spring, for he was very, very thirsty. besides, he was very, very hot, and he hurried still faster as he thought of the cool bath he would have when he found that spring. xix grandfather frog jumps into more trouble some people are heedless and run into trouble. some people are stupid and walk into trouble. grandfather frog was both heedless and stupid and jumped into trouble. when striped chipmunk told him where the spring was, it seemed to him that he couldn't wait to reach it. you see, grandfather frog had spent all his life in the smiling pool, where he could get a drink whenever he wanted it by just reaching over the edge of his big green lily-pad. whenever he was too warm, all he had to do was to say "chugarum!" and dive head first into the cool water. so he wasn't used to going a long time without water. jump, jump, jump! grandfather frog was going as fast as ever he could in the direction striped chipmunk had pointed out. every three or four jumps he would stop for just a wee, wee bit of rest, then off he would go again, jump, jump, jump! and each jump was a long one. peter rabbit certainly would have been envious if he could have seen those long jumps of grandfather frog. at last the ground began to grow damp. the farther he went, the damper it grew. presently it became fairly wet, and there was a great deal of soft, cool, wet moss. how good it did feel to grandfather frog's poor tired feet! "must be i'm most there," said grandfather frog to himself, as he scrambled up on a big mossy hummock, so as to look around. right away he saw a little path from the direction of the long lane. it led straight past the very hummock on which grandfather frog was sitting, and he noticed that where the ground was very soft and wet, old boards had been laid down. that puzzled grandfather frog a great deal. "it's a sure enough path," said he. "but what under the blue, blue sky does any one want to spoil it for by putting those boards there?" you see, grandfather frog likes the soft wet mud, and he couldn't understand how any one, even farmer brown's boy, could prefer a hard dry path. of course he never had worn shoes himself, so he couldn't understand why any one should want dry feet when they could just as well have wet ones. he was still puzzling over it when he heard a sound that made him nearly lose his balance and tumble off the hummock. it was a whistle, the whistle of farmer brown's boy! grandfather frog knew it right away, because he often had heard it over by the smiling pool. the whistle came from over in the long lane. farmer brown's boy had had his dinner and was on his way back to look for grandfather frog where he had been dropped. grandfather frog actually grinned as he thought how surprised farmer brown's boy was going to be when he could find no trace of him. suddenly the smile seemed to freeze on grandfather frog's face. that whistle was coming nearer! farmer brown's boy had left the long lane and was coming along the little path. the truth is, he was coming for a drink at the spring, but grandfather frog didn't think of this. he was sure that in some way farmer brown's boy had found out which way he had gone and was coming after him. he crouched down as flat as he could on the big hummock and held his breath. farmer brown's boy went straight past. just a few steps beyond, he stopped and knelt down. peeping through the grass, grandfather frog saw him dip up beautiful clear water in an old cup and drink. then grandfather frog knew just where the spring was. a few minutes later, farmer brown's boy passed again, still whistling, on his way to the long lane. grandfather frog waited only long enough to be sure that he had really gone. then, with bigger jumps than ever, he started for the spring. a dozen long jumps, and he could see the water. two more jumps and then a long jump, and he had landed in the spring with a splash! "chugarum!" cried grandfather frog. "how good the water feels!" and all the time, grandfather frog had jumped straight into more trouble. xx grandfather frog loses heart look before you leap; the water may be deep. that is the very best kind of advice, but most people find that out when it is too late. grandfather frog did. of course he had heard that little verse all his life. indeed, he had been very fond of saying it to those who came to the smiling pool to ask his advice. but grandfather frog seemed to have left all his wisdom behind him when he left the smiling pool to go out into the great world. you see, it is very hard work for any one whose advice has been sought to turn right around and take advice themselves. so grandfather frog had been getting into scrapes ever since he started out on his foolish journey, and now here he was in still another, and he had landed in it head first, with a great splash. of course, when he had seen the cool, sparkling water of the spring, it had seemed to him that he just couldn't wait another second to get into it. he was so hot and dry and dreadfully thirsty and uncomfortable! and so--oh, dear me!--grandfather frog didn't look at all before he leaped. no, sir, he didn't! he just dived in with a great long jump. oh, how good that water felt! for a few minutes he couldn't think of anything else. it was cooler than the water of the smiling pool, because, as you know, it was a spring. but it felt all the better for that, and grandfather frog just closed his eyes and floated there in pure happiness. presently he opened his eyes to look around. then he blinked them rapidly for a minute or so. he rubbed them to make sure that he saw aright. his heart seemed to sink way, way down towards his toes. "chugarum!" exclaimed grandfather frog, "chugarum!" and after that for a long time he didn't say a word. you see, it was this way. all around him rose perfectly straight smooth walls. he could look up and see a little of the blue, blue sky right overhead and whispering leaves of trees and bushes. over the edge of the smooth straight wall grasses were bending. but they were so far above his head, so dreadfully far! _there wasn't any place to climb out!_ grandfather frog was in a prison! he didn't understand it at all, but it was so. of course, farmer brown's boy could have told him all about it. a long time before farmer brown himself had found that spring, and because the water was so clear and cold and pure, he had cleared away all the dirt and rubbish around it. then he had knocked the bottom out of a nice clean barrel and had dug down where the water bubbled up out of the sand and had set the barrel down in this hole and had filled in the bottom with clean white sand for the water to bubble up through. about half-way up the barrel he had cut a little hole for the water to run out as fast as it bubbled in at the bottom. of course the water never could fill the barrel, because when it reached that hole, it ran out. this left a straight, smooth wall up above, a wall altogether too high for grandfather frog to jump over from the inside. poor old grandfather frog! he wished more than ever that he never, never had thought of leaving the smiling pool to see the great world. round and round he swam, but he couldn't see any way out of it. the little hole where the water ran out was too small for him to squeeze through, as he found out by trying and trying. so far as he could see, he had just got to stay there all the rest of his life. worse still, he knew that farmer brown's boy sometimes came to the spring for a drink, for he had seen him do it. that meant that the very next time he came, he would find grandfather frog, because there was no place to hide. when grandfather frog thought of that, he just lost heart. yes, sir, he just lost heart. he gave up all hope of ever seeing the smiling pool again, and two big tears ran out of his big goggly eyes. xxi the merry little breezes try to comfort grandfather frog when the merry little breezes of old mother west wind had left grandfather frog in the long lane where farmer brown's boy had dropped him, and had hurried as fast as ever they could to try to find some of his friends to help him, not one of them had been successful. no one was at home, and no one was in any of the places where they usually were to be found. the merry little breezes looked and looked. then, one by one, they sadly turned back to the long lane. they felt so badly that they just hated to go back where they had left grandfather frog. when they got there, they found striped chipmunk, who now was scolding farmer brown's boy as fast as his tongue could go. "where is he?" cried the merry little breezes excitedly. striped chipmunk stopped scolding long enough to point to farmer brown's boy, who was hunting in the grass for some trace of grandfather frog. "we don't mean him, you stupid! we can see him for ourselves. where's grandfather frog?" cried the merry little breezes, all speaking at once. "i don't know," replied striped chipmunk, "and what's more, i don't care!" now this wasn't true, for striped chipmunk isn't that kind. it was mostly talk, and the merry little breezes knew it. they knew that striped chipmunk really thinks a great deal of grandfather frog, just as they do. so they pretended not to notice what he said or how put out he seemed. after a while, he told them that he had set grandfather frog free and that then he had started for the spring on the other side of the long lane. the merry little breezes were delighted to hear the good news, and they said such a lot of nice things to striped chipmunk that he quite forgot to scold farmer brown's boy. then they started for the spring, dancing merrily, for they felt sure that there grandfather frog was all right, and they expected to find him quite at home. "hello, grandfather frog!" they shouted, as they peeped into the spring. "how do you like your new home?" grandfather frog made no reply. he just rolled his great goggly eyes up at them, and they were full of tears. "why--why--why, grandfather frog, what is the matter now?" they cried. "chugarum," said grandfather frog, and his voice sounded all choky, "i can't get out." then they noticed for the first time how straight and smooth the walls of the spring were and how far down grandfather frog was, and they knew that he spoke the truth. they tried bending down the grasses that grew around the edge of the spring, but none were long enough to reach the water. if they had stopped to think, they would have known that grandfather frog couldn't have climbed up by them, anyway. then they tried to lift a big stick into the spring, but it was too heavy for them, and they couldn't move it. however, they did manage to blow an old shingle in, and this gave grandfather frog something to sit on, so that he began to feel a little better. then they said all the comforting things they could think of. they told him that no harm could come to him there, unless farmer brown's boy should happen to see him. [illustration: "that's just what i'm afraid of!" croaked grandfather frog. _page 109_.] "that's just what i am afraid of!" croaked grandfather frog. "he is sure to see me if he comes for a drink, for there is no place for me to hide." "perhaps he won't come," said one of the little breezes hopefully. "if he does come, you can hide under the piece of shingle, and then he won't know you are here at all," said another. grandfather frog brightened up. "that's so!" said he. "that's a good idea, and i'll try it." then one of the merry little breezes promised to keep watch for farmer brown's boy, and all the others started off on another hunt for some one to help grandfather frog out of this new trouble. xxii grandfather frog's troubles grow head first in; no way out; it's best to know what you're about! grandfather frog had had plenty of time to realize how very true this is. as he sat on the old shingle which the merry little breezes had blown into the spring where he was a prisoner, he thought a great deal about that little word "if." _if_ he hadn't left the smiling pool, _if_ he hadn't been stubborn and set in his ways, _if_ he hadn't been in such a hurry, _if_ he had looked to see where he was leaping--well, any one of these _ifs_ would have kept him out of his present trouble. it really wasn't so bad in the spring. that is, it wouldn't have been so bad but for the fear that farmer brown's boy might come for a drink and find him there. that was grandfather frog's one great fear, and it gave him bad dreams whenever he tried to take a nap. he grew cold all over at the very thought of being caught again by farmer brown's boy, and when at last one of the merry little breezes hurried up to tell him that farmer brown's boy actually was coming, poor old grandfather frog was so frightened that the merry little breeze had to tell him twice to hide under the old shingle as it floated on the water. at last he got it through his head, and drawing a very long breath, he dived into the water and swam under the old shingle. he was just in time. yes, sir, he was just in time. if farmer brown's boy hadn't been thinking of something else, he certainly would have noticed the little rings on the water made by grandfather frog when he dived in. but he was thinking of something else, and it wasn't until he dipped a cup in for the second time that he even saw the old shingle. "hello!" he exclaimed. "that must have blown in since i was here yesterday. we can't have anything like that in our nice spring." with that he reached out for the old shingle, and grandfather frog, hiding under it, gave himself up for lost. but the anxious little breeze had been watching sharply and the instant he saw what farmer brown's boy was going to do, he played the old, old trick of snatching his hat from his head. the truth is, he couldn't think of anything else to do. farmer brown's boy grabbed at his hat, and then, because he was in a hurry and had other things to do, he started off without once thinking of the old shingle again. "chugarum!" cried grandfather frog, as he swam out from under the shingle and climbed up on it, "that certainly was a close call. if i have many more like it, i certainly shall die of fright." nothing more happened for a long time, and grandfather frog was wondering if it wouldn't be safe to take a nap when he saw peeping over the edge above him two eyes. they were greenish yellow eyes, and they stared and stared. grandfather frog stared and stared back. he just couldn't help it. he didn't know who they belonged to. he couldn't remember ever having seen them before. he was afraid, and yet somehow he couldn't make up his mind to jump. he stared so hard at the eyes that he didn't notice a long furry paw slowly, very slowly, reaching down towards him. nearer it crept and nearer. then suddenly it moved like a flash. grandfather frog felt sharp claws in his white and yellow waistcoat, and before he could even open his mouth to cry "chugarum," he was sent flying through the air and landed on his back in the grass. pounce! two paws pinned him down, and the greenish yellow eyes were not an inch from his own. they belonged to black pussy, farmer brown's cat. xxiii the dear old smiling pool once more black pussy was having a good time. grandfather frog wasn't. it was great fun for black pussy to slip a paw under grandfather frog and toss him up in the air. it was still more fun to pretend to go away, but to hide instead, and the instant grandfather frog started off, to pounce upon him and cuff him and roll him about. but there wasn't any fun in it for grandfather frog. in the first place, he didn't know whether or not black pussy liked frogs to eat, and he was terribly frightened. in the second place, black pussy didn't always cover up her claws, and they pricked right through grandfather frog's white and yellow waistcoat and hurt, for he is very tender there. at last black pussy grew tired of playing, so catching up grandfather frog in her mouth, she started along the little path from the spring to the long lane. grandfather frog didn't even kick, which was just as well, because if he had, black pussy would have held him tighter, and that would have been very uncomfortable indeed. "it's all over, and this is the end," moaned grandfather frog. "i'm going to be eaten now. oh, why, why did i ever leave the smiling pool?" just as black pussy slipped into the long lane, grandfather frog heard a familiar sound. it was a whistle, a merry whistle. it was the whistle of farmer brown's boy. it was coming nearer and nearer. a little bit of hope began to stir in the heart of grandfather frog. he didn't know just why, but it did. always he had been in the greatest fear of farmer brown's boy, but now--well, if farmer brown's boy should take him, he might get away from him as he did before, but he was very sure that he never, never could get away from black pussy. the whistle drew nearer. black pussy stopped. then she began to make a queer whirring sound deep down in her throat. "hello, black pussy! have you been hunting? come here and show me what you've got," cried a voice. black pussy arched up her back and began to rub against the legs of farmer brown's boy, and all the time the whir, ring sound in her throat grew louder and louder. farmer brown's boy stooped down to see what she had in her mouth. "why," he exclaimed, "i do believe this is the very same old frog that got away from me! you don't want him, puss. i'll just put him in my pocket and take him up to the house by and by." with that he took grandfather frog from black pussy and dropped him in his pocket. he patted black pussy, called her a smart cat, and then started on his way, whistling merrily. it was dark and rather close in that pocket, but grandfather frog didn't mind this. it was a lot better than feeling sharp teeth and claws all the time. he wondered how soon they would reach the house and what would happen to him then. after what seemed like a long, long time, he felt himself swung through the air, and then he landed on the ground with a thump that made him grunt. farmer brown's boy had taken off his coat and thrown it down. the whistling stopped. everything was quiet. grandfather frog waited and listened, but not a sound could he hear. then he saw a little ray of light creeping into his prison. he squirmed and pushed, and all of a sudden he was out of the pocket. the bright light made him blink. as soon as he could see, he looked to see where he was. then he rubbed his eyes with both hands and looked again. he wasn't at farmer brown's house at all. where do you think he was? why, right on the bank of the smiling pool, and a little way off was farmer brown's boy fishing! "chugarum!" cried grandfather frog, and it was the loudest, gladdest chugarum that the smiling pool ever had heard. "chugarum!" he cried again, and with a great leap he dived with a splash into the dear old smiling pool, which smiled more than ever. and never again has grandfather frog tried to see the great world. he is quite content to leave it to those who like to dwell there. and since his own wonderful adventures, he has been ready to believe anything he is told about what happens there. nothing can surprise him, not even the astonishing things that happened to chatterer the red squirrel, about which it takes a whole book to tell. university of kansas museum of natural history vol. 20, no. 1, pp. 1-45, 22 figs. february 20, 1970 a taxonomic revision of the leptodactylid frog genus syrrhophus cope by john d. lynch university of kansas lawrence 1970 university of kansas publications, museum of natural history editors of this number: frank b. cross, philip s. humphrey, william e. duellman volume 20, no. 1, pp. 1-45, 22 figs. published february 20, 1970 university of kansas lawrence, kansas printed by the university of kansas printing service lawrence, kansas 1970 a taxonomic revision of the leptodactylid frog genus syrrhophus cope by john d. lynch introduction cope (1878) proposed the genus _syrrhophus_ for a medium-sized leptodactylid frog from central texas; in the ensuing 75 years the genus was expanded to include a heterogeneous group of frogs ranging from texas to peru. taylor (1952) and firschein (1954) limited the genus to several species of frogs occurring in guatemala, méxico, and texas. lynch (1968) provided a definition of the previously loosely-defined genus. with the exception of taylor (1952), who treated the costa rican species, none of these authors dealt with the present status of the nineteen species erroneously assigned to _syrrhophus_. these species are listed in tables 1 and 2 with the name currently applied. some of them are new combinations and their justifications will be published elsewhere. gorham (1966) is the most recent author to include south american species in the genus _syrrhophus_. smith and taylor (1948) recognized two species groups of the genus in méxico, an eastern and a western group (here termed complexes for purposes of discussion), separated on the basis of the number of palmar (metacarpal) tubercles (three palmar tubercles in the members of the eastern complex and two in those of the western complex). duellman (1958) reviewed the species of the genus occurring in western méxico and concluded that there were five species (two polytypic). dixon and webb (1966) described an additional species from jalisco, méxico. the distributions of some species have been extended, but otherwise the western complex of species remains unchanged since duellman's review. smith and taylor (1948) recognized seven species of the genus in eastern méxico. firschein revised the eastern complex (as then understood), and in so doing added one new species and treated _syrrhophus verruculatus_ as a _nomen dubium_. dixon (1957) redefined the related genus _tomodactylus_ and transferred _t. macrotympanum_ taylor to the genus _syrrhophus_. neill (1965) described a new subspecies of _s. leprus_ from british honduras. two species (_s. gaigeae_ and _s. marnockii_) were recognized in texas until milstead, mecham, and mcclintock (1950) synonymized _s. gaigeae_ with _s. marnockii_. thus, at present, nine species (one polytypic) are recognized on the eastern slopes and lowlands from central texas to british honduras. these are currently placed on one species group equivalent to the western complex reviewed by duellman (1958). table 1--species described as members of the genus _syrrhophus_ but now placed in other genera. ======================================================================= trivial name and author current combination ---------------------------------------------------------------------- _areolatus_ boulenger, 1898 _eleutherodactylus areolatus_ _calcaratus_ andersson, 1945 _eleutherodactylus anderssoni_ _caryophyllaceus_ barbour, 1928 _eleutherodactylus caryophyllaceus_ _coeruleus_ andersson, 1945 _eleutherodactylus coeruleus_ _ineptus_ barbour, 1928 _eleutherodactylus diastema_ _juninensis_ shreve, 1938 _eupsophus juninensis_ _lutosus_ barbour and dunn, 1921 _eleutherodactylus lutosus_ _molinoi_ barbour, 1928 _eleutherodactylus molinoi_ _montium_ shreve, 1938 _niceforonia montia_ _mystaceus_ barbour, 1922 _eleutherodactylus rhodopis_ _obesus_ barbour, 1928 _eleutherodactylus punctariolus_ _omiltemanus_ gunther, 1900 _eleutherodactylus omiltemanus_[1] _pardalis_ barbour, 1928 _eleutherodactylus pardalis_ ======================================================================= [1] new combination. table 2--species incorrectly regarded as members of the genus _syrrhophus_ but described as members of other genera. ========================================================================== trivial name, original generic assignment, and author current combination ------------------------------------------------------------------------- _chalceus_ (_phyllobates_) peters, 1873 _eleutherodactylus chalceus_ _festae_ (_paludicola_) peracca, 1904 _niceforonia festae_ _hylaeformis_ (_phyllobates_) cope, 1875 _eleutherodactylus hylaeformis_ _palmatus_ (_phyllobates_) werner, 1899 _colostethus palmatus_ _ridens_ (_phyllobates_) cope, 1866 _eleutherodactylus ridens_ _simonsii_ (_paludicola_) boulenger, 1900 _niceforonia simonsii_ ========================================================================== table 3--nominal species of _syrrhophus_ (_sensu strictu_) and the name used herein. ======================================================================= original combination current combination ---------------------------------------------------------------------- _campi_, _syrrhophus_ _cystignathoides campi_ _cholorum_, _syrrhophus leprus_ _leprus_ _cystigathoides_, _phyllobates_ _cystignathoides cystignathoides_ _dennisi_, _syrrhophus_ _dennisi_ new species _gaigeae_, _syrrhophus_ _guttilatus_ _guttilatus_, _malachylodes_ _guttilatus_ _interorbitalis_, _syrrhophus_ _interorbitalis_ _latodactylus_, _syrrhophus_ _longipes_ _leprus_, _syrrhophus_ _leprus_ _longipes_, _batrachyla_ _longipes_ _macrotympanum_, _tomodactylus_ _verrucipes_ _marnockii_, _syrrhophus_ _marnockii_ _modestus_, _syrrhophus_ _modestus_ _nebulosus_, _syrrhophus_ _pipilans nebulosus_ _nivocolimae_, _syrrhophus_ _nivocolimae_ _pallidus_, _syrrhophus modestus_ _pallidus_ _petrophilus_, _syrrhophus_ _guttilatus_ _pipilans_, _syrrhophus_ _pipilans pipilans_ _rubrimaculatus_, _syrrhophus_ _rubrimaculatus_ _smithi_, _syrrhophus_ _guttilatus_ _teretistes_, _syrrhophus_ _teretistes_ _verrucipes_, _syrrhophus_ _verrucipes_ _verruculatus_, _phyllobates_ _nomen dubium_ ======================================================================= in the course of preparing an account of the species of _eleutherodactylus_ occurring in méxico and northern central america, it became necessary to reëxamine the status of the genus _syrrhophus_ and its nominal species. it soon became evident that there were more names than species, that some previously regarded species were geographic variants, and that the eastern and western groups (complexes here) were artificial divisions of the genus. i conclude that there are seven species (one polytypic) of _syrrhophus_ in eastern méxico, texas, and el petén of guatemala, and seven species (one polytypic) in western méxico. the current status of each of the 23 names correctly assigned to the genus is presented in table 3. the fourteen species recognized by me are placed in five species groups. two of these groups are presently placed in the western complex (_modestus_ and _pipilans_ groups) and three in the eastern complex (_leprus_, _longipes_ and _marnockii_ groups). the two complexes do not correspond exactly with the eastern and western groups of smith and taylor (1948), firschein (1954), and duellman (1958) since _s. rubrimaculatus_ is now associated with the eastern _leprus_ group. the definitions and contents of the five species groups are as follows: _leprus_ group: digital pads not or only slightly expanded, rounded in outline; first finger longer or shorter than second; snout acuminate or subacuminate, not rounded; outer metatarsal tubercle conical; digits lacking distinct lateral fringes. content: _cystignathoides_, _leprus_ and _rubrimaculatus_. _longipes_ group: digital pads widely expanded, triangular in outline; first finger shorter than second; snout acuminate; outer metatarsal tubercle not conical; digits bearing lateral fringes. content: _dennisi_ and _longipes_. _marnockii_ group: digital pads expanded, rounded to truncate in outline; first finger equal in length to second or slightly shorter; snout rounded; outer metatarsal tubercle not conical; digits lacking lateral fringes; generally stout-bodied frogs. content: _guttilatus_, _marnockii_, and _verrucipes_. _modestus_ group: digital pads expanded, truncate in outline; first and second fingers subequal in length, first usually slightly shorter than second; snout subacuminate; inner metatarsal tubercle twice as large (or larger) as outer metatarsal tubercle; digits bearing poorly-defined lateral fringes. content: _interorbitalis_, _modestus_, _nivocolimae_, _pallidus_, and _teretistes_. _pipilans_ group: digital pads not or only slightly expanded, truncate in outline; first finger equal in length to second; snout subacuminate; metatarsal tubercles subequal in size; digits lacking lateral fringes. content: _pipilans_. _acknowledgments._--for loan of specimens, i am indebted to richard j. baldauf, texas a & m university (tcwc); w. frank blair, university of texas (tnhc); charles m. bogert and richard g. zweifel, american museum of natural history (amnh); james e. böhlke and edmond v. malnate, academy of natural sciences of philadelphia (ansp); robert f. inger and hymen marx, field museum of natural history (fmnh); ernest a. liner (eal); michael ovchynnyk, michigan state university collection (msu); james a. peters, united states national museum (usnm); douglas a. rossman, louisiana state university museum of zoology (lsumz); hobart m. smith, university of illinois museum of natural history (uimnh); charles f. walker, university of michigan museum of zoology (ummz); and john w. wright, los angeles county museum (lacm). specimens in the collection at the university of kansas museum of natural history are identified as ku. the abbreviations eht-hms refer to the edward h. taylor-hobart m. smith collection and fas to the frederick a. shannon collection. the type-specimens from these collections are now in the field museum of natural history and the university of illinois museum of natural history. i have profited from discussions concerning this problem with several persons, most notably william e. duellman, hobart m. smith, edward h. taylor and charles f. walker. nevertheless, the ideas and conclusions presented here should not be construed as necessarily reflecting their opinions. david m. dennis executed all of the figures, and my wife, marsha, typed the manuscript. _materials and methods._--in the course of this study, 1003 specimens of the genus were examined. the holotypes of 21 of the 23 nominal species are extant; i have examined 19 of these. nine measurements were taken, and five ratios computed for each of 338 specimens. females are available for all species but one; thus, measurements were taken on individuals of both sexes. analysis of characters _size and proportions._--frogs of this genus range in size from 16 to 40 mm. in snout-vent length. five species are relatively small: _s. cystignathoides_, _modestus_, _nivocolimae_, _pallidus_ and _rubrimaculatus_; one, _s. longipes_, is relatively large, and the remaining eight species are intermediate in size (22-30 mm.). males are generally smaller than females and have proportionately longer heads and usually larger tympani. no significant differences were found among proportions, except that _s. longipes_ has a larger tympanum/eye ratio than any other species. frogs in the _syrrhophus marnockii_ group tend to have shorter shanks and feet, thereby giving those species a more stocky appearance. however, the differences are not significant. a summary of the data on size and proportions for the frogs of the genus _syrrhophus_ is given in tables 4, 5, and 6. _hands and feet._--taylor and smith (1945), smith and taylor (1948), firschein (1954) and duellman (1958) discussed the value of the palmar tubercles in identifying frogs of this genus. the eastern complex in general has a well-developed outer palmar tubercle (fig. 1) in distinction to the western complex in which the outer palmar tubercle is reduced or absent (fig. 2). dixon and webb (1966) imply that the outer palmar tubercle is rarely absent but is usually smaller than the first supernumerary tubercle of the fourth finger. my study of the western species demonstrates that the outer palmar tubercle is indeed usually present and smaller than the first supernumerary tubercle. differences in interpretation of the terms "unexpanded" and "narrow," as well as differences in techniques of preservation, have led to confusion of the reported digital shapes in various species. constant specific differences are evident in the hands (fig. 1). except in the cases of excessive uptake of fluids, all species have a terminal transverse groove at the tip of each digit. taylor (1940b) stated that _s. smithi_ lacked grooves, but examination of the holotype reveals faint grooves at the tops of the digits. _syrrhophus guttilatus_, _leprus_, _pipilans_, and _verrucipes_ lack lateral fringes on the fingers. lateral fringes are well developed in the _longipes_ and _modestus_ groups but poorly defined or absent in the other members of the genus. the digital pads of the frogs of the _longipes_ group are much broader than those of the other species and are narrowest in the frogs of the _leprus_ group. supernumerary tubercles are present on the palmar surfaces of all species of the genus. table 4--size and proportions in the frogs of the _syrrhophus leprus_ group. a: _cystignathoides campi_ b: _c. cystignathoides_ c: _leprus_ d: _rubrimaculatus_ ========================================================================== snout-vent tibia head tympanum/ eyelid/ length length/ width/ eye interorbital species sex n (svl) svl svl ------------------------------------------------------------------------- a [m] 33 16.3-23.5 41.3-49.6 34.0-40.1 43.7-66.5 43.2-89.6 (45.8) (37.0) (56.2) (61.5) [f] 12 16.0-25.8 41.5-51.0 33.0-38.0 42.8-60.0 48.2-69.2 (45.8) (35.0) (51.2) (60.1) b [m] 15 16.8-22.1 45.1-50.4 33.2-40.7 44.3-68.7 44.6-65.4 (47.3) (37.8) (54.8) (60.0) [f] 6 19.6-24.2 46.4-50.0 34.1-38.1 43.3-56.5 53.2-65.4 (47.6) (36.2) (46.9) (59.2) c [m] 14 20.6-26.4 42.3-52.3 35.0-40.3 47.5-62.5 58.2-72.5 (46.8) (37.4) (56.5) (67.3) [f] 15 22.1-29.2 43.4-53.3 32.6-38.9 38.6-57.9 50.2-86.9 (47.1) (35.8) (47.1) (68.1) d [m] 12 18.2-23.5 40.4-46.2 31.8-35.5 35.5-46.5 65.1-78.5 (43.4) (33.8) (41.7) (71.7) ========================================================================== table 5--size and proportions in the frogs of the _syrrhophus longipes_ and _s. marnockii_ groups. a: _dennisi_ b: _longipes_ c: _guttilatus_ d: _marnockii_ e: _verrucipes_ ========================================================================== snout-vent tibia head tympanum/ eyelid/ length length/ width/ eye interorbital species sex n (svl) svl svl ------------------------------------------------------------------------- a [m] 16 22.8-28.4 43.9-49.7 35.3-41.2 53.9-64.2 55.3-74.0 (47.4) (38.8) (58.9) (65.1) [f] 10 25.9-32.0 46.3-50.8 35.6-40.3 50.6-58.7 58.1-70.9 (48.2) (37.7) (54.9) (63.6) b [m] 22 22.1-33.2 45.8-51.7 38.7-44.4 61.1-87.2 61.5-83.0 (48.4) (41.8) (72.0) (72.0) [f] 19 26.8-39.6 44.3-51.0 36.3-40.8 49.5-72.1 55.3-85.9 (47.2) (39.1) (59.5) (67.9) c [m] 19 20.6-29.0 41.2-48.1 36.9-44.9 55.1-75.7 53.3-79.5 (44.5) (40.6) (64.1) (66.0) [f] 5 25.7-31.0 41.4-46.8 35.9-42.3 47.6-61.7 62.3-79.8 (43.6) (38.5) (54.0) (72.9) d [m] 14 18.4-28.9 42.3-47.2 36.1-43.0 47.2-68.3 51.6-74.4 (44.1) (39.6) (61.2) (66.3) [f] 29 20.4-35.4 38.7-46.4 35.9-41.3 45.8-73.3 52.1-70.5 (42.7) (38.2) (60.3) (60.7) e [m] 29 17.5-29.2 42.7-49.5 36.2-42.4 56.1-82.2 56.8-82.8 (46.3) (39.1) (67.8) (70.4) [f] 6 26.5-31.7 42.4-47.7 36.0-38.1 45.8-57.8 61.0-77.9 (44.6) (37.0) (53.9) (69.0) ========================================================================== table 6--size and proportions in the frogs of the _syrrhophus pipilans_ and _s. modestus_ groups. a: _pipilans nebulosus_ b: _pipilans pipilans_ c: _modestus_ d: _pallidus_ e: _teretistes_ f: _nivocolimae_ g: _interorbitalis_ ========================================================================== snout-vent tibia head tympanum/ eyelid/ length length/ width/ eye interorbital species sex n (svl) svl svl ------------------------------------------------------------------------- a [m] 17 22.9-28.5 38.1-42.0 34.4-37.2 36.6-47.8 56.1-82.4 (40.0) (35.4) (43.6) (68.2) [f] 3 21.1-22.7 42.1-44.5 33.2-35.8 36.6-47.6 64.3-65.4 b [m] 18 22.6-27.8 37.9-44.0 32.2-36.5 38.0-54.0 56.1-79.5 (41.4) (33.0) (46.2) (67.3) [f] 1 29.4 38.4 32.5 44.6 55.0 c [m] 8 15.8-20.1 38.5-42.6 32.1-38.1 26.8-39.3 57.0-86.9 (40.6) (34.2) (31.5) (69.1) [f] 1 18.5 44.2 36.0 24.0 52.1 d [m] 6 17.9-19.3 41.0-44.9 32.6-36.2 27.0-35.6 59.4-67.7 (43.4) (35.2) (30.9) (65.2) e [m] 18 19.2-23.2 41.5-45.3 32.5-36.4 28.6-43.8 51.2-75.0 (43.7) (34.0) (33.7) (62.2) [f] 1 24.8 41.8 30.8 37.9 60.5 f [m] 15 18.9-21.1 42.2-48.6 30.9-37.1 30.0-39.3 42.6-69.1 (45.0) (33.7) (34.7) (55.0) [f] 1 24.1 40.9 33.5 27.6 56.5 g [m] 1 25.6 43.0 --- 39.4 57.6 [f] 9 20.2-26.7 39.9-47.1 32.6-39.3 29.1-41.2 58.2-76.9 (43.2) (35.8) (36.4) (69.2) ========================================================================== [illustration: fig. 1: palmar views of hands of six species of the eastern complex of _syrrhophus_. (a) _verrucipes_ (uimnh 15995), (b) _rubrimaculatus_ (ku 58911), (c) _dennisi_ sp. nov. (holotype, ummz 101121), (d) _guttilatus_ (uimnh 55520), (e) _marnockii_ (tcwc 4782), and (f) _longipes_ (tcwc 12179). all ×6.5.] [illustration: fig. 2: palmar views of hands of two species of the western complex of _syrrhophus_. _pipilans_ (left, ku 58908, ×6) and _teretistes_ (center, ku 75269, and right, ku 75263, respectively, ×9).] in _s. cystignathoides_ and _leprus_, the first finger is longer than the second, and the first two fingers are equal in length in _guttilatus_ and _marnockii_. in the other species the first finger is shorter than the second. supernumerary tubercles are well developed on the plantar surfaces in all species, except _s. guttilatus_, in which they are poorly defined (fig. 3). the relative sizes of the metatarsal tubercles has been used in the classification of the species and species groups of _syrrhophus_. the metatarsal tubercles are similar in all species of the eastern complex (including _rubrimaculatus_); the outer tubercle is always about one-half the size of the ovoid inner metatarsal tubercle. in the _leprus_ group the outer tubercle is conical and compressed. the metatarsal tubercles of _pipilans_ are about the same size, or the outer is slightly smaller than the inner. in the _modestus_ group the outer metatarsal tubercle is about one-third the size of the inner. all species, except _guttilatus_, have well-defined to poorly defined lateral fringes on the toes. all species have expanded toe pads. the fifth toe is usually shorter than the third, but the second is equal in length to the fifth in some specimens of _s. cystignathoides_ and _s. marnockii_. _syrrhophus nivocolimae_ is the only species with tubercles along the outer edge of the tarsus; this is merely a reflection of the highly tuberculate nature of the skin in this species. _skin texture._--the skin of the dorsum is smooth or very weakly pustular in all species of the genus except _nivocolimae_ and _verrucipes_. the dorsal surfaces of _nivocolimae_ are warty; in _verrucipes_ the skin is pustular. the skin of the venter is areolate in _cystignathoides cystignathoides_, _dennisi_ and _verrucipes_ but is smooth in all other species of the genus. [illustration: fig. 3: plantar views of feet of four species of the eastern complex of _syrrhophus_. (a) _guttilatus_ (uimnh 55519, ×6), (b) _leprus_ (uimnh 42726, ×6), (c) _verrucipes_ (uimnh 15995, ×6), and (d) _longipes_ (tcwc 12179, ×4.6).] _color pattern._--as is evident in the diagnoses, the color patterns of given populations have been regarded as useful in separating the species and subspecies. duellman (1958) suggested that the coloration, with the exception of _modestus_, was a dark ground color with pale markings. it is a moot point whether the frogs have light spots on a dark background or have a light background with an extensive reticulate dark pattern. the venters are gray or white, and the vocal sac is nearly black in some species. interorbital dark bars or triangles are absent in only two species of the eastern complex, _cystignathoides campi_ and _marnockii_; the latter lacks a supratympanic stripe, which is present in the other members of the eastern complex. _syrrhophus interorbitalis_ and _nivocolimae_ have light interorbital bars; these bars occur in only one other population of the genus (_s. c. cystignathoides_). bars on the thighs are ill defined or absent in the members of the _marnockii_ and part of the _modestus_ groups. the color in life is noted in the species accounts. _voice._--the voices of all _syrrhophus_ can be described as a single short chirp or peep; without audiospectrographic analyses the significance of the differences between a chirp, peep, or short whistle cannot be appreciated. martin (1958) and wright and wright (1949) reported multi-noted calls, and one collector of _s. verrucipes_ noted the frog "trilled." fouquette (1960) presented analyses of two species (_marnockii_ and _pipilans nebulosus_). the voices were very similar; both frogs were reported to "trill" and "chirp." systematic account the genus _syrrhophus_ has been defined (lynch, 1968) and limited to the group of species occurring in guatemala, méxico and the united states. the closest relatives of _syrrhophus_ are the frogs of the genus _tomodactylus_ (dixon, 1957; firschein, 1954). lynch (1968) implied there were no osteological bases for the separation of _eleutherodactylus_, _syrrhophus_, and _tomodactylus_. at that time, i believed such to be the case and derived _syrrhophus_ and _tomodactylus_ from the _rhodopis_ complex of _eleutherodactylus_, with which they share terrestrial habits and relatively short limbs. in the _rhodopis_ complex there is a tendency for the loss of the outer palmar tubercle, a not uncommon condition in _syrrhophus_ and _tomodactylus_. however, the skulls of _syrrhophus_ and _tomodactylus_ show departures from the pattern observed in the middle american _eleutherodactylus_, as well as many of those species in western south america. baldauf and tanzer (1965) reported that the frontoparietals and prootics were fused in _syrrhophus marnockii_ and that the prootics and exoccipitals appeared to be one bone (otoccipital). the otoccipital is not uncommon in eleutherodactyline frogs, but the fusion of the frontoparietals with the prootics (regardless of the fusion of the latter with the exoccipital) is uncommon in the family. i have found the frontoparietal-prootic fusion only in _syrrhophus_ (all species), _tomodactylus_ (all species), and _eleutherodactylus_ (west indies species). none of the middle american _eleutherodactylus_ has the two bones fused. examination of the character is difficult in dried skeletal preparations. cleared and stained or macerated preparations are satisfactory for checking this character. thus, in addition to the presence of numerous plantar supernumerary tubercles in the frogs of the genera _syrrhophus_ and _tomodactylus_, these two genera can be separated from other middle american eleutherodactylines by the fusion of the frontoparietals and prootics. this character not only further strengthens the argument that the two genera are closely related but poses a problem of zoogeographic analysis of the distribution of the character, which will be discussed fully elsewhere. key to the species of the frog genus _syrrhophus_ 1. three large, well-developed palmar tubercles 2 two large palmar tubercles; outer (third) palmar tubercle reduced in size or absent 9 2. digital pads more than twice (usually three or more) times width of digit 3 digital pads less than twice width of digit 4 3. males having vocal slits; dorsum vermiculate; diameter of tympanum in males about one-half diameter of eye _s. dennisi_ males lacking vocal slits; dorsum flecked, spotted, or blotched; diameter of tympanum in male about three-fourths that of eye _s. longipes_ 4. first finger longer than second 5 first finger shorter than or equal to second 7 5. venter smooth; dorsum spotted or vermiculate _s. leprus_ venter areolate, or if smooth, dorsum flecked and interorbital bar lacking 6 6. venter areolate; interorbital bar present; ground color yellowish _s. cystignathoides cystignathoides_ venter smooth; interorbital bar absent; ground color brown _s. cystignathoides campi_ 7. first finger shorter than second; digital tips only slightly dilated; green in life with darker green spots _s. verrucipes_ first finger equal to second; digital tips slightly to moderately expanded 8 8. dorsum vermiculate; interorbital bar present; ground color cream to brown in life _s. guttilatus_ dorsum punctate or flecked; interorbital bar absent; ground color green in life _s. marnockii_ 9. dorsum dark with pale (red in life) spots; digital pads not expanded _s. rubrimaculatus_ dorsum pale with dark markings and digital pads slightly to widely expanded 10 10. digital tips not widely expanded; tympanum well-defined; outer metatarsal tubercle more than one-half size of inner 11 digital tips widely expanded, truncate in outline; tympanum poorly defined; outer metatarsal tubercle less than one-half size of inner 12 11. dorsum dark brown with large light spots or blotches; tympanum/eye ratio usually greater than 43 percent _s. pipilans pipilans_ dorsum dark brown with small light spots; tympanum/eye ratio less than 48 percent _s. pipilans nebulosus_ 12. light interorbital bar present 13 light interorbital bar absent 14 13. adults small, less than 22 mm. snout-vent length with a broad mid-dorsal stripe; dark bands on shank narrower than light interspaces _s. nivocolimae_ adults larger, more than 22 mm. snout-vent length; dorsum vermiculate; dark bands on shank broader than light interspaces _s. interorbitalis_ 14. dorsum spotted with discrete black spots; pattern definite _s. modestus_ dorsum reticulate or vermiculate, pattern poorly defined 15 15. adults small, less than 21 mm. snout-vent length; upper arm not banded _s. pallidus_ adults larger, usually greater than 21 mm. snout-vent length; upper arm banded _s. teretistes_ species accounts the following accounts do not include complete descriptions of each taxon, because a more than adequate number of descriptions is available in the recent (1940-1966) literature. an abbreviated synonymy, in which are listed all combinations and emendations of names and significant contributions to our knowledge of the taxon, is given for each. for each species and subspecies the following are given: descriptive diagnosis, statement of range, remarks on taxonomy, list of specimens examined, illustration of color pattern, and distribution map. =syrrhophus cystignathoides= (cope) _phyllobates cystignathoides_ cope, 1877:89-90 [syntypes.--originally usnm 32402-32409, (32405 now in mcz) from potrero, near córdoba, veracruz, méxico, francis sumichrast collector.] _diagnosis._--adults small, males 16.0 to 23.5 mm. in snout-vent length, females 16.0-25.8 mm. in snout-vent length; vocal slits present in males; finger tips slightly expanded; first finger longer than second; outer metatarsal tubercle one-half size of inner, conical, compressed; skin of dorsum weakly pustular, that of venter smooth to areolate; tympanum 44 to 69 per cent diameter of eye (mean 55.5 per cent); ground color yellow to brown in life with brown to black fleckings on dorsum and flanks; limbs banded; interorbital bar present or not. _remarks._--two geographic races (subspecies) are herein recognized; previously these were held by various authors to be species (_campi_ and _cystignathoides_). intergradation occurs in southern tamaulipas and eastern san luis potosí, méxico. the two subspecies can be distinguished on the basis of color pattern and the condition of the skin of the venter. _distribution._--low to moderate elevations from the río grande embayment to central veracruz, méxico (fig. 5). =syrrhophus cystignathoides campi= stejneger, new combination _syrrhophus campi_ stejneger, 1915:131-32. [holotype.--usnm 52290, from brownsville, cameron co., texas; r. d. camp collector, march 31, 1915]. smith and taylor, 1948:52. martin, 1958:50. _diagnosis._--venter smooth; usually no interorbital light and dark bars present; ground color brown in life (fig. 4a). _remarks._--martin (1958) was the first author to point out that _s. campi_ was probably a subspecies of the more southern _s. cystignathoides_. various references in the literature might lead one to believe that the two were sympatric over much of northeastern méxico; this error was created by the use of a single character (condition of the skin of the venter) to characterize the two populations. specimens from southern texas have a smooth venter, lack interorbital bars and have, in general, a brown ground color, whereas specimens from central veracruz have an areolate venter, interorbital light and dark bars and a yellow ground color. in southern tamaulipas and eastern san luis potosí, these characters vary discordantly, thereby strongly suggesting that the two populations intergrade. both populations agree in other morphological characters; therefore, they are here treated as geographic variants. _etymology._--named for the collector of the type specimens, mr. r. d. camp of brownsville, texas. _distribution._--lower río grande embayment in texas to central nuevo león and tamaulipas, méxico. intergrades are known from southern tamaulipas and adjacent san luis potosí, méxico (fig. 5). _specimens examined._--(113) texas, cameron co.: mcz 10277-85, 10286 (10); brownsville, amnh 3215, 3218-20, 3221 (3), 5376, 62117, fmnh 105336, ku 8135-39, mcz 3738-42, 3743 (10), tcwc 5908, 7139, tnhc 92-94, 20909, ummz 51760, 54031 (5), usnm 52290 (holotype); 22 mi. se brownsville, tnmc 14223; 8 mi. sw brownsville, ummz 101127 (3); harlingen, amnh 62118, ummz 105200-205, 105206 (5), 105207 (4). _hidalgo co._: bentsen-río grande state park, ummz 114378; 6 mi. s mcallen, tnhc 7136-39; santa ana refuge, tcwc 13495-96; weslaco, tcwc 17658-60. mexico, _nuevo león_: salto cola de caballo, amnh 57953-54, fmnh 30644-45, 37169-70; monterrey, uimnh 13324; 40 km. se monterrey, uimnh 3686. _tamaulipas_: 80 km. matamoros, fmnh 27150 (13). intergrades [_s. c. cystignathoides_ × _s. c. campi_ (88)] méxico, _san luis potosí_: 5 km. e ciudad del maiz, ummz 106435; 16 km. w naranjo, fmnh 104584; salto de agua, 34 km. wsw antigua morelos, tcwc 6980. _tamaulipas_: 5 km. w acuña, 1060 m., ummz 101172, 101173 (16), 101174-76, 101177 (6); 14.5 km. nnw chamal, 430 m., ummz 111337 (2); 20 km. nnw chamal, 700 m., ummz 111338 (11); 8 km. n gómez farías, 450 m., ummz 101165; 8 km. ne gómez farías, pano ayuctle, ummz 102264, 102924 (6); 8 km. nw gómez farías, 1060 m., lsumz 11084, ummz 101199, 102928 (5), 102929-32, 110124 (3); río guayala, near magiscatzin, mcz 24138-42, 85071-81, ummz 88242 (2); magiscatzin, tcwc 6981; las yucas, north of aldama, mcz 29665-68; 16 km. ne zamorina, ummz 101124. [illustration: fig. 4: _syrrhophus cystignathoides campi_ (left, tcwc 13490) and _s. c. cystignathoides_ (right, ku 105500). dorsal views ×2, sides of heads ×3.] =syrrhophus cystignathoides cystignathoides= (cope), new combination _phyllobates cystignathoides_ cope, 1877:89-90 [syntypes.--usnm 32402-32409, from potrero, near córdoba, veracruz, méxico, collected by francis sumichrast]. boulenger, 1882:196. _syrrhophus cystignathoides_: cope, 1879:268. kellogg, 1932: 126-27. taylor and smith, 1945: 582-83. smith and taylor, 1948:50. martin, 1958:49. _syrrhaphus cystignathoides_: günther, 1900:218. _syrraphus cystignathoides_: díaz de león, 1904:10. _syrrhopus cystignathoides_: barbour and loveridge, 1946:170. [illustration: fig. 5: distribution of _syrrhophus cystignathoides campi_ (solid symbols) and the nominate subspecies (open symbols).] _diagnosis._--venter areolate; interorbital light and dark bars present; ground color yellow to brownish-yellow in life (fig. 4b). _remarks._--firschein (1954) briefly considered the status of peters' (1871) _phyllobates verruculatus_ and noted that if it was a _syrrhophus_ it would probably be referrable to _s. cystignathoides_. peters' (1871) original description corresponds well with _s. cystignathoides_, and the type-locality ("huanusco" = huatusco) is within the range of that species. firschein (1954) expressed doubt that _verruculatus_ was a _syrrhophus_, because peters placed it in another genus. however, peters described _verruculatus_ a decade before cope diagnosed the genus syrrhophus. most frogs now called _syrrhophus_, plus a number of lower central american frogs now placed in a variety of genera were placed in _phyllobates_ by boulenger, cope, and peters. the types of _phyllobates verruculatus_ were destroyed during world war ii (günther peters, _in litt._); the specimens subsequently assigned to the taxon by kellogg (1932) are _syrrhophus cystignathoides_. because the type specimens are lost and because the name antedates the more established name, _cystignathoides_, i favor retaining _phyllobates verruculatus_ peters as a _nomen dubium_. smith and taylor (1948) reported _s. verruculatus_ from tianguistengo, hidalgo, méxico. these specimens are examples of _verrucipes_. smith (1947) reported a specimen of _verruculatus_ from san lorenzo, veracruz. firschein (1954) referred it to _cystignathoides_, and duellman (1960) concluded that both authors were in error and that the specimen (usnm 123530) was a _leprus_. _etymology._--the trivial name is the diminutive of _cystignathus_, a once-used generic name for several leptodactylid frogs. _distribution._--low and moderate elevations in the foothills along the sierra madre oriental from eastern san luis potosí to central veracruz, méxico (fig. 5). _specimens examined._--(130), méxico, _puebla_: necaxa, ummz 69519-20. _san luis potosí_: 5 km. w aguismón, lsumz 4962-63; along río axtla, road to xilitla, ummz 105500; tamazunchale, uimnh 3199; 6.5 km. n tamazunchale, ummz 104039; 8 km. n tamazunchale, ummz 119490. _veracruz_: coatepec, 1210 m., fmnh 704966-67; 11 km. se coatepec, 850 m., fmnh 70468-70; below córdoba, fmnh 104588, uimnh 13321; cuautlapam, 1000 m., fmnh 106477-80, ku 100364, uimnh 58200-03, ummz 105392; fortín de las flores, uimnh 13322, 13339; 1.6 km. n fortín de las flores, uimnh 42799-808, ummz 105389; 3.2 km. n fortín de las flores, uimnh 26633-35; 4.8 km. n fortín de las flores, uimnh 71967-68; 3.2 km. w fortín de las flores (barranca metlac), 910 m., uimnh 49294-95, ummz 115444-46, 118221, 119893 (2); huatusco, ku 100363; jalapa, 1400 m., fmnh 70440, 70443-51, 70454-65; 16 km. ne jalapa, 1300 m., fmnh 70452-53; 8 km. e jalapa, uimnh 13338; 9.5 km. s jalapa, ummz 122083 (2); mirador, ku 23967; paraja nuevo, el suchil, ummz 85490 (7), 85491 (2), 90315; la passa, uimnh 49293, 49297; 1 km. e plan del río, 240 m., ummz 102067 (2); potrero viejo, fmnh 104583, 104586, 105326-27, ku 26789, 100357-62, uimnh 13323, 13340-43; usnm 32402 (lectotype), 32403-04, 32406-09; 9.6 km. s santa rosa, tcwc 12785; 24 km. ne tezuitlán (puebla), ummz 105388; teocelo, fmnh 70437-38, ku 26080, 26790; 3.2 km. n teocelo, fmnh 70439, 70441-42; 9.6 km. nw tihuatlán, uimnh 3684-85; 15 km. ene tlacotepec, ku 23966; 26 km. nw tuxpan, ummz 126419. =syrrhophus leprus= cope _syrrhophus leprus_ cope, 1879:268-69 [holotype.--usnm 10040, from santa efigena, oaxaca, méxico, francis sumichrast collector]. kellogg, 1932:124-5, 128. taylor and smith, 1945:582. smith and taylor, 1948:50-51. duellman, 1958:8, pl. 1, fig. 2; 1960:56-57. gorham, 1966:165. _syrrhaphus leprus_: günther, 1900:217. _syrrhophus leprus leprus_: neill, 1965:85-86. _syrrhophus leprus cholorum_ neill, 1965:85-86 [holotype.--wilfred t. neill collection 1525, from 3.9 mi. n san antonio, toledo district, british honduras, collected october 28, 1959, by r. a. allen, t. c. allen, and w. t. neill]. _diagnosis._--medium-sized frogs, males 20.5-26.5 mm. in snout-vent, females 22.0-29.3 mm. in snout-vent length; vocal slits present in males; tips of fingers dilated slightly; first finger longer than second; inner metatarsal tubercle twice size of small, conical outer metatarsal tubercle; skin of dorsum pustular, that of venter smooth; snout subacuminate; diameter of tympanum 47.5-62.5 per cent of eye in males, 38.6-57.9 per cent in females; dorsum yellowish-green with chocolate brown blotches or spots forming reticulations in most specimens; venter white to gray; flanks brown, spotted with white or not; limbs banded; interorbital bar obscured by dorsal pattern. [illustration: fig. 6: dorsal views of _syrrhophus leprus_ showing variation in dorsal pattern (left, ummz 121244, ×2; right, ku 26106, ×1.7). side of head (uimnh 42726, ×7).] [illustration: fig. 7: distribution of three species of eastern complex _syrrhophus_: _leprus_ (circles), _rubrimaculatus_ (triangles), and _verrucipes_ (squares).] _remarks._--my distribution map (fig. 7) differs somewhat from that of duellman (1958), who was unaware of specimens reported by taylor and smith (1945) from central veracruz, méxico. duellman (1958, 1960) regarded _s. leprus_ as having a gray venter. neill (1965) characterized his new subspecies on the basis of white venter and spots on the dorsum. some specimens from throughout the range have only small round spots, instead of vermiculations (fig. 6). the gray ventral coloration is largely restricted to the population in los tuxtlas, veracruz, but only about 80 per cent of the specimens from the los tuxtlas have gray venters, whereas specimens from guatemala, oaxaca, tabasco, and central veracruz, méxico, have white venters (rarely gray). since the specimens from british honduras are not distinct from specimens throughout most of the range, there is no reason to recognize them as a subspecies. _etymology._--greek, _lepra_, leprosy, in reference to the mottled color pattern. _distribution._--discontinuous; central veracruz to british honduras to low elevations in the foothills of the sierra madre oriental, los tuxtlas, sierra madre de chiapas (isthmus of tehuantepec (fig. 7)). _specimens examined._--(84). guatemala, _alta verapaz_: chinajá, ku 55961-62. _el petén_: 15 km. nw chinajá, ku 55963; piedras negras, usnm 114085-92; tikal, ummz 117035; uaxactún, amnh 55121-22. méxico, _oaxaca_: cerro san pedro del isthmo, uimnh 35510; finca la gloria, usnm 114093; 30.5 km. n matías romero, uimnh 39459, 71969; santa efigenia, usnm 10040 (holotype). _tabasco_: teapa, ummz 113799-800; 13.5 km. w teapa, ummz 120253. _veracruz_: 27.5 km. n acayucan, uimnh 42726; atoyac, uimnh 13331, 49296; 3.2 km. n catemaco, uimnh 71976-77; coyame, uimnh 38995, 38998, 40342; dos amates, tcwc 21211; fortín de las flores, fmnh 113751, 113753; paraja nuevo, el suchil, ummz 90315; potrero viejo, fmnh 113743-50, 126114-18, ku 26104-06, uimnh 13332-37, ummz 88837; san andrés tuxtla, uimnh 27123-31, 28611, 71975, ummz 115450 (5); san lorenzo, usnm 123530; 4.5 km. nw santiago tuxtla, jdl 992 (skeleton), uimnh 27122; 32 km. s sayula, eal 1696; tepalapan, 1.6 km. s catemaco, ummz 118222 (2); volcán san martín, south slope, ummz 118223; volcán san martín, rancho el tular, uimnh 35399-400, 40340-41. =syrrhophus rubrimaculatus= taylor and smith _syrrhophus rubrimaculatus_ taylor and smith, 1945:583-85 [holotype.--usnm 114070, from la esperanza, near escuintla, chiapas, méxico, collected may 13, 1940, by h. m. and r. smith]. duellman, 1958:1-4, 7, 12, 14. gorham, 1966:167. _syrrhophus rubrimaculata_: smith and taylor, 1948:48-49. [illustration: fig. 8: _syrrhophus rubrimaculatus_ (upper right, ku 58911, ×1.6; lower right, ku 58910, ×4) and _s. verrucipes_ (upper left, uimnh 15995, ×1.6; lower left, uimnh 15989, ×3.7).] _diagnosis._--small frogs, males 18.2-23.5 mm. snout-vent, females 19.0-22.5 mm. snout-vent length (small sample); vocal slits in males; digital tips scarcely expanded (fig. 1); first finger shorter than second; outer palmar tubercle reduced in size; inner metatarsal tubercle elongate, twice the size of small, conical outer metatarsal tubercle; diameter of tympanum 35.5-46.5 per cent that of eye in both sexes; dorsum brown with small pale spots (red in life); venter gray. _remarks._--previous authors who treated _syrrhophus_ placed this species in the western complex, because it occurs on the pacific versant and has a reduced outer palmar tubercle. duellman (1958) placed _rubrimaculatus_ apart from the other western species, because of its relatively unexpanded digital tips and coloration. the digital tips are like those in _leprus_, which _rubrimaculatus_ resembles. except for the reduction of the outer palmar tubercle, _rubrimaculatus_ could be a member of the _leprus_ group. _syrrhophus rubrimaculatus_ is probably best treated as a pacific derivative of the _leprus_ group, even though the palmar tubercles do not agree. the removal of _rubrimaculatus_ from the western complex results in a more homogeneous remainder and does not greatly increase the heterogeneity of the eastern complex. _etymology._--latin, meaning spotted with red; in reference to the colors in life. _distribution._--low to moderate elevations on the pacific versant of southeastern chiapas, méxico (fig. 7); probably extending into adjacent guatemala. _specimens examined._--(48) méxico, _chiapas_: escuintla, ummz 88283; 6 km. ne escuintla, ummz 87876-80; la esperanza, uimnh 13285, ummz 88496-97, usnm 114070 (holotype), 114054-69, 114072; monte cristo, ummz 88353; 1.3 km. n puerto madero, ku 58910-11; finca san jerónimo, 600-650 m., uimnh 55299-312, 55313-16 (cleared and stained). =syrrhophus guttilatus= (cope) _malachylodes guttilatus_ cope, 1879:264 [holotype.--usnm 9888, from guanajuato, guanajuato, méxico; collected in 1877 by alfredo duges]. _syrrhopus guttulatus_: boulenger, 1888:204-06. _syrrhaphus guttulatus_: günther, 1900:317. _syrraphus guttulatus_: díaz de león, 1904:11. _syrrhophus guttilatus_: nieden, 1923:399-400. kellogg, 1932:125, 127-28. smith and taylor, 1948:49, 51. firschein, 1954:52-54. gorham, 1966:164. _syrrhophus smithi_ taylor, 1940b:43-45, pl. 1 [holotype.--usnm 108594, from 15 mi. sw galeana, nuevo león, méxico, 1575 m.; collected on october 13, 1939, by hobart m. smith]. smith and taylor, 1948:49, 51. firschein, 1954:54-55. martin, 1958:50. gorham, 1966:167. _syrrhophus gaigeae_ schmidt and smith, 1944:80 [holotype.--fmnh 27361, from the basin, chisos mountains, brewster co., texas; collected on july 24, 1937, by walter l. necker]. _syrrhophus petrophilus_ firschein, 1954:50-52 [holotype.--uimnh 7807, from 5 km. sw san luis potosí, san luis potosí, méxico; collected on july 18, 1949, by david langebartel]. gorham, 1966:166. _syrrhophus marnocki_: milstead, mecham, and mcclintock, 1950:548 (in part). _diagnosis._--medium-sized frogs, males 20.6-29.0 mm. snout-vent, females 25.7-31.0 mm. snout-vent length; vocal slits in males; digital tips slightly expanded (fig. 1); first and second fingers equal; skin of dorsum smooth to moderately pustular, that of venter smooth; snout blunt; diameter of tympanum 55.1-75.7 per cent that of eye in males, 47.6-61.7 in females; dorsum and flanks cream to gray with light brown to black flecking and vermiculations; thighs usually not banded; interorbital bar present (fig. 8). [illustration: fig. 9: _syrrhophus guttilatus_ (upper left, uimnh 55519, ×1.4; lower left, uimnh 55519, ×2.3) and _s. marnockii_ (upper right, tcwc 9317, ×1.4; lower right, tcwc 13510, ×2.1).] _remarks._--cope (1879) distinguished _malachylodes_ from _syrrhophus_ on the basis of the presence of a frontoparietal fontanelle in the holotype of _guttilatus_. the holotype is a juvenile female and as is the case in the juveniles of nearly all leptodactylids, a frontoparietal fontanelle is present. firschein (1954) used the presence of the fontanelle to distinguish _guttilatus_ from his _petrophilus_. as is clearly evident from the length of the synonymy, i consider a number of currently used names to be synonymous with _guttilatus_. i have seen the holotypes of all four names and am unable to recognize more than a single species. the holotype of _petrophilus_ is a male, whereas that of _smithi_ is a female. the supposed differences are a reflection of sexual dimorphism in the size of the eye (table 5). the two holotypes, as well as those of _gaigeae_ and _malachylodes guttilatus_ agree in color pattern. schmidt and smith (1944) named _syrrhophus gaigeae_ from the chisos mountains of the big bend region of texas and compared it only with _s. marnockii_. milstead, mecham and mcclintock (1950) synonymized _gaigeae_ and _marnockii_ because they were unable to verify the characters wright and wright (1949) used to separate them. specimens from the big bend region differ from those of the edward and stockton plateaus in having a vermiculate pattern, an interorbital bar, and a supratympanic stripe. in these respects they agree with specimens from northern méxico. based on limited observations, the mexican population is yellowish to brownish in life whereas the central texas population is green in life. lacking evidence of genetic exchange, the two are held to be specifically distinct. nearly every specimen examined was infested with chiggers of the genus _hannemania_. the greatest concentrations are on the venter, in the groin, and on the thighs. many specimens have chiggers on the digits and tarsi. the same, or a related, chigger was found on many specimens of _syrrhophus marnockii_ and a few _s. verrucipes_, but on no other species of the genus. mr. willy wrenn told me that he has seen heavy infestations of _hannemania_ on _syrrhophus pallidus_. infestation by _hannemania_ probably reflects similar ecologies rather than close relationships. [illustration: fig. 10: distribution of _syrrhophus guttilatus_.] _etymology._--latin, _guttula_, meaning spotting or flecking, in reference to the color pattern. _distribution._--moderate to intermediate elevations (600 to 2000 m.) along the sierra madre oriental from the big bend region of texas to guanajuato, méxico (fig. 10). _specimens examined._--(32) texas, _brewster co._: juniper canyon, chisos mts., fmnh 27361 (holotype of _s. gaigeae_), 27360, 27362-63, mcz 15346, 27801, ummz 66080, 66082, 66085-91, usnm 76876; upper green gulch, tcwc 15943. méxico: _coahuila_: 8 km. s saltillo, uimnh 55518-21. _guanajuato_: guanajuato, usnm 9888 (holotype of _malachulodes guttilatus_); 8 km. e guanajuato, amnh 73425; cerro cubilete, amnh 73424. _nuevo león_: 3 km. s galeana, jdl 1215 (skeleton), uimnh 58204; 24 km. sw galeana. 1575 m., usnm 108594 (holotype of _syrrhophus smithi_). _san luis potosí_: 5 km. sw san luis potosí, uimnh 7807 (holotype of _s. petrophilus_). _tamaulipas_: 1.6 km. nw la joya de salas, 1530 m., ummz 110736 (4). =syrrhophus marnockii= cope _syrrhophus marnockii_ cope, 1878:253 [syntypes.--ansp 10765-68, from "near san antonio," bexar co., texas; collected by g. w. marnock]. _syrrhophus marnocki_: yarrow, 1882:24, 193. milstead, mecham, and mcclintock, 1950:550. _diagnosis._--medium-sized frogs, males 18.4-28.9 mm. snout-vent, females 20.4-35.4 mm. snout-vent length; vocal slits in males; digital tips widened (fig. 1); first and second fingers equal; skin of dorsum smooth to weakly pustular, that of venter smooth; snout blunt, rounded; diameter of tympanum 47.2-68.3 per cent that of eye in males, 45.8-73.3 in females; dorsum tan to light brown in preservative with rusty-brown flecks, venter white; ground color green in life; thighs banded; interorbital bar absent. _remarks._--specimens from the southern edge of the edwards plateau and the eastern edge of the stockton plateau have larger flecks on the back that tend to form a vermiculate pattern like that of _s. guttilatus_. the vermiculation is never well developed (see plate 38 in conant, 1958). most of the specimens from the edwards plateau have a punctate pattern (fig. 9). fossils are known from the sangamon interglacial deposits in foard and knox counties, texas (lynch, 1964; tihen, 1960). _etymology._--a patronym for the collector of the type specimens. _distribution._--the edwards plateau and the extreme eastern edge of the stockton plateau in texas (fig. 11). the fossil records lie some 200 miles to the north. two specimens (fmnh 103216-17) from brownsville, cameron co., texas, were formerly in the eht-hms collection (nos. 31348-49). data given in taylor's field catalogue (housed in the division of reptiles, field museum) are "brownsville, a. j. kirn collector, april 15, 1934." until verification by recently collected material is available, this record must be disregarded. _specimens examined._--(103) texas, _bandera co._: 10 mi. sw medina, tcwc 13508-10; 8 mi. w medina, ku 60243; 13 mi. w medina, ku 60242, tcwc 13506-07. _bexar co._: uimnh 34694; classen ranch, near san antonio. ummz 98891; helotes, eal 1560, mcz 11837 (2), ummz 64045, usnm 13635; 2 mi. n helotes, tcwc 9234-35; 3.5 mi. n helotes, lsumz 10363; 8 mi. n helotes, tcwc 1549, 4364; san antonio, fmnh 15553-56, tcwc 13497-99. _blanco co._: 8 mi. ne blanco, tcwc 4782. _comal co._: new braunfels, tcwc 13500-05; 5 mi. ne new braunfels, ummz 71016 (10). _hays co._: san marcos, amnh 22661-64, 32700, fmnh 15245-46, 26250, 26253-57, 37617, 37665, mcz 15649-50, 23268-69; 6 mi. sw san marcos, tcwc 5070-71, 7140, 9232-33, 9236, 9316-17, 9320. _kendall co._: 11 mi. e boerne, amnh 54660-61, 54662 (2); 10 mi. w boerne, ku 18441; kendalia, uimnh 21434. _kerr co._: kerr w. m. area, tcwc 15859; 40 mi. nw kerrville, tcwc 6555. _medina co._: uimnh 13287-88; 12 mi. n castroville, uimnh 21423; 14 mi. n castroville, uimnh 21424-25; 16 mi. n castroville, uimnh 21421-22; 17 mi. n castroville, uimnh 21428-29; 18 mi. n castroville, uimnh 21426-27, 21430-33; 6.5 mi. nw rio medina, ku 18440. _real co._: rio frio, fmnh 55156-57. _travis co._: austin, amnh 44221-22; mount bonnell, 5 mi. s austin, ummz 101453 (10). _uvalde co._: 13 mi. from uvalde, uimnh 62322. _val-verde co._: 40 mi. n del rio, jdl 214 (skeleton). [illustration: fig. 11: distribution of _syrrhophus marnockii_ (circles). starred localities are late pleistocene records.] =syrrhophus verrucipes= cope _syrrhophus verrucipes_ cope, 1885:383 [holotype.--ansp 11325, from near zacualtipán, hidalgo, méxico (1800 feet lower in a rocky gorge of a stream near its junction with the río san miguel), collected by dr. santiago bernard]. kellogg, 1932:126-29. smith and taylor, 1948:52-53. firschein, 1954:55-57. gorham, 1966:167. _syrrhaphus verrucipes_: günther, 1900:216-17. _tomodactylus macrotympanum_ taylor, 1940e:496-99, pl. 55, figs. 2a-b. [holotype.--fmnh 100049 (formerly eht-hms 6838), from la placita, 8 km. s jacala, hidalgo, méxico, 1850 m.; collected on july 2, 1936, by edward h. taylor]. smith and taylor, 1948:47-48. _syrrhophus macrotympanum_: dixon, 1957:384. gorham, 1966:165. _diagnosis._--medium-sized frogs, males 17.5-26.1 mm. snout-vent, females 28.0-31.7 mm. snout-vent length; vocal slits in males; digital tips slightly expanded; first finger shorter than second; skin of dorsum pustular, that of venter areolate; snout elongate, subacuminate; diameter of tympanum 56.1-76.7 per cent that of eye in males, 54.3-56.8 in females; in preservative, dorsum reddish brown with numerous small black or dark brown spots (fig. 8); venter white to cream; in life dorsum green with darker green spots, belly white; iris gold above, bronze below. _remarks._--cope's (1885) original description was not sufficiently clear to enable subsequent authors to recognize this species. taylor (1940e) described it as a _tomodactylus_, but dixon (1957) pointed out that _t. macrotympanum_ differed from the other species of the genus in having a poorly developed lumbo-inguinal (inguinal) gland, and placed the species in the genus _syrrhophus_. comparison of the holotypes of _s. verrucipes_ and _t. macrotympanum_ leaves no doubt in my mind that a single species is involved. this same species was reported by smith and taylor (1948) as _s. verruculatus_. _syrrhophus verrucipes_ bears resemblance to members of both the _leprus_ and _marnockii_ groups. in snout shape it is closer to the _leprus_ group, whereas in digital pad, the shape of the general body form, and contiguity of habitat it is most similar to the _marnockii_ group (_s. guttilatus_). _etymology._--latin, meaning warty foot, probably in reference to the numerous plantar supernumerary tubercles. _distribution._--moderate elevations in southeastern san luis potosí, queretaro, and northwestern hidalgo, méxico (fig. 7). _specimens examined_--(43) méxico, _hidalgo_: jacala, ummz 106434; 9.6 km. ne jacala, puerto de la zorra, 1820 m., ku 60240-41, tcwc 11090, 11147; 8 km. s jacala, la placita, 1850 m., fmnh 100049 (holotype of _tomodactylus macrotympanum_), 100791-803, 105334-35, 114287, uimnh 15989-92, 15995-96, ummz 117252, usnm 137202; tianguistengo, fmnh 113705-09, uimnh 13328-30; near zacualtipán, ansp 11325 (holotype of _syrrhophus verrucipes_). _queretaro_: 3.5 km. s san juan del río, eal 1343. _san luis potosí_: 9.6 km. w ahuacatlán, lsumz 4968-70. =syrrhophus dennisi= new species _syrrhophus latodactylus_: martin, 1958:49 (in part). _holotype._--ummz 101121, adult male from a cave near el pachón, 8 km. n antiguo morelos, tamaulipas, méxico, 250 m., collected on march 13, 1949, by paul s. martin. _paratopotypes._--(26). ummz 101122 (10), 101123 (2), 101126, 126993 (12). _diagnosis._--medium-sized frogs, males 22.8-28.4 mm. snout-vent, females 25.9-32.0 mm. snout-vent; vocal slits in males; digital tips greatly expanded, more than twice width of digit; first finger shorter than second; skin of dorsum shagreened to pustular, that of venter weakly to moderately areolate; toes webbed basally; dorsum light brown to tan with brown vermiculations; venter white; diameter of tympanum 53.9 to 64.2 per cent that of eye in males, 50.6 to 58.7 per cent in females. _description and variation._--(fig. 12). head wider than body; head as wide or wider than long in males, sometimes longer than wide in females; snout acuminate in dorsal view, elongate and rounded in lateral profile; canthus rostralis rounded but distinct; loreal region slightly concave, sloping abruptly to lip; lips not flared; eyelid about two-thirds interorbital distance; length of eye less than distance between eye and nostril; diameter of tympanum 53.9 to 64.2 per cent that of eye in males, 50.6 to 58.7 per cent in females; tympanum round and distinct in both sexes; supratympanic fold moderately distinct; choanae within border of jaws, completely visible from directly below, rounded to slightly oval; dentigerous processes of prevomers and teeth absent; tongue free for posterior one-half, generally oval in outline; vocal slits present in males. many scattered pustules on dorsum; flanks areolate; skin of venter areolate or not (variability may be due to differences in preservation); ventral disc distinct on chest and lower abdomen; inguinal gland present or not, when present varying from very large and distinct to poorly defined; axillary gland absent. first finger shorter than second; all fingers bearing truncate tips with pads, each pad having a terminal groove; fingers fringed; fingers three and four having dilated pads two to three times width of digit; subarticular tubercles large, conical, rounded, simple; supernumerary tubercles numerous on thenar surface, none on digits; three palmar tubercles, outer slightly smaller than largest supernumerary tubercles; row of tubercles on outer edge of forearm variable, weak to very distinct; tips of toes wider than digits, rounded to truncate at tips, each pad having terminal groove; toes having lateral fringes, bases of toes united by web, web not extending to basal subarticular tubercle; subarticular tubercles smaller than those of hand, round, conical, simple; supernumerary tubercles numerous on plantar surfaces, extending between metatarsal tubercles, present on toes between basal two subarticular tubercles in some specimens; outer metatarsal tubercle round, conical, one-half as large as ovoid, non-compressed inner metatarsal tubercle; tarsal tubercles or folds absent. ground color pale reddish-brown to tan dorsally, creamy on flanks; dorsal pattern consisting of reddish-brown to brown vermiculations extending onto flanks; distinct interorbital light bar present; loreal region darker than snout, reddish-brown compared to tan or pale reddish-brown; arms colored like dorsum; thighs banded, unicolor brown on posterior surfaces; shanks and tarsi banded; venter white to cream punctated with brown in some specimens. the variation in proportions is summarized in table 5. _remarks._--martin (1958) expressed some doubt that this series of 26 specimens was identical with "_s. latodactylus_." my study indicates that the specimens from el pachón represent a distinctive but allied species. males of the two species can be readily separated by the relative sizes of the tympani, presence or absence of vocal slits, and color pattern. females of the two species can be separated by color pattern. within the type-series, the pattern varies from weakly to strongly vermiculate but is always recognizable as vermiculate rather than spotted as in _s. longipes_ (= _s. latodactylus_ of taylor and martin). [illustration: fig. 12: _syrrhophus dennisi_ sp. nov., holotype, ummz 101121 (dorsum ×1.8, side of head ×6.1).] _etymology._--the specific name is a patronym for david m. dennis, whose drawings greatly enhance the worth of this paper. _distribution._--known only from the type series. =syrrhophus longipes= (baird), new combination _batrachyla longipes_ baird, 1859:35, pl. 37, fig. 1-3 [holotype.--apparently usnm 3237 (cited as 3207 by cope, 1887:16), now lost, from 40 leagues from (probably north) méxico city; collected by john potts]. kellogg, 1932:107. _epirhexis longipes_: cope, 1866:96. _eleutherodactylus longipes_: kellogg, 1932:107 (part). smith and taylor, 1948:61. lynch, 1963:580-581. gorham, 1966:82. _syrrhophus latodactylus_ taylor, 1940d:396-401, pl. 43, figs. a-f, text fig. 7 [holotype.--fmnh 100063 (formerly eht-hms 6807), from huasteca canyon, 15 km. w monterrey, nuevo león, méxico, 680 m.; collected on june 20, 1936, by edward h. taylor]. smith and taylor, 1948:50-52. martin, 1958:48-50. gorham, 1966:165. _diagnosis._--large frogs, males 22.1-33.2 mm. snout-vent, females 26.8-39.6 mm. snout-vent length; vocal slits lacking in males; digital tips greatly expanded (more than twice the width of digit); first finger shorter than second; skin of dorsum pustular, that of venter smooth; diameter of tympanum in males 61.1-87.2 per cent that of eye, 49.5-72.1 per cent in females; dorsum tan with large or small spots and blotches; limbs banded; interorbital bar or triangle present. _remarks._--i have applied baird's _batrachyla longipes_ to the frog taylor (1940d) called _syrrhophus latodactylus_ because the color pattern (fig. 13) predominant in the southern part of the range agrees with that described (figured) for _batrachyla longipes_. the color pattern of individuals in the southern part of the range of this species consists of large spots or blotches, whereas in the northwestern part the pattern is made up of smaller spots. in the northeastern part of the range, the pattern is more reduced and tends to consist of heavy flecking. the interorbital bar is narrower in specimens from nuevo león and tamaulipas and is triangular in specimens from hidalgo and queretaro. the status of the name _batrachyla longipes_ is currently that of a _nomen dubium_ (lynch, 1963). at that time, i was unaware of the geographic variation in color pattern in _syrrhophus latodactylus_. the exact type-locality of _batrachyla longipes_ is not known. if it is 40 leagues north of méxico city, the locality would be in an area where the species has a blotched instead of a flecked or spotted pattern. no justifiable evidence was presented to place _batrachyla longipes_ in _eleutherodactylus_ instead of _syrrhophus_. barbour (1923) and kellogg (1932) associated another species (_e. batrachylus_) with _longipes_. taylor (1940a) noted this as a case of misidentification and corrected the error but left _longipes_ in the genus _eleutherodactylus_. lynch (1963) noted several points of morphological agreement between _syrrhophus_ and _b. longipes_ but did not place _longipes_ in _syrrhophus_. baird's (1859) figures of the holotype do not illustrate prevomerine teeth, but according to cope (1866) they were present in the holotype. the digital tips of the frog in the figure are somewhat narrower than those typically seen in _s. latodactylus_. if the specimen was slightly desiccated, as possibly was the case, the digits would appear narrower. there is no evidence contrary to placing _syrrhophus latodactylus_ in the synonymy of _batrachyla longipes_. [illustration: fig. 13: dorsal views of _syrrhophus longipes_ illustrating geographic variation in pattern (left, tcwc 12179, ×1.5; right, ku 92572, ×1.8); side of head (tcwc 10966, ×6).] application of baird's name _batrachyla longipes_ to the species of frog heretofore called _syrrhophus latodactylus_ poses one serious problem. _batrachyla longipes_ is the type-species (by original designation) of the genus _epirhexis_ cope, 1866, which has priority over _syrrhophus_ cope, 1878. if _batrachyla longipes_ is left in the status of a _nomen dubium_, _epirhexis_ can be forgotten, for the two names are tied together. however, since it seems almost certain that _batrachyla longipes_ and _syrrhophus latodactylus_ are conspecific, the former name should not be left as a _nomen dubium_. _epirhexis_ never came into general usage (cope cited the name four times, but no one else has used it), whereas _syrrhophus_ is well established in the zoological literature. it would serve only to confuse the literature to adhere strictly to the law of priority and replace _syrrhophus_ with _epirhexis_. therefore, _syrrhophus_ is used in this paper, even though _epirhexis_ has priority. a request for the suppression of _epirhexis_ cope, 1866, has been submitted to the international commission of zoological nomenclature (lynch, 1967). _etymology._--latin, meaning long-footed; taylor's _latodactylus_ refers to the wide digital pads. [illustration: fig. 14: distribution of _syrrhophus dennisi_ (triangle) and _s. longipes_ (circles).] _distribution._--moderate elevations (650 to 2000 meters) along the sierra madre oriental from central nuevo león to northern hidalgo, méxico (fig. 14). _specimens examined._--(122) méxico, _hidalgo_: 3 km. ne jacala, amnh 52977; 9.6 km. ne jacala, 1800 m., tcwc 10966-70, 12179; 8 km. s jacala, la placita, 1850 m., fmnh 100266-68, 103244, uimnh 13291, 13327. _nuevo león_: salto cola de caballo, ku 92572; huasteca canyon, 15 km. w monterrey, 680 m., fmnh 100063 (holotype of _s. latodactylus_), uimnh 13290; 6.5 km. n pablillo, eal 1319; sabinas hidalgo, usnm 139728. _queretaro_: cueva de los riscos, 8 km. sw jalpan, ku 106300. _san luis potosí_: 13 km. e santa barberita, lsumz 2295; second camp, san luis potosí road, uimnh 13326; xilitla, cueva sin nombre, ummz 125892. _tamaulipas_: 4 km. w el carrizo, 500 m., ummz 111343 (31); 8 km. n chamal, bee cave, ku 106299; 14.5 km. nnw chamal, 420 m., ummz 111339-40, 111342 (4), 111344 (11); 19 km. nnw chamal, 700 m., ummz 111341 (3); el chihue, 1880 m., ummz 111289 (4); 11 km. n gómez farías, 1060 m., ummz 101166; 11 km. wnw gómez farías, 1800 m., ummz 108507 (3); 8 km. nw gómez farías, 1060-1400 m., lsumz 11085, ummz 101167 (3), 101168 (4), 101169 (2), 101170 (3), 101171 (2), 101360-61, 102860, 102933 (4), 102934 (2), 102935-38, 102939 (2), 102940-43, 108800 (3), 110735, 111345-46. =syrrhophus pipilans= taylor _syrrhophus pipilans_ taylor, 1940c:95-97, pl. 1 [holotype.--fmnh 100072 (formerly eht-hms 6843), 14.6 km. s mazatlán, guerrero, méxico; collected on july 22, 1936, by edward h. taylor]. _diagnosis._--medium sized frogs, males 22.6-28.5 mm. snout-vent, females 21.1-29.4 mm. snout-vent length; vocal slits present in males; finger tips slightly expanded, truncate in outline; inner metatarsal tubercle less than twice the size of outer; skin of dorsum smooth to shagreened, that of venter smooth; tympanum 36.5-54.0 per cent diameter of eye; dorsum dark brown with large or small light brown, orange-brown, or yellowish spots or blotches; limbs banded; interorbital bar absent. [illustration: fig. 15: dicegrams of ear size relative to eye diameter in the two subspecies of _syrrhophus pipilans_. n = 17 in _nebulosus_, 18 in _pipilans_.] _remarks._--two subspecies were recognized by duellman (1958). previously both had been treated as species. the two populations were distinguished on the basis of color pattern and the size of the tympanum. measurements of 17 males of _s. p. nebulosus_ from central chiapas and 18 males of _s. p. pipilans_ from southcentral oaxaca and guerrero, méxico, demonstrates that the supposed difference in tympanum size is not significant (fig. 15). there is, however, a tendency for the western population of _s. pipilans_ to have larger tympani. based on the present examination of 112 specimens of this species the two populations are held to be sufficiently distinct to warrant taxonomic recognition as subspecies (fig. 16). [illustration: fig. 16: _syrrhophus pipilans nebulosus_ (left, ku 58908) and _s. p. pipilans_ (right, ku 86885). ×2.7.] the parotoid glands attributed to this species by taylor (1940c:95) are merely the superficial expression of the _m. depressor mandibulae_ and scapula. no true glands are present in the parotoid region. =syrrhophus pipilans nebulosus= taylor _syrrhophus nebulosus_ taylor, 1943:353-55, pl. 27, figs. 3-5 [holotype.--fmnh 100095 (formerly eht-hms 3774), near tonolá, chiapas, méxico; collected on august 27, 1935, by hobart m. smith and edward h. taylor]. smith and taylor, 1948:49, 51. _syrrhophus pipilans nebulosus_: duellman, 1958:2-4, 9, 12, 14. stuart, 1963:32-33. gorham, 1966:166-67. _diagnosis._--diameter of tympanum 36.6-47.8 per cent that of eye; dorsum dark brown with numerous small light brown to yellowish spots. _remarks._--the distribution of this subspecies is adequately described by duellman (1958). fouquette (1960) described the vocalization of this frog. _etymology._--latin, _nebula_, in reference to the clouded dorsal pattern. _distribution._--low to moderate elevations along the pacific versant of chiapas and in the grijalva valley of chiapas and guatemala (fig. 17). _specimens examined._--(54) guatemala, _huehuetenango_: jacaltenango, ummz 117036; 35 km. se la mesilla, tnhc 29652. méxico, _chiapas_: 11.2 km. n arriaga, 300 m., ummz 125891; 11.8 km. n arriaga, ummz 117279; 12.8 km. n arriaga, ummz 117280; 17.5 km. s arriaga, uimnh 57108-109; 1.5 km. s bochil, 1250 m., ku 58898-908; cerro hueco, 7 km. s tuxtla gutierrez, ummz 123007; 3.2 km. s ixtapa, ummz 124000; linda vista, ca. 2 km. nw pueblo nuevo solistahuacán, ku 58897; hda. monserrate, 40 km. nw arriaga, ummz 102258; near san ricardo, fmnh 100720; tapachula, fmnh 75792, 103242, 100695-96, uimnh 13292; 56 km. e tapanatepec, oaxaca, tnhc 26942, tonolá, fmnh 100095 (holotype), 100686-92, uimnh 13293-95; tuxtla gutierrez, fmnh 100693-94, uimnh 13297; 19 km. n tuxtla gutierrez, tnhc 25229-30; 15.5 km. ne tuxtla gutierrez, ummz 119892 (3); 19 km. ne tuxtla gutierrez, ummz 119891 (3); 8 km. nnw tuxtla gutierrez, ku 37809; unión de juarez, fmnh 105294. =syrrhophus pipilans pipilans= taylor _?syrrhopus verruculatus_: gadow, 1905:194. _syrrhophus pipilans_ taylor, 1940c:95-97, pl. 1 [holotype.--fmnh 100072 (formerly eht-hms 6843), from 14.6 km. s mazatlán, guerrero, méxico; collected on july 22, 1936, by edward h. taylor]. taylor and smith, 1945:581-82. smith and taylor, 1948:49, 50-51. _syrrhophus pipilans pipilans_: duellman, 1958:1-4, 8-9, 13-14, pl. 2, fig. 1. gorham, 1966:166. _diagnosis._--diameter of tympanum 40.6-54.0 per cent that of eye; dorsum dark brown with large light spots or blotches. _remarks._--duellman's (1958) synopsis of this subspecies is adequate; the distribution has not been extended, but several records are now available which fill in gaps. [illustration: fig. 17: distribution of _syrrhophus pipilans_: _nebulosus_ (open circles) and _pipilans_ (solid circles).] gadow's (1905) record of _s. verruculatus_ from "buena vista, s. guerrero" is most likely applicable to this species. gadow simply included the name in a list of the species he had collected during his trip in méxico (1902-04); no further comment was made on this species although references to _syrrhopus_ (sic) appear in several places in the paper and would appear to apply to the species he had. _etymology._--latin, _pipilo_, chirping, peeping, in reference to the call of the male. _distribution._--sea level to about 1800 meters along the pacific versant of western méxico from central guerrero to the isthmus of tehuantepec (fig. 17). _specimens examined._--(62). méxico, _guerrero_: acapulco, ummz 110125; 6.4 km. n acapulco, fmnh 100389, 100525; agua del obispo, 980-1000 m., fmnh 75791, 100518-21, 100526, ku 86884-86, uimnh 13315, ummz 119152, 125890 (4); 13.3 km. nw coyuca, uimnh 38367, 71982-83; 14.5 km. s mazatlán, fmnh 100072 (holotype), 100408, 100511-17, uimnh 13302-309; tierra colorado, 300 m., ku 67961, uimnh 13313-14; near el treinte, fmnh 126639; xaltinanguis, fmnh 100522-24, 126640. _oaxaca_: cacahuatepec, uimnh 52853; 8 km. nw río canoa, 53 km. ese cuajinicuilapa, uimnh 52852; 6.4 km. n el candelaria, uimnh 9501; 11.2 km. s el candelaria, uimnh 9502; 17 km. ne juchatengo, 1600 m., ku 86887; 31.5 km. n pochutla, ummz 123999 (2); 32.9 km. n pochutla, 850 m., ummz 123996; 37.1 km. n pochutla, ummz 123998 (2); 41.4 km. n pochutla, ummz 123997 (2); cerro quiengola, fmnh 105653; 3.8 km. n santiago chivela, ummz 115449; 14.5 km. w tehuantepec, ummz 115448 (2). =syrrhophus interorbitalis= langebartel and shannon _syrrhophus interorbitalis_ langebartel and shannon, 1956: 161-65, figs. 1-2 [holotype.--uimnh 67061 (formerly fas 9378), 36 mi. n mazatlán, sinaloa, méxico, collected on november 17, 1955, by e. c. bay, j. c. schaffner, and d. a. langebartel]. duellman, 1958:1-4, 10, 12, 14. gorham, 1966:164-65. _syrrhophis interorbitalis_: campbell and simmons, 1962:194, fig. 1. [illustration: fig. 18: left to right. _syrrhophus interorbitalis_ (uimnh 38095, ×1.5), _s. nivocolimae_ (lacm 3203, ×1.3), and _s. teretistes_ (ku 75263, ×1.5).] _diagnosis._--medium sized frogs, only known male 25.6 mm. snout-vent, females 20.0-26.7 mm. snout-vent length (small sample); vocal slits in males; finger tips expanded; first finger shorter than second; outer metatarsal tubercle one-third size of inner; skin of dorsum shagreened, that of venter smooth; diameter of tympanum 37.7-42.4 per cent that of eye in both sexes; pale yellow-brown ground color mottled with brown; limb bands broad, much wider than narrow light interspaces; interorbital bar very long, edged with dark brown to black (fig. 18). _remarks._--duellman's (1958) measurements and proportions of _s. interorbitalis_ were based exclusively on the type series, which is composed of only females; therefore his _interorbitalis_ data are not comparable with the data for the other species in his table. campbell and simmons (1962) collected the only known male. the type series was collected beneath rocks in a stream bed; the collectors heard calling frogs in the bushes but were unable to obtain specimens (langebartel and shannon, 1956). campbell and simmons (1962) reported that their specimen had a poorly developed interorbital bar in life; in preservative the bar compares favorably with the bar in the female (fig. 18). _etymology._--latin, in reference to the pale interocular band. _distribution._--pacific lowlands of sinaloa, méxico (fig. 20). _specimens examined._--(10). méxico, _sinaloa_: 36 mi. n mazatlán, uimnh 38094-96, 67061 (holotype), 71970-74; 65 mi. n mazatlán, lacm 13773. =syrrhophus modestus= taylor _syrrhophus modestus_ taylor, 1942:304-06, pl. 29 [holotype.--fmnh 100048 (formerly eht-hms 3756), from hacienda paso del río, colima, méxico; collected on july 8, 1935, by hobart m. smith]. smith and taylor, 1948:49-50. _syrrhophus modestus modestus_: duellman, 1958:2-5, 7, 14, pl. 1, fig. 1. gorham, 1966:166. _diagnosis._--small frogs, males 15.8-20.1 mm. snout-vent length, single female 18.5 mm.; vocal slits present in males; finger tips widely expanded; first finger shorter than second; inner metatarsal tubercle about three times size of outer; skin of dorsum shagreened, that of venter smooth; tympanum concealed; pale cream in preservative with dark brown spots; limbs banded; bands on forearm and thigh poorly developed or absent; interorbital bar absent. _remarks._--the tympanum is concealed in _s. modestus_, _s. nivocolimae_, _s. pallidus_, _s. teretistes_, and to a lesser degree in _s. interorbitalis_. however, if the specimen is permitted to dry slightly, the annulus tympanicus becomes visible through the skin and a tympanum/eye ratio can be computed. one of the few cases of sympatry within the genus _syrrhophus_ involves this species; _modestus_ and _nivocolimae_ are known to be sympatric at one locality in southwestern jalisco, méxico. duellman (1958) used the trinomial for this population and named a new subspecies, _pallidus_, from nayarit. i consider _pallidus_ to be specifically distinct from _modestus_ because there is no evidence of genetic exchange, and there is no overlap in the distinguishing morphological features. i do consider the two populations to be closely related but feel the interrelationships between _modestus_, _pallidus_, _nivocolimae_, and _teretistes_ are more complex than would be indicated by the use of trinomials. the sympatric occurrence of _modestus_ and _nivocolimae_ is significant; morphologically, they might otherwise be regarded as subspecies. although allopatric, similar arguments could be advanced for the morphologically similar _pallidus_ and _teretistes_. the four are here afforded species rank since morphological similarity and allopatry are not sufficient grounds for the assumption of genetic exchange. [illustration: fig. 19: _syrrhophus modestus_ [left, ummz 115447 (wed 11155)] and _s. pallidus_ (right, ummz 115453). ×2.2.] _etymology._--latin, meaning unassuming, modest, in reference to the small size of the species. _distribution._--low elevations (up to 700 meters) in the lowlands and foothills of colima and southwestern jalisco, méxico (fig. 20). _specimens examined._--(14). méxico, _colima_: hda. paso del río, fmnh 100048 (holotype), 100167, 100299, uimnh 13300, ummz 110877 (2), usnm 139729; 7.2 km. sw tecolapa, ummz 115477 (4); _jalisco_: 17.6 km. sw autlan, 606 m., ku 102627; 3.2 km. n la resolana, ummz 102100; bahía tenacatita, ummz 84264. =syrrhophus nivocolimae= dixon and webb _syrrhophus nivocolimae_ dixon and webb, 1966:1-4, fig. 1 [holotype.--lacm 3200, from nevado de colima (6 airline miles west of atenquique), jalisco, méxico, 7800 feet; collected on july 20, 1964, by robert g. webb]. _diagnosis._--small frogs, males 18.5-21.1 mm. snout-vent length, only known female 24.1 mm. snout-vent; vocal slits present in males; finger tips widely expanded; first finger shorter than second; inner metatarsal tubercle about three times size of outer; skin of dorsum warty, that of venter smooth; tympanum concealed, its diameter 30.0-39.3 per cent that of eye in males; mid-dorsal brown band from interorbital bar to anus; bands on limbs narrow, dark bands less than one-half width of light bands, upper arm not banded; narrow interorbital light bar. _remarks._--this species is closely related to _s. modestus_ and differs in color pattern and degree of wartiness of the skin. dixon and webb (1966) held that _nivocolimae_ had no close relatives, but the condition of the tympanum, size, nature of the outer palmar tubercle, relative sizes of the metatarsal tubercles, and shape and size of the digital pads all point to a close relationship between _s. modestus_, _s. nivocolimae_, and _s. pallidus_. [illustration: fig. 20: distribution of the species of the _modestus_ group: _interorbitalis_ (open circles), _teretistes_ (solid circles), _modestus_ (open triangles), _pallidus_ (solid triangles) and _nivocolimae_ (square). arrow indicates locality of sympatry between _modestus_ and _nivocolimae_. solid line about the localities for _interorbitalis_ is a range estimate based on call records and specimens examined.] dixon and webb (1966) reported that _s. nivocolimae_ has a large tympanum (50.0-59.0 per cent diameter of eye). however, my examination of the type series and several other specimens from jalisco reveals that the largest tympanum/eye ratio is 39.3 per cent. therefore, the tympanum/eye ratio in _s. nivocolimae_ is in agreement with those for _s. modestus_, _s. pallidus_, and _s. teretistes_ (table 6). _etymology._--_niv_, latin, and colima (nevado de), meaning high on the volcano, in reference to the higher distribution of this species (around 2000 meters) than other members of the group. _distribution._--known from southwestern jalisco, méxico, at moderate to high elevations (600-2400 meters). _specimens examined._--(48) méxico, _jalisco_: 17.6 km. sw autlán, 606 m., ku 102626, 102631; 6.4 km. w atenquique, 2060 m., ku 102628-30, 102632; 8 km. w atenquique, 1970 m., lacm 3210-12; 9.6 km. w atenquique, 2360 m., lacm 3200 (holotype), 3201-09; 14.5 km. w atenquique, 2000 m., lacm 25424-36, 25439-41, 25446; 15 km. w atenquique, lacm 37044-46, 37244-47; 16 km. w atenquique, 2105 m., lacm 25443-45; 17 km. w atenquique, 2180 m., lacm 25442. =syrrhophus pallidus= duellman, new combination _syrrhophus modestus_: davis and dixon, 1957:146. _syrrhophus modestus pallidus_ duellman, 1958:2-3, 5-7, 14, pl. 3 [holotype.--ummz 115452, from san blas, nayarit, méxico, sea level; collected on august 13, 1956, by william e. and ann s. duellman]. zweifel, 1960:86-88, 91, 93-94, 118, 120-22. gorham, 1966:166. _syrrhophis modestus pallidus_: campbell and simmons, 1962:194. _diagnosis._--small frogs, males 17.9-19.3 mm. snout-vent length; vocal slits in males; finger tips widely expanded; first finger shorter than second; inner metatarsal tubercle about three times size of outer; skin of dorsum shagreened, that of venter smooth; tympanum concealed, its diameter 27.0-35.6 per cent of eye in males; ground color cream vermiculated with brown, upper arm and thigh lacking, or with few, indistinct, bands; interorbital bar absent. _remarks._--considerable debate has been waged relative to the value of subspecies and to the reasons for recognizing distinct disjunct populations as species versus subspecies. lacking evidence of genetic exchange, i prefer to retain disjunct populations that are distinctive as species. all known specimens of _pallidus_ can be separated from those of _modestus_ by color pattern. the two nominal species exhibit overlap in proportions but the same can be said about nearly every species of _syrrhophus_; therefore, overlap in proportions can be disregarded in assessing specific versus subspecific rank. until contrary evidence is forthcoming, i consider the disjunct populations heretofore held to be subspecies of _modestus_ to be specifically distinct. the specimens of the disjunct population of _pallidus_ on the tres marias do not differ from the mainland population in nayarit. this evidence, though perhaps secondary, supports my contention that two species should be recognized. _etymology._--latin, in reference to the pale ground color in comparison with that of _s. modestus_. _distribution._--low elevations in coastal nayarit and on islas tres marias (fig. 20). _specimens examined._--(12) méxico, _nayarit_: 18.8 mi. nw ahuacatlán, uimnh 7808; san blas, ummz 115452 (holotype), 115453-57; 17 km. ne san blas, 150 m., msu 5085; 12.8 km. e san blas, uimnh 71979; 31 km. e san blas, uimnh 71978; 13.5 km. n tepic, uimnh 71980-81. =syrrhophus teretistes= duellman _syrrhophus teretistes_ duellman, 1958:2-3, 10-14, pl. 2, fig. 2 [holotype.--ummz 115451, from 4.8 km. nw tepic, nayarit, méxico, 840 m.; collected on august 12, 1956, by william e. duellman]. gorham, 1966:167. _diagnosis._--medium-sized frogs, males 19.2-23.2 mm. snout-vent length, single known female 24.8 mm. snout-vent; vocal slits in males; finger tips widely expanded; first finger shorter than second; inner metatarsal tubercle about three times size of outer; skin of dorsum shagreened, that of venter smooth; tympanum partially concealed, its diameter 28.6-43.8 per cent of eye in males; ground color brown vermiculated with dark brown to nearly black; upper arm and thigh banded; interorbital light bar absent. _remarks._--_s. teretistes_ appears to be most closely related to _s. pallidus_; i consider it to be an upland derivative of _pallidus_. morphologically, the differences between the two are few, but lacking evidence of genetic exchange they are retained as species. _etymology._--greek, in reference to the whistle-like nature of the call. _distribution._--moderate elevations (840-1200 meters) in the sierra occidental of nayarit, sinaloa, and durango, méxico (fig. 20). _specimens examined._--(13) méxico, _nayarit_: 4.8 km. nw tepic, 840 m., ummz 115451 (holotype). _sinaloa_: santa lucía, 1090 m., ku 75263-72; 1 km. ne santa lucía, 1156 m., ku 78257; 2.2 km. ne santa lucía, 1156 m., ku 78258. discussion there are relatively few clear-cut morphological differences among the fourteen species now assigned to _syrrhophus_. the majority of the species are allopatric and differ primarily in color patterns. sympatric occurrence serves as an indicator of specific distinctness and is one of the more practical tests of species validity when cross-breeding experiments are not possible. two cases of sympatric occurrence are known for the species of _syrrhophus_ in western méxico: _modestus_ and _nivocolimae_ are sympatric in southern jalisco and _pipilans nebulosus_ and _rubrimaculatus_ are sympatric in southeastern chiapas. in eastern méxico, _longipes_ and _verrucipes_ are sympatric in southern hidalgo, and _longipes_ is sympatric with _cystignathoides_, _dennisi_, and _guttilatus_ in southern tamaulipas. _syrrhophus cystignathoides_ and _leprus_ are apparently sympatric in central veracruz. subspecific assignments have been made only when there is evidence of intergradation. the sympatric occurrence of morphologically similar species in this genus has led me to adopt a conservative approach to the degree of difference philosophy. i have therefore recognized all morphologically distinct allopatric populations as species. [illustration: fig. 21: generic distributions of _syrrhophus_ (stipple) and _tomodactylus_ (hatching). black areas are zones of intergeneric sympatry.] _syrrhophus_ is closely allied to another mexican leptodactylid genus, _tomodactylus_, which was revised by dixon (1957), who along with numerous other authors noted the close relationship between the two genera. there is an almost complete lack of sympatry between the two genera; in very few places in méxico do they coexist (fig. 21). _tomodactylus_ has its greatest diversity in the cordillera volcánica and sierra madre del sur, whereas _syrrhophus_ reaches its greatest diversity in the sierra madre oriental and eastern foothills. the species of both genera are about the same size and presumably have similar requirements insofar as food, breeding sites, and habitat selection. four cases of intergeneric sympatry are known for the two genera: 1) the chilpancingo region of guerrero, 2) the lowlands of colima and the mountains just inland in jalisco, 3) the lowlands of central nayarit, and 4) the sierra madre occidental on the durango-sinaloan border. the apparent sympatry in the chilpancingo region involves four species: _s. pipilans_, _t. albolabris_, _t. dilatus_, and _t. nitidus_. of the four, _t. dilatus_ appears to be completely allopatric in that it occurs at higher altitudes (above 2000 meters), whereas the other three occur below 1800 meters in the region (davis and dixon, 1965). in the colima-jalisco region, _tomodactylus_ tends to occur higher (dixon and webb, 1966) than some of the _syrrhophus_, but one subspecies of _tomodactylus nitidus_ is a lowland frog, occurring sympatrically with the lowland _syrrhophus modestus_. a similar situation is observed in nayarit; the lowland _tomodactylus_ occurs sympatrically with the small _syrrhophus pallidus_. in both cases the _syrrhophus_ is smaller than the _tomodactylus_. [illustration: fig. 22: altitudinal distributions of _syrrhophus_ and _tomodactylus_. widths of the columns are proportional to the numbers of species at a given altitude; narrowest width equals one species.] frogs of the genus _syrrhophus_ tend to occur at lower elevations than do their close relatives of the genus _tomodactylus_ (fig. 22). this generalization is complicated by the occurrence in the sierra madre oriental in relatively high altitude _syrrhophus_ (up to 2000 m.) and the occurrence in michoacán of low altitude _tomodactylus_ (to sea level). there are no _tomodactylus_ in the sierra madre oriental, whereas the genus _syrrhophus_ is represented in the lowlands of western méxico (_modestus_ group). _syrrhophus_ and _tomodactylus_ exhibit essentially parapatric distributions. the two genera as now composed can be characterized as low to moderate elevation frogs (_syrrhophus_) and moderate to intermediate elevation frogs (_tomodactylus_). literature cited baird, s. f. 1859. reptiles of the boundary. united states and mexican boundary survey, pp. 1-35, pls. 1-41. barbour, t. 1923. the reappearance of batrachyla longipes. proc. new england zool. club, 8:81-83. barbour, t., and a. loveridge 1946. typical reptiles and amphibians; supplement. bull. mus. comp. zool., 96:59-214. boulenger, g. a. 1882. catalogue of the batrachia salientia ... british museum., 2nd ed. 1888. note on the classification of the ranidae. proc. zool. soc. london, 1888, pt. 2:204-06. campbell, h. w., and r. s. simmons 1962. notes on some reptiles and amphibians from western mexico. bull. so. california acad. sci., 61:193-203. conant, r. 1958. a field guide to reptiles and amphibians. houghton-mifflin co. boston. 366 pp. cope, e. d. 1866. on the structures and distribution of the genera of the arciferous anura. j. acad. nat. sci. philadelphia, n. ser., 6:67-112. 1877. tenth contribution to the herpetology of tropical america. proc. amer. philos. soc., 17:85-98. 1878. new genus of cystignathidae from texas. amer. nat., 12:252-53. 1879. eleventh contribution to the herpetology of tropical america. proc. amer. philos. soc., 18:261-77. 1885. a contribution to the herpetology of mexico. _ibid._, 22:379-404. davis, w. b., and j. r. dixon 1957. notes on mexican amphibians, with description of a new _microbatrachylus_. herpetologica, 13:145-47. 1965. amphibians of the chilpancingo region, mexico. _ibid._, 20:225-33. díaz de león, j. 1904. indice de los batracios que se enquentran en la republica méxicana. imprenta de ricardo rodriquez romo. aguascalientes. 40 pp. dixon, j. r. 1957. geographic variation and distribution of the genus tomodactylus in mexico. texas j. sci., 9:379-409. dixon, j. r., and r. g. webb 1966. a new _syrrhophus_ from mexico (amphibia: leptodactylidae). cont. sc., los angeles co. mus., 102:1-5. duellman, w. e. 1958. a review of the frogs of the genus _syrrhophus_ in western mexico. occ. pap. mus. zool. univ. michigan, 594:1-15. 1960. a distributional study of the amphibians of the isthmus of tehuantepec, mexico. univ. kansas publs. mus. nat. hist., 13:19-72. firschein, i. l. 1954. definition of some little-understood members of the leptodactylid genus _syrrhophus_, with a description of a new species. copeia, (1):48-58. fouquette, m. j. 1960. call structure in frogs of the family leptodactylidae. texas j. sci., 12:201-15. gadow, h. 1905. the distribution of mexican amphibians and reptiles. proc. zool. soc. london, 1905, pt. 2:191-244. gorham, s. w. 1966. liste der rezenten amphibien und reptilien.... das tierreich. lief, 85:1-222. günther, a. c. l. g. 1885-1902. biologia centrali-americana. reptilia and batrachia. 326 pp., 76 pls. syrrhophus section dated 1900. kellogg, r. 1932. mexican tailless amphibians in the united states national museum. bull. u.s. natl. mus., 160. langebartel, d. a., and f. a. shannon 1956. a new frog (syrrhophus) from the sinoloan lowlands of mexico. herpetologica, 12:161-65. lynch, j. d. 1963. the status of _eleutherodactylus longipes_ (baird) of mexico (amphibia: leptodactylidae). copeia, (3):580-81. 1964. additional hylid and leptodactylid remains from the pleistocene of texas and florida herpetologica. 20:141-42. 1967. _epirhexis_ cope, 1866 (amphibia: salientia): request for suppression under the plenary powers. i. n. (s). bull. zool. nomencl., 24:313-15. 1968. genera of leptodactylid frogs in méxico. univ. kansas publs., mus. nat. hist., 17:503-15. martin, p. s. 1958. a biogeography of reptiles and amphibians in the gomez farias region, tamaulipas, mexico. misc. publs. mus. zool. univ. michigan, 101:1-102. milstead, w. m., j. s. mecham, and h. mcclintock 1950. the amphibians and reptiles of the stockton plateau in northern terrell county, texas. texas j. sci., 2:543-62. neill, w. t. 1965. new and noteworthy amphibians and reptiles from british honduras. bull. florida state mus., 9:77-130. nieden, f. 1923. anura i ... das tierreich. lief., 46:1-584. peters, w. 1871. über neue amphibien ... des konigl. zoologischen museums. monatsb. k. k. preuss. akad. wiss. berlin, 1870:641-52. schmidt, k. p., and t. f. smith 1944. amphibians and reptiles of the big bend region of texas. field mus. nat. hist., zool. ser., 29:75-96. smith, h. m. 1947. notes on mexican amphibians and reptiles. j. washington acad. sci., 37:408-12. smith, h. m., and e. h. taylor 1948. an annotated checklist and key to the amphibia of mexico. bull. u.s. natl. mus., 194:1-118. stejneger, l. 1915. a new species of tailless batrachian from north america. proc. biol. soc. washington, 28:131-32. taylor, e. h. 1940a. a new eleutherodactylid frog from mexico. proc. new england zool. club, 18:13-16. 1940b. two new anuran amphibians from mexico. proc. u.s. natl. mus., 89:43-47, 1 pl. 1940c. a new syrrhophus from guerrero, mexico. proc. biol. soc. washington, 53:95-98, 1 pl. 1940d. new species of mexican anura. univ. kansas sci. bull., 26:385-405. 1940e. herpetological miscellany no. i. _ibid._, 26:489-571. 1942. new caudata and salientia from méxico. _ibid._, 28:295-323. 1943. herpetological novelties from mexico. _ibid._, 29:343-61. 1952. a review of the frogs and toads of costa rica. _ibid._, 35:577-942. taylor, e. h., and h. m. smith 1945. summary of the collections of amphibians made in mexico under the walter rathbone bacon traveling scholarship. proc. u.s. natl. mus., 95:521-613. tihen, j. a. 1960. notes on late cenozoic hylid and leptodactylid frogs from kansas, oklahoma and texas. southwest. nat., 5:66-70. wright, a. h., and a. a. wright 1949. handbook of frogs and toads. 3rd ed. comstock. 640 pp. yarrow, h. c. 1882. checklist of north american reptilia and batrachia, with catalogue of specimens in u.s. national museum. bull. u.s. natl. mus., 24:1-249. zweifel, r. g. 1960. results of the puritan-american museum of natural history expedition to western mexico. 9. herpetology of the tres marias islands. bull. amer. mus. nat. hist., 119:81-128. transcriber's notes although _syrrhophus marnocki_ and _syrrhophus marnockii_ both appear in this text, a literature search shows that both spellings have been used and the two instances where there is only one "i" at the end are in reference to priviously published names. therefore, they were left as is. with the exception of the list below and a number of silent corrections, the text presented is that of the original printed version. typographical corrections page correction ==== ====================== 3 otherwse => otherwise 5 poltypic => polytypic 12 interorbtal => interorbital 14 neublosus => nebulosus 16 cuidad => ciudad 16 1946-170 => 1946:170 22 rubrimacultaus => rubrimaculatus 27 resemblence => resemblance text emphasis _text_ italics =text= bold university of kansas publications museum of natural history volume 17, no. 12, pp. 517-557, pls. 13-16, 9 figs. march 20, 1968 middle american frogs of the hyla microcephala group by william e. duellman and m. j. fouquette, jr. university of kansas lawrence 1968 university of kansas publications, museum of natural history editors: e. raymond hall, chairman, henry s. fitch, frank b. cross volume 17, no. 12, pp. 517-557, 4 pls. 9 figs. published march 20, 1968 university of kansas lawrence, kansas printed by robert r. (bob) sanders, state printer topeka, kansas 1968 31-9419 middle american frogs of the hyla microcephala group by william e. duellman and m. j. fouquette, jr. contents page introduction 519 acknowledgments 520 materials and methods 520 hyla microcephala group 521 key to species and subspecies 522 accounts of species and subspecies 523 cranial osteology 540 analysis of mating calls 544 life history 550 phylogenetic relationships 552 literature cited 556 introduction the small yellow tree frogs, _hyla microcephala_ and its relatives, are among the most frequently heard and commonly collected frogs in the lowlands of southern méxico and central america. the similarities in size, proportions, and coloration of the different species have resulted not so much in a multiplicity of specific names, but in differences of opinion on the application of existing names to the various taxa. for example, the populations on the atlantic lowlands have been known by three names, two of which have been applied to other taxa. much of the confusion has been the result of previous workers' unfamiliarity with the animals in life and unawareness of the intraspecific geographic variation in the most widespread species. independently we undertook studies of these frogs in the field. the second author worked on the interspecific relationships and isolating mechanisms in panamá (fouquette, 1960b) and later studied the species in southern méxico. as part of his survey of the hylids of middle america, the first author accumulated field and laboratory data on the frogs throughout their ranges in méxico and central america. the purpose of this report is to present our findings on the four species of middle american frogs that we place in the _hyla microcephala_ group. in addition to conventional taxonomic characters, we have utilized the features of the cranial osteology and have relied heavily on the data obtained from an analysis of the mating calls. furthermore, we have included ecological and distributional data in our synthesis of interspecific relationships. acknowledgments examination of specimens was made possible by the provision of working space at various institutions or through the loan of specimens. for their generosity in this manner we are grateful to richard j. baldauf, charles m. bogert, james e. böhlke, doris m. cochran, robert f. inger, john m. legler, alan e. leviton, gerald raun, jay m. savage, hobart m. smith, robert c. stebbins, wilmer w. tanner, charles f. walker, ernest e. williams, and richard g. zweifel. duellman is especially grateful to charles w. myers, linda trueb, jerome b. tulecke, and john wellman for their assistance in the field and to linda trueb for her work on the cranial osteology that is incorporated in this report. fouquette is indebted to h. morgan smith and a. c. collins for assistance in the field, to a. j. delahoussaye for assistance in the laboratory, and to w. frank blair for use of the facilities of the sound laboratory at the university of texas and for much help in the early stages of this study. the research reported herein was accomplished mainly through support by the national science foundation (grants nsf g-9827 and gb-1441 to duellman and gb-599 to fouquette). the latter's field work in méxico was assisted in part by nsf grant g-4956 to w. frank blair. some of the field studies carried out in panamá by duellman were supported by a grant from the national institutes of health (nih gm-12020). we are grateful to many persons, too numerous to mention, who in various ways aided our field work in middle america. we are especially indebted to dr. rodolfo hernandez corzo and the late ing. luis macías arellano of the dirección general de la fauna silvestre of the mexican government for providing permits to collect in méxico. materials and methods for this report, data has been obtained from 2829 preserved frogs, 42 skeletal preparations, 8 lots of tadpoles and young, and 4 lots of eggs. much of the material was collected in our independent field work, which has extended over a period of 11 years. measurements were taken in the manner described by duellman (1956). osteological data were obtained from specimens that were cleared in potassium hydroxide, stained with alizarin red, and stored in glycerine. recordings were made by means of magnemite portable tape recorders (amplifier corp. america). the calls recorded by fouquette were analyzed on a sonagraph (kay electric co.) at the university of texas; those recorded by duellman were analyzed mainly on a vibralyzer (kay electric co.) at the university of kansas and in part on a sonagraph at the university of southwestern louisiana. sample calls were analyzed on all three instruments; the slight differences in results were found to be less than the error in measurement, so the data from all sources were combined without correction. the techniques and terminology of the calls are those defined by fouquette (1960a, 1960b). in the accounts of the species we have attempted to give a complete synonymy. at the end of each species account the localities from which specimens were examined are listed alphabetically within each state, province, or department, which in turn are listed alphabetically within each country. the countries are arranged from north to south. localities preceded by an asterisk (*) are not plotted on the accompanying maps due to the crowding of symbols that would have resulted. abbreviations for museum specimens are listed below: amnh --american museum of natural history ansp --academy of natural sciences of philadelphia bmnh --british museum (natural history) byu --brigham young university cas --california academy of sciences fmnh --field museum of natural history ku --university of kansas museum of natural history mcz --museum of comparative zoology mvz --museum of vertebrate zoology su --stanford university uimnh--university of illinois museum of natural history ummz --university of michigan museum of zoology usc --university of southern california usnm --united states national museum uu --university of utah tcwc --texas cooperative wildlife collection tnhm --texas natural history museum hyla microcephala group _definition._--small hylids attaining a maximum snout-vent length of 27 mm. in males and 32 mm. in females; dorsum yellowish tan with brown markings; thighs uniformly yellow, vocal sac in breeding males yellow; snout truncate in lateral profile; tympanum distinct, usually slightly smaller than one-half diameter of eye; vocal sac single, median, subgular; fingers about one-third webbed; toes webbed nearly to bases of discs, except only to middle of antepenultimate or base of penultimate phalanx of fourth toe; tarsal fold weak; inner metatarsal tubercle low, flat, elliptical; axillary membrane present; pupil horizontally elliptical; palpebral membrane unmarked; cranial elements reduced in ossification; sphenethmoid small, short; frontoparietal fontanelle large; tegmen tympani not extensive; quadratojugal greatly reduced; anterior arm of squamosal extending only about one-fourth distance to maxillary; posterior arm of squamosal not having bony connection with proötic; nasals lacking maxillary processes; medial ramus of pterygoid not having bony attachment to proötic; maxillary, premaxilary, and prevomerine teeth present; palatine and parasphenoid teeth absent; mentomeckelians ossified; tadpoles having xiphicercal tails with deep caudal fins and terminal mouth lacking teeth; mating call consisting of one primary note followed by a series of shorter secondary notes; haploid number of chromosomes, 15 (known only in _h. microcephala_ and _h. phlebodes_.) _content._--as recognized here the _hyla microcephala_ group contains four species, one having two subspecies. an alphabetical list of the specific and subspecific names that we consider to be applicable to the _hyla microcephala_ group are listed below. names proposed valid names _hyla cherrei_ cope, 1894 ? = _h. m. microcephala_ _hyla microcephala_ cope, 1886 = _h. m. microcephala_ _hyla microcephala_ boulenger, 1898 (_nec_ cope, 1886) = _h. microcephala underwoodi_ _hyla microcephala martini_ smith, 1951 = _h. microcephala underwoodi_ _hyla microcephala sartori_ smith, 1951 = _h. sartori_ _hyla phlebodes_ stejneger, 1906 = _h. phlebodes_ _hyla robertmertensi_ taylor, 1937 = _h. robertmertensi_ _hyla underwoodi_ boulenger, 1899 = _h. microcephala underwoodi_ _discussion._--the color pattern is the most useful character in distinguishing the species of the _hyla microcephala_ group from one another. except in _hyla microcephala_, little geographic variation in color pattern is noticeable. the features of color pattern that are helpful in identifying the species are: 1) presence or absence of lateral dark brown stripe; 2) longitudinal extent and width of lateral stripe, if present; 3) presence or absence of a narrow white line just dorsal to the lateral dark stripe; 4) presence or absence of an interorbital dark mark; 5) the arrangement of dark markings on the back, either as longitudinal lines or series of dashes, or in the form of various kinds of transverse markings; 6) presence of dark flecks, longitudinal line, or transverse marks on shanks. few consistent differences in measurements and proportions exist among the species (table 1). the most obvious morphological difference is that the head is noticeably narrower in _h. robertmertensi_ than in the other species. _hyla phlebodes_ is the smallest species; adult males attain snout-vent lengths of only 23.6 mm. the body is slender in _h. microcephala_ and _robertmertensi_, slightly wider in _phlebodes_, and noticeably broader in _sartori_. _distribution._--the composite range of the middle american frogs of the _hyla microcephala_ group includes the lowlands of southern méxico and central america, in some places to elevations of 1200 meters, southeastward from southern jalisco and southern veracruz, excluding arid regions (northern yucatán peninsula, balsas-tepalcatepec basin, plains of tehuantepec, grijalva valley, salamá basin, and upper motagua valley) to the pacific lowlands and the cauca and magdalena valleys in colombia. key to species and subspecies 1. lateral dark stripe, bordered above by narrow white line, extending from snout at least to sacral region 2 lateral dark stripe, if present, not extending posteriorly to sacral region and not bordered above by narrow white line 4 2. lateral dark stripe continuous to groin; dark flecks or longitudinal line on shanks; interorbital dark bar absent; dorsal pattern usually consisting of pair of longitudinal dark lines or series of dashes 3 lateral dark stripe usually extending only to sacral region; dark transverse bars on shanks; interorbital bar usually present; dorsal pattern usually consisting of interconnecting dark lines, sometimes forming transverse marks _h. microcephala underwoodi_ 3. lateral dark stripe narrow, covering only upper edge of tympanum; dorsal longitudinal stripes continuous, extending to vent _h. microcephala microcephala_ lateral dark stripe wide, encompassing entire tympanum; dorsal markings consisting of longitudinal series of flecks or dashes, or of two lines, usually not extending to vent _h. robertmertensi_ 4. lateral dark stripe indistinct, present only above tympanum and insertion of arm; dorsal markings consisting of narrow lines and dashes, sometimes interconnected; transverse bars on shanks narrow relative to interspaces _h. phlebodes_ lateral dark stripe absent; dorsal markings consisting of two broad chevron-shaped marks; transverse bars on shanks wide relative to interspaces _h. sartori_ accounts of species and subspecies _hyla microcephala_ cope _diagnosis._--lateral dark stripe narrow, covering only upper edge of tympanum, bordered above by narrow white stripe; dorsal pattern consisting of pair of longitudinal brown lines and no interorbital bar (eastern populations), or of irregular dark markings forming an xor )(-shaped mark in scapular region and an interorbital bar (western populations). _content._--the populations inhabiting the pacific lowlands of southeastern costa rica eastward to colombia are recognized herein as _hyla microcephala microcephala_ cope; the populations in western costa rica northward to méxico are assigned to _hyla microcephala underwoodi_ boulenger. _distribution._--southern veracruz and northern oaxaca southeastward through the atlantic lowlands of central america to north-central nicaragua, thence southeastward on the pacific lowlands to eastern panamá, and thence into the cauca and magdalena valleys (caribbean drainage) of colombia (fig. 1). [illustration: fig. 1. map showing locality records for _hyla microcephala_.] table 1.--variation in certain measurements and properties in the hyla microcephala group. (all data based on adult males; mean and standard error of mean below observed range.) ======================================================================== locality | n | snout-vent | tibia length |foot length| | | length | -----------| --------| | | (s-v l) | s-v l | s-v l | ----------------------------------------------------------------------- | _h. m. microcephala_ | panamá: canal zone | 25 | 21.5-24.1 | 50.2-56.0 | 40.9-46.6 | | | 22.8±0.20 | 52.9±0.37 | 43.5±0.28 | | | | | | costa rica: golfito | 25 | 18.5-24.5 | 49.1-54.4 | 41.8-48.0 | | | 22.4±0.27 | 51.6±0.26 | 45.1±0.32 | | | _h. m. underwoodi_ | | nicaragua: la cumplida | 25 | 23.0-25.6 | 51.0-55.7 | 41.3-46.5 | | | 24.1±0.19 | 52.9±0.25 | 43.7±0.25 | | | | | | guatemala: finca chamá | 25 | 21.8-25.0 | 51.0-57.2 | 41.2-47.8 | | | 23.5±0.16 | 54.3±0.39 | 44.4±0.30 | | | | | | tabasco: teapa | 25 | 22.7-25.8 | 48.0-54.5 | 40.7-46.8 | | | 24.3±0.14 | 51.5±0.29 | 43.3±0.25 | | | | | | oaxaca: donají-sarabia | 25 | 22.1-25.9 | 49.8-55.6 | 40.5-46.6 | | | 23.8±0.19 | 52.8±0.33 | 43.4±0.27 | | | | | | veracruz: alvarado | 25 | 21.9-25.4 | 49.6-54.4 | 40.7-47.5 | | | 24.1±0.17 | 51.1±0.28 | 42.6±0.34 | | | _h. robertmertensi_ | guatemala: la trinidad | 21 | 21.8-24.6 | 47.1-52.8 | 40.9-51.3 | | | 23.4±0.15 | 49.9±0.34 | 43.5±0.17 | | | | | | chiapas: acacoyagua | 25 | 21.4-25.7 | 47.8-52.4 | 41.7-46.3 | | | 24.1±0.20 | 50.4±0.45 | 43.9±0.23 | | | | | | oaxaca: tapanatepec | 25 | 22.4-26.4 | 44.1-48.3 | 39.1-44.5 | | | 24.7±0.18 | 46.4±0.23 | 41.7±0.23 | | | _h. phlebodes_ | panamá: canal zone | 25 | 19.6-23.2 | 49.1-56.9 | 41.9-47.1 | | | 22.2±0.16 | 52.8±0.35 | 45.4±0.26 | | | | | | costa rica: turrialba | 25 | 19.7-23.6 | 47.4-55.7 | 38.1-46.4 | | | 22.0±0.18 | 51.1±0.35 | 42.8±0.38 | | | _h. sartori_ | guerrero: tierra colorada| 25 | 23.7-26.0 | 47.2-51.4 | 42.4-47.8 | | | 24.8±0.13 | 49.6±0.23 | 45.2±0.27 | -----------------------------------------------------------------------table 1. (continued) =============================================================== locality | head length | head width | tympanum | ----------| ---------| ------- | s-v l | s-v l | eye -------------------------------------------------------------- | _h. m. microcephala_ | panamá: canal zone | 28.5-32.8 | 28.1-30.9 | 44.0-54.1 | 31.0±0.22 | 29.4±0.11 | 49.0±0.55 | costa rica: golfito | 30.2-35.5 | 29.0-32.7 | 40.0-57.8 | 33.1±0.25 | 30.8±0.16 | 48.4±1.10 | | _h. m. underwoodi_ | nicaragua: la cumplida | 29.7-33.5 | 28.9-31.8 | 42.3-60.0 | 31.6±0.19 | 30.4±0.17 | 49.3±0.97 | guatemala: finca chamá | 30.8-35.3 | 29.6-33.6 | 37.5-56.4 | 33.0±0.16 | 31.3±0.36 | 45.2±0.89 | tabasco: teapa | 29.5-33.0 | 28.7-31.8 | 40.7-53.8 | 31.7±0.17 | 30.3±0.16 | 45.5±0.38 | oaxaca: donají-sarabia | 30.4-34.8 | 28.9-32.6 | 37.0-54.1 | 32.8±0.19 | 30.8±0.17 | 45.1±0.76 | veracruz: alvarado | 29.9-33.8 | 29.1-32.9 | 40.7-53.8 | 31.4±0.18 | 30.5±0.17 | 46.6±0.65 | | _h. robertmertensi_ | guatemala: la trinidad | 30.0-33.3 | 27.3-29.8 | 44.4-50.0 | 31.3±0.20 | 28.5±0.23 | 47.4±0.46 | | | chiapas: acacoyagua | 29.1-32.7 | 26.0-30.3 | 42.8-53.8 | 31.2±0.29 | 28.1±0.20 | 46.5±0.50 | | | oaxaca: tapanatepec | 26.1-30.4 | 25.4-28.1 | 45.8-58.3 | 28.4±0.16 | 26.8±0.14 | 52.9±0.77 | | _h. phlebodes_ | panamá: canal zone | 33.6-37.4 | 32.3-36.0 | 37.9-46.4 | 34.8±0.18 | 33.8±0.18 | 41.6±0.49 | | | costa rica: turrialba | 32.6-35.9 | 30.5-35.0 | 35.7-48.2 | 34.1±0.16 | 32.9±0.17 | 40.1±0.53 | | _h. sartori_ | guerrero: tierra colorada| 29.4-31.8 | 28.9-31.0 | 42.3-52.0 | 30.6±0.13 | 30.0±0.12 | 47.4±0.59 --------------------------------------------------------------_hyla microcephala microcephala_ cope _hyla microcephala_ cope, proc. amer. philos. soc., 23:281, february 11, 1886 [syntypes.--usnm 13473 (2 specimens, now lost) from chiriquí, panamá; mr. macneil collector]; bull. u.s. natl. mus., 32:14, 1887. günther, biologia-centrali americana, reptilia and batrachia, p. 265, june, 1901. dunn, occas. papers boston soc. nat. hist., 5:413, october 10, 1931; occas. papers boston soc. nat. hist., 8:72, june 7, 1933. stebbins and hendrickson, univ. california publ. zool., 56:524, february 17, 1959. fouquette, evolution, 14:484, december 16, 1960. busack, copeia, 2:371, june 21, 1966. ? _hyla cherrei_ cope, proc. acad. nat. sci. philadelphia, 1894, p. 195, 1894 [holotype.--location unknown, apparently lost; type-locality: "alajuela, costa rica;" r. alfaro collector]. günther, biologia centrali-americana: reptilia and batrachia, p. 264, june, 1901. taylor, univ. kansas sci. bull., 35:846, july 1, 1952. _hyla underwoodi_, ruthven, misc. publ. mus. zool., univ. michigan, 8:55, september 15, 1922. barbour, proc. new england zool. club, 10:31, march 2, 1928. _hyla microcephala microcephala_, smith, herpetologica, 7:185, december 31, 1951. taylor, univ. kansas sci. bull., 39:23, november 18, 1958. _diagnosis._--brown lateral stripe narrow, extending from nostril along canthus, along upper edge of tympanum to groin, bordered above by narrow white line; pair of dark brown longitudinal lines on dorsum extending to vent; shanks having dark longitudinal line or flecks, no transverse bars; interorbital dark mark lacking. _description and variation._--the color pattern is nearly constant. of 103 males from the canal zone, all lack an interorbital dark bar, and all have a dark longitudinal line on the dorsal surface of the shank and a narrow lateral dark stripe, bordered above by a narrow white line, extending to the groin. the longitudinal dark lines on the dorsum are continuous to the groin in 95 specimens and fragmented in two specimens. in two others the lines converge and fuse in the scapular region, and in four specimens auxiliary, fragmented lines are present dorsolaterally. in all specimens from southeastern costa rica (golfito, palmar sur, and villa neilly) the pattern is constant, except that in about 10 per cent of the specimens the longitudinal line on the dorsal surface of the shank is replaced by a row of brown flecks. of the limited number of colombian specimens examined, all are patterned normally, except three from sautata, chocó, three from curumani, and three from arcataca, magdalena, which have flecks on the dorsal surfaces of the shanks, and one from espinal, tolima, which has no markings on the shanks. when active at night most individuals are pale yellowish tan dorsally; the white dorsolateral line is noticeable, but the brown lateral stripe, dorsal brown lines, and lines on shanks are so pale that often they are barely discernible. by day the dorsum changes to tan or pale reddish brown; the stripes are dark brown, and the dorsolateral stripe that is white at night becomes creamy yellow (pl. 13). small brown flecks are present on the dorsum of most individuals. the venter always is white, and the iris is pale bronze with a brown tint immediately anterior and posterior to the pupil. in breeding males the vocal sac is pale yellow. _tadpoles._--tadpoles of this species have been found in weed-choked ponds in eastern panamá province. the following description is based on ku 104097, a specimen in developmental stage 34 (gosner, 1960). total length, 20.5 mm.; body length, 8.2 mm.; body slightly wider than deep; snout pointed; nostrils large, situated dorsally, much closer to snout than eyes, directed anteriorly; eyes moderately small, situated dorsolaterally and directed laterally; spiracle sinistral, located just posteroventral to eye; anal tube dextral. tail xiphicercal; caudal musculature moderately deep, becoming slender posteriorly, extending beyond caudal fin; fins deepest at about one-third distance from body to tip of tail; dorsal fin extending onto body, deeper than deepest part of caudal musculature; ventral fin slightly shallower than musculature. mouth small, terminal, lacking teeth and fringing papillae, but having finely serrate beaks. in preservative, top of head pale brown; dark stripe from tip of snout through eye to posterior edge of body, narrowing to thin line on proximal one-fourth of tail; venter white; tail creamy tan with fine black flecks most numerous posteriorly; posterior two-thirds of fins edged with black. in life, top of head yellowish tan; lateral stripe brown; belly white; anterior half of tail lacking pigment; posterior half deep orange; iris pale bronze (pl. 15). _remarks._--evidence for intergradation of _hyla microcephala_ with _h. underwoodi_ is provided by four specimens [usc 818 (2), 6081-2] from 6.1 kilometers northeast of the mouth of the río tarcoles, and nine specimens [usc 8254 (2), 8255, 8256 (4), 8258 (2)] from parrita, both in puntarenas province, costa rica. these localities lie about two-thirds the distance from the northwesternmost locality for _h. m. microcephala_ (palmar sur) to the southeasternmost locality for _h. m. underwoodi_ (barranca). although in most aspects of coloration the frogs are more nearly like _h. m. underwoodi_ than _h. m. microcephala_, some specimens have longitudinal lines on their shanks, such as are characteristic of _h. m. microcephala_. the dorsal pattern varies from nearly complete longitudinal lines to broken lines, fused into an x-shaped scapular mark or not. as noted by rivero (1961:135), _hyla microcephala_ seems to be closely related to _hyla misera_ werner, a species having a wide distribution east of the andes in south america. despite the similarity in color pattern, size, and structure, we are reluctant to place the two taxa in the same species until data on coloration in life, mating calls, and life history are available for _hyla misera_ and compared with those of _hyla microcephala_. the status of cope's _hyla cherrei_ is questionable. since the type, the only specimen ever referred to the species, apparently is lost, the only extant information regarding the taxon is contained in the original description (cope, 1894). there the species was characterized as having a narrow dorsolateral white stripe and lacking pigment on the upper arms and thighs. these characteristics of the color pattern combined with the statements "vomerine teeth few, opposite the middle of the very large choanae" and "tympanic drum distinct, one half the area of eye" serve to distinguish _h. cherrei_ from all other costa rican hylids, except _h. m. microcephala_ and _h. m. underwoodi_. no statements in the type description will definitely associate _cherrei_ with one or the other of these subspecies. since it seems obvious that _h. cherrei_ can be associated with _h. microcephala_, we prefer to place the name in the synonymy of the nominate subspecies, thereby preserving the commonly used name _h. underwoodi_ (boulenger, 1899) as a subspecies of _h. microcephala_. _distribution._--_hyla microcephala microcephala_ inhabits coastal lowlands from the area of golfo dulce (apparently absent from the osa peninsula) in southeastern costa rica eastward in panamá, including the azuero peninsula to northern colombia and thence southward in the valleys of the río cauca and río magdalena in colombia (fig. 1). except for the central area of the canal zone the subspecies is unknown from the caribbean drainage in central america, but in colombia the subspecies occurs only in the caribbean drainage. in central america this frog occurs mostly on the coastal lowlands; the highest recorded elevation is 560 meters at el valle, coclé, panamá. throughout most of its range _hyla microcephala microcephala_ occurs in disturbed habitats--cut-over forests, secondary growth, and pastureland. it does not seem to be an inhabitant of either primary forest or of _curatella_-savanna. _specimens examined._--522, as follows: +costa rica+: puntarenas: golfito, ku 32172-207; 3 km. e golfito, ku 86399, usc 2757-8; palmar sur, ku 64591-608, usc 2650 (14), uu 3907-32; *1.5-2.5 km. ese palmar sur, ku 68293-7 (skeletons), 93957-62; parrita, usc 8254 (2), 8255, 8256 (4), 8258 (2) [intergrades with _h. m. underwoodi_]; 3 km. nw piedras blancas, ku 103689; 6.1 km. ne mouth of río tárcoles, usc 818 (2), 6081-2 [intergrades with _h. m. underwoodi_]; villa neilly, usc 2651; *1-5 km. wnw villa neilly, usc 6182-4, 8003 (4), 8031 (3), 8032; *10.5 km. wnw villa neilly, ku 64609-27, 68398 (eggs). +panama+: canal zone: albrook air base, tnhc 23389, 23497; balboa, ansp 19555-6; *fort clayton, uimnh 42008-12; *2.8 km. sw fort kobbe, ku 96015-25; *frijoles, mcz 19208; *bamboa, mcz 21507; *8.3 km. n gatún locks, tnhc 23441; *juan diaz, mcz 13747; *juan mina, amnh 55436-7, ansp 21811-2, ummz 126734, 126735 (6), uu 3900-6; *8-14 km. n miraflores locks, tnhc 23374-88, 23390-409, 23411-38, 23440, 23442-60, 23462-76; 23478-83, 23492, 23555-60, 23562-76; *río chagres, amnh 55430, 55439; *río cocolí, 3.5 km. n miraflores locks, tnhc 23410; *summit, ansp 23365-71, fmnh 22966-9, ku 97783-87. chiriqui: 5.5 km. e concepción, amnh 69772; *14.4 km. e concepción, amnh 69773-8; 2 km. s david, amnh 69779; *progreso, ummz 58252, 58253 (2), 58254, 58436; río gariché, 8.3 km. ese paso canoas, ku 103065-8. coclé: 1 km. se el caño, ku 103042-51; el valle de antón, amnh 59614-18 (10), 69785, ansp 23502-5, ku 77201-14, mvz 66578-83, uimnh 46532. colón: cement plant, transisthmian highway, fmnh 60394-5. darién: el real, ku 80454-5, 103052-64, ummz 125036 (10), usnm 140567-8; río canclon at río chucunaque, ummz 125035; *río chucunaque, near yavisa, amnh 59523. los santos: tonosí, ku 101606-9. panamá: 5 km. s bejuco, amnh 69782; 3 km. w chepo, ku 77172-4, 104097-8 (tadpoles); *6 km. wsw chepo, ku 77175; *chico, río la jagua, usnm 129070; *la joya, cacora, ansp 25129-33; madden dam, fmnh 67819; nueva gorgona, amnh 69780-1; *1.6 km. w nueva gorgona, amnh 69783-4; 1.5 km. w pacora, 77176-200; *río la laja, near chamé, ansp 21845; *río tapia, mcz 10048; *tapia, amnh 18930, 18950, 18952-3; *18 km. e tocumen, mvz 78662. +colombia+: chocó: sautatá, atrato, fmnh 74918 (2), 74919. magdalena: aracataca, ansp 19755-7; curumani, mcz 21465-74, uimnh 28855; ummz 90168, usnm 118247; el banco, río magdalena, ansp 25061; fundación, ummz 48281-2. tolima: espinal, mcz 15068; mariquita, fmnh 81822-3. valle: sevilla, mcz 13751-3. _hyla microcephala underwoodi_ boulenger _hyla microcephala_ boulenger, proc. zool. soc. london, p. 481, october 1, 1898 [syntypes.--bmnh 94. 11. 1532-33 from bebedero, guanacaste province, costa rica; c. f. underwood collector] (not _hyla microcephala_ cope, proc. amer. philos. soc., 23:281, february 11, 1886, from chiriquí, panamá). _hyla underwoodi_ boulenger, ann. mag. nat. hist., ser. 7, 3:277, april, 1899 (substitute name for _hyla microcephala_ boulenger, preoccupied). günther, biologia-centrali americana, reptilia and batrachia, p. 278, september, 1901. dunn and emlen, proc. acad. nat. sci. philadelphia, 84:25, march 22, 1932. stuart, misc. publ. mus. zool., univ. michigan, 29:39, october 1, 1935. taylor, proc. biol. soc. washington, 50:44, april 21, 1937. stuart, occas. papers mus. zool., univ. michigan, 471:15, may 17, 1943. taylor and smith, proc. u. s. natl. mus., 95:586, june 30, 1945. stuart, misc. publ. mus. zool., univ. michigan, 69:35, june 12, 1948. smith and taylor, bull. u. s. natl. mus., 194:85, june 17, 1948; univ. kansas sci. bull., 33:316, march 20, 1950. stuart, contr. lab. vert. biol., univ. michigan, 45:48, may, 1950. taylor, univ. kansas sci. bull., 35:891, july 1, 1952; univ. kansas sci. bull., 39:25, november 18, 1958. _hyla phlebodes_, cole and barbour, bull. mus. comp. zool., 50:154, november, 1906. kellogg, bull. u. s. natl. mus., 160:172, march 31, 1932. _hyla microcephala martini_ smith, herpetologica, 7:187, december 31, 1951 [holotype.--uimnh 20965 from encarnacion, campeche, méxico; h. m. smith collector]. stuart, contr. lab. vert. biol., univ. michigan, 68:46, november, 1954. fugler and webb, herpetologica, 13:105, july 10, 1957. stuart, contr. lab. vert. biol., univ. michigan, 75:17, june, 1958. neill and allen, publ. research div., ross allen's reptile inst., 2:26, november 10, 1959. duellman, univ. kansas publ., mus. nat. hist., 13:62, august 16, 1960. stuart, herpetologica, 17:74, july 11, 1961. hensley and smith, herpetologica, 18:70, april 9, 1962. stuart, misc. publ. mus. zool., univ. michigan, 122:36, april 2, 1963. holman and birkenholz, herpetologica, 19:144, july 3, 1963. duellman, univ. kansas publ., mus. nat. hist., 15:225, october 4, 1963; univ. kansas publ., mus. nat. hist., 15:588, june 22, 1965. _hyla microcephala underwoodi_, smith, herpetologica, 7:188, december 31, 1951. _diagnosis._--brown lateral stripe narrow, extending to groin or only to sacral region, bordered above by narrow white line; dorsal pattern bold, consisting of xor )(-shaped mark in scapular region or pair of interconnected dark lines on back; interorbital dark mark usually present; shanks usually having dark transverse bars. _description and variation._--the dorsal color pattern is highly variable. the various permutations of the x-shaped scapular mark and dark sacral marks differ proportionately in different samples. the variation in color pattern in 12 samples is summarized in table 2. in samples from the southern part of the range (southern nicaragua and guanacaste province, costa rica) more (40-93%) individuals have the lateral stripes extending to the groin than in northern samples (0-42%) from southern méxico and guatemala. likewise, the percentage of specimens lacking bars on the shanks and a dark interorbital bar is higher in the costa rican samples than elsewhere in the range. the xor )(-shaped scapular markings and /\or / \-shaped sacral markings are most prevalent in northern samples, whereas to the south the dorsal markings are more commonly arranged in a pattern of paired lines, which usually are discontinuous and usually extend posteriorly only to the sacral region. thus, the color pattern in _h. m. underwoodi_ in the southern part of its range shows trends towards the pattern characteristic of _h. m. microcephala_. intergrades between these two subspecies have been discussed in the account of the nominate subspecies. table 2.--variation in color pattern in hyla microcephala underwoodi ========================================================================== population | n | shanks || interorbital || dorsolateral | | | || bar || stripe | | |-------------||----------------||--------------| | | bars |flecks|| present| absent|| groin| sacrum| -------------------------------------------------------------------------oaxaca: | 27 | 22 | 5 || 27 | 0 || 0 | 27 | donají-sarabia | | | || | || | | | | | || | || | | tabasco: | 55 | 46 | 9 || 55 | 0 || 0 | 55 | teapa-villahermosa| | | || | || | | | | | || | || | | guatemala: | 51 | 51 | 0 || 51 | 0 || 17 | 34 | la libertad | | | || | || | | | | | || | || | | guatemala: | 32 | 32 | 0 || 32 | 0 || 0 | 32 | finca chamá | | | || | || | | | | | || | || | | guatemala: | 31 | 31 | 0 || 31 | 0 || 14 | 17 | puerto barrios | | | || | || | | | | | || | || | | honduras: | 13 | 13 | 0 || 13 | 0 || 9 | 4 | lago yojoa | | | || | || | | | | | || | || | | nicaragua: | 56 | 44 | 12 || 54 | 2 || 13 | 43 | la cumplida | | | || | || | | | | | || | || | | nicaragua: | 10 | 10 | 0 || 10 | 0 || 8 | 2 | tipitapa | | | || | || | | | | | || | || | | nicaragua: | 10 | 10 | 0 || 10 | 0 || 8 | 2 | santo thomás | | | || | || | | | | | || | || | | costa rica: | 12 | 0 | 12 || 6 | 6 || 7 | 5 | tenorio-tilarán | | | || | || | | | | | || | || | | costa rica: | 38 | 21[a]| 15 || 34 | 4 || 25 | 13 | las cañas-liberia | | | || | || | | | | | || | || | | costa rica: | 32 | 26 | 6 || 29 | 3 || 30 | 2 | esparta | | | || | || | | -------------------------------------------------------------------------========================================================================== population | scapular markings || sacral | | || markings | |----------------------------||----------------------| | x | )( | ][ | other || /\ | / \ | other | -------------------------------------------------------------------------oaxaca: | 23 | 4 | 0 | 0 || 7 | 6 | 14 | donají-sarabia | | | | || | | | | | | | || | | | tabasco: | 53 | 2 | 0 | 0 || 19 | 11 | 23 | teapa-villahermosa| | | | || | | | | | | | || | | | guatemala: | 45 | 6 | 0 | 0 || 16 | 14 | 21 | la libertad | | | | || | | | | | | | || | | | guatemala: | 32 | 0 | 0 | 0 || 26 | 2 | 4 | finca chamá | | | | || | | | | | | | || | | | guatemala: | 23 | 0 | 4 | 4 || 6 | 4 | 21 | puerto barrios | | | | || | | | | | | | || | | | honduras: | 3 | 2 | 3 | 5 || 2 | 1 | 10 | lago yojoa | | | | || | | | | | | | || | | | nicaragua: | 11 | 35 | 8 | 2 || 0 | 19 | 37 | la cumplida | | | | || | | | | | | | || | | | nicaragua: | 0 | 5 | 3 | 2 || 0 | 3 | 7 | tipitapa | | | | || | | | | | | | || | | | nicaragua: | 3 | 0 | 7 | 0 || 0 | 5 | 5 | santo thomás | | | | || | | | | | | | || | | | costa rica: | 0 | 0 | 12 | 0 || 0 | 0 | 12 | tenorio-tilarán | | | | || | | | | | | | || | | | costa rica: | 0 | 11 | 19 | 8 || 0 | 0 | 38 | las cañas-liberia | | | | || | | | | | | | || | | | costa rica: | 0 | 0 | 14 | 18 || 0 | 0 | 32 | esparta | | | | || | | | -------------------------------------------------------------------------[footnote a: longitudinal stripes present in two specimens.] when this frog is active at night its dorsum is pale yellow; faint flecks are present in some individuals. the white dorsolateral line usually is evident in the tympanic region, but in many individuals a dorsal pattern of lines and other marks is not evident. by day the dorsum changes to yellowish tan or pale brown with dark brown or reddish brown markings (pl. 13). the venter is white, and the vocal sac in breeding males is yellow. the iris is pale bronze with a brown tint anterior and posterior to the pupil. _remarks._--_hyla microcephala underwoodi_ has had a confused nomenclatural history. the taxon was first named _hyla microcephala_ by boulenger (1898); this name was preoccupied by _hyla microcephala_ cope (1886). cole and barbour (1906) and kellogg (1932) used the name _hyla phlebodes_ stejneger (1906) for specimens of this frog from méxico. dunn (1931, 1933, 1934) applied the name _hyla underwoodi_ to panamanian specimens that we identify as _hyla phlebodes_. smith (1951) named _hyla microcephala martini_ from southern méxico and guatemala and considered the northern populations to represent a subspecies distinct from the costa rican _hyla microcephala underwoodi_, despite the fact the stuart (1935:39) stated that comparisons of specimens from el petén, guatemala, with the holotype of _hyla underwoodi_ showed only trivial differences. much of the confusion regarding the name _hyla underwoodi_ stems from the illustration given by boulenger (1898:pl. 39, fig. 3) and reproduced by taylor (1952:892), which shows a frog having a unicolor dorsum, dorsolateral white lines, and dark flanks. this pattern is in marked contrast to the pattern seen in most preserved specimens, which have the dorsum variously marked by dark brown lines or irregular marks. smith (1951:185), in his description of _hyla microcephala martini_ from southern méxico, considered _h. underwoodi_ to be a subspecies of _h. microcephala_ that lacked dorsal dark markings. data accumulated in 1961 through field studies by the senior author at the type locality, bebedero, and other localities in guanacaste and puntarenas provinces in costa rica provide a reasonable explanation of the differences in color pattern. as noted in the preceding description of this subspecies, at night the dorsal markings are not evident in many living individuals, whereas by day the dorsal markings are prominent. most collectors prepare their specimens by day; consequently the majority of specimens have a pronounced dorsal pattern. of the frogs collected in costa rica in 1961, some specimens were preserved at night; others from the same series were preserved by day. the differences are striking. in those preserved at night, dorsal markings are faint, if present at all. some specimens closely match the figure given by boulenger (1898). it is extremely doubtful if the frog described and illustrated by boulenger could be associated with either _hyla phlebodes_ or _h. microcephala microcephala_. individuals of the former species lack a dorsolateral white line and always have some dorsal markings evident at night; furthermore, _h. phlebodes_ is not known to occur on the pacific lowlands. _hyla microcephala microcephala_ occurs farther southeast. since there is no reason to doubt the type locality of _h. underwoodi_, since specimens from the area around the type locality that have been preserved at night are like the holotype in pattern, and since the characteristics of the populations of the frogs in guanacaste are the same as, or gradually blend into those of, populations in northern central america and southern méxico, the frogs from throughout the entire range can be referred to one taxon, the earliest name for which is _hyla underwoodi_ boulenger, which herein is considered to be a subspecies of _h. microcephala_ cope. _distribution._--_hyla microcephala underwoodi_ inhabits the atlantic slopes and lowlands from southern veracruz and extreme northern oaxaca eastward across the base of the yucatan peninsula (possibly the species is extant in the northern part of the peninsula) to british honduras and thence southeastward through the caribbean lowlands and interior valleys in honduras to central nicaragua, where it apparently avoids the forested caribbean lowlands and the dry pacific lowlands of northwestern nicaragua, but in the vicinity of managua invades the pacific lowlands and continues southward into northwestern costa rica as far as the puntarenas peninsula (fig. 1). in méxico and guatemala the species has not been taken at elevations of more than 350 meters, whereas farther south it occurs at higher elevations--780 meters at silencio, costa rica, 830 meters on montaña de guaimaca, honduras, 960 meters at finca tepeyac, nicaragua, and 1200 meters at finca venecia, nicaragua. _specimens examined._--1270, as follows: +mexico+: campeche: balchacaj, fmnh 100406, uimnh 20944-6; encarnación, fmnh 27069-70, 75784, mcz 28360, 29637, uimnh 20948-58, 20965, usnm 134264-5; escárcega, ummz 122999; *7.5 km. w escárcega, ku 71229-43; laguna alvarado, 65 km. s xpujil, ku 75084-9; pacaitún, río candelaria, fmnh 83118-20; *tres brazos, fmnh 113101-22, uimnh 20947; 10 km. w xpujil, ku 75082-3. chiapas: palenque, uimnh 47984, 49139-50, usnm 114973-8. oaxaca: *5 km. n chiltepec, ku 87015-23; 3 km. n donají ummz 115249 (9); *3.7 km. n donají, ummz 115250 (5); *43 km. n matías romero, uimnh 42550-68; *3.5 km. n palomares, tnhc 25185, 25321-31, 25341-68; 4.6 km. n sarabia, ummz 115247 (2); *6.1 km. n sarabia, ummz 115248 (11), *3 km. n tolocita, ku 39655; tuxtepec, ku 87024-40. tabasco: 24 km. n frontera, mcz 35665-70; 0.8 km. e río tonolá, tnhc 25189; teapa, ummz 119218 (4); *2.7 km. n teapa, ummz 119216 (4); *10 km. n teapa, ummz 119217 (6); *11.5 km. n teapa, ummz 119219; *15.2 km. n teapa, ummz 119220 (4); *17.6 km. n teapa, ummz 119221 (12), 3.3 km. s villahermosa, ummz 119215 (12), *17.6 km. s villahermosa, ummz 119214 (12). veracruz: 2.1 km. n acayucan, uimnh 42547-9; *6.4 km. nw acayucan, ummz 115254 (14); 1.6 km. ese alvarado, ummz 115258 (39); *2.4 km. ese alvarado, ummz 115251 (2); *4.5 km. s aquilera, ummz 115252 (21); *8 km. sw coatzacoalcos, ummz 119213 (10); 2.2 km. e cosoleacaque, ummz 119222 (26); 10 km. se hueyapan, ummz 115255; 0.8 km. s lerdo de tejada, ummz 122778; *3.6 km. ne minatítlán, tnhc 25150-2; 1.9 km. s naranja, ummz 115253 (3); 4.5 km. ne novillero, ummz 115256; san andrés tuxtla, fmnh 113124-8, uimnh 20942-3. yucatán: chichén-itzá, fmnh 36570, mcz 2463 (2). +british honduras+: cayo: 6.2 km. s el cayo, mcz 37885-92. stann creek: stann creek, fmnh 49068. +guatemala+: alta verapaz: 28.3 km. n campur, ku 64578-90; chinajá, ku 57425; cubilquitz, ummz 90887, 90888 (4); finca chamá, ummz 90879 (13), 90880 (4), 90881, 90882 (28), 90883 (12), 90884 (46), 90885 (39), 90886 (20); *finca tinaja, byu 16032; panzós, ummz 90889 (2). chiquimula: chiquimula, ummz 98113; 2 km. n esquipulas, ummz 106844. el petén: la libertad, ku 57447-97, 59907-11 (skeletons), mcz 21461, ummz 75332 (13), 75333 (11), 75334 (14), 75335 (10); piedras negras, fmnh 113123, uimnh 20966; *5 km. s piedras negras, usnm 114951-72; tikal, ummz 117981 (2); toocog, 15 km. se la libertad, ku 57426-46. el quiché: finca tesoro, ummz 89165 (5). huehuetenango: finca san rafael, 16 km. se barillas, fmnh 40917-9. izabal: puerto barrios, fmnh 20004-7; 8 km. s puerto barrios, ku 57507-37, 59991 (eggs), 59992 (tadpoles); quirigua, cas 69657-701; 2.5 km. ne río blanco, ku 57539; san felípe, fmnh 35065. zacapa: 14 km. ene mayuelas, ku 57502-6; 8 km. ene río hondo, ku 57498-501. +honduras+: atlantidad: la ceiba, ummz 91948 (2), usnm 117593-600; lancetilla, mcz 17981. cortes: lago yojoa, amnh 54917-9, 54957, 55134, ku 64563-77. el paraiso: valle de jamastran, amnh 54807-12. francisco-moranza: el zamorano, amnh 54873-81, ku 103223, ummz 123101; montaña de guaimaca, amnh 54900-4 (8); ranch san diego, 19 km. sw guaimaca, amnh 53939. itibucá: vieja itibucá, amnh 54912-3. +nicaragua+: chontales: 3 km. sw santo tomás, ku 64770-9, 68308 (skeleton). esteli: finca venecia, 7 km. n, 16 km. e condega, ku 85296; 2.4 km. n estelí, mcz 28933-7. managua: 12-13 km. e managua, ku 85297-301; *10 km. sw tipitapa, ummz 119977 (10). matagalpa: *finca tepeyac, 10.5 km. n, 9 km. e matagalpa, ku 85302-3; hacienda la cumplida, ku 64780-96, 68309-11 (skeletons), ummz 116482 (8), 116483 (23), 116484 (3), 116485 (5), 119984 (3). rivas: *finca amayo, 13 km. s, 14 km. e rivas, ku 85304-7; 16 km. s rivas, mcz 29011-7; *20.5 km. se rivas, ku 85308-10; 5 km. se san pablo, ku 43111-4. +costa rica+: guanacaste: arenal, usc 6254 (2); *3 km. w bagaces, usc 7019 (10); *3 km. ne boca del barranca, usc 8017 (21), *finca san bosco, usc 6272 (6), 6276 (3); *guayabo de bagaces, usc 7022 (4), 7023 (3), 7025; 12 km. s la cruz, usc 8091 (2); *laguna arenal, usc 6262; *27 km. n las cañas, usc 8171 (6); *16 km. e las cañas, ku 102252-8; 16 km. sse las cañas, ku 65090-5; *20 km. se las cañas, ku 102251; liberia, ku 30827-39; *7.3 km. n liberia, usc 8096 (4); *10 km. n liberia, usc 8085 (9); *7.5 km. se liberia, ku 65102-8, 68621-2 (skeletons); *14.7 km. s liberia, usc 8238 (3); *4 km. w liberia, ku 36847-57; 2 km. s nicoya, usc 8230; *3-10 km. ese playa del coco, usc 8012 (16), 8137 (14); *21.6 km. ese playa del coco, usc 8138 (13); *peñas blancas, ku 102247-50; *río bebedero, 5 km. s bebedero, ku 65089; *río higuerón, usc 7168 (2); santa cruz, usc 8232 (2); *silencio, usc 6248; *tenorio, ku 32313; tilarán, ku 36858-60; *2 km. e tilarán, ku 86403, *5 km. ne tilarán, ku 36840-6 usc 6269. puntarenas: barranca, ku 32305-12, *5 km. wnw barranca, ummz 119976 (2); *10 km. e esparta, ku 86400-2; 1 km. wnw esparta, ku 65101; *4 km. wnw esparta, ku 65088; *10 km. wnw esparta, ku 65063-87, 68616-20 (skeletons); *12 km. wnw esparta, ku 65096-100, usc 8251; 21.8 km. w san ramón, usc 8242 (15). +hyla robertmertensi+ taylor _hyla robertmertensi_ taylor, proc. biol. soc. washington, 50:43, april 21, 1937 [holotype.--cnhm 100096 (formerly eht-hms 2270) from tapachula, chiapas, méxico; h. m. smith and e. h. taylor collectors]. smith and taylor, bull. u. s. natl. mus., 194:84, june 17, 1948; univ. kansas sci. bull., 33:326, march 20, 1950. mertens. senckenbergiana, 33:170, june 15, 1952; senckenbergischen naturf. gesell., 487:30, december 1, 1952. stuart, contr. lab. vert. biol., univ. michigan, 68:47, november, 1954. duellman, univ. kansas publ., mus. nat. hist., 13:63, august 16, 1960. duellman and hoyt, copeia, 1961 (2): 417, december 19, 1961. porter, herpetologica, 18:168, october 17, 1962. stuart, misc. publ. mus. zool., univ. michigan, 122:36, april 2, 1963. duellman and trueb, univ. kansas publ., mus. nat. hist., 17:348, july 14, 1966. _diagnosis._--brown lateral stripe wide, including loreal region and entire tympanum, extending to groin, bordered above by narrow white line; dorsum unicolor or with pair of dark lines (or rows of dashes) usually extending only to the sacral region; shanks having dark flecks, no transverse bars; interorbital bar lacking. _description and variation._--males attain a maximum snout-vent length of 26.4 mm. in oaxaca, whereas in a sample from acacoyagua, chiapas, the largest male has a snout-vent length of 25.7 mm., and from la trinidad, guatemala, 24-6 mm. specimens from the western part of the range (eastern oaxaca) have slightly smaller heads and proportionately larger tympani than the more eastern populations (table 1). the color pattern shows little variation, except in the nature of the dorsal markings. in a few specimens from throughout the range, but especially in the eastern part of the range, the dorsum lacks markings between the dorsolateral white lines. in most specimens the dorsal pattern consists of flecks or dashes arranged in two parallel longitudinal rows, and in some specimens the marks are fused into parallel lines. small brown flecks are present on the dorsal surfaces of the shanks; in some specimens these flecks tend to form a longitudinal stripe on the shank. an interorbital dark mark is invariably absent. when active at night _hyla robertmertensi_ is pale yellow above with a white dorsolateral line and pale brown lateral stripe; the dorsal markings are faint. by day the dorsum is yellowish tan with brown markings. the dorsolateral stripe is creamy white, and the lateral stripe is dark brown (pl. 14). the venter is white, and the iris is dull bronze. in breeding males the vocal sac is yellow. _remarks._--although this species superficially resembles _hyla microcephala microcephala_, the latter is easily distinguished by the narrow brown lateral stripe, as compared with the much wider stripe in _h. robertmertensi_. no other hylids in northern central america and southern méxico can be confused with this species. _distribution._--_hyla robertmertensi_ inhabits the pacific slopes (to elevations of 700 meters) and lowlands from eastern oaxaca (east of the plains of tehuantepec) southeastward to central el salvador. the species also occurs in the cintalapa valley (atlantic drainage) in southwestern chiapas (fig. 2.) the distribution seems to be limited on the northwest and southeast by arid environments. the region in which _hyla robertmertensi_ lives is characterized by higher rainfall and more luxuriant vegetation than occur on the plains of tehuantepec or on the pacific lowlands of eastern el salvador and southern honduras. in addition to the localities listed below, mertens (1952:30) recorded the species from hacienda cuyan-cuya, depto. sonsonate, el salvador. [illustration: fig. 2. map showing locality records for _hyla robertmertensi_.] _specimens examined._--490, as follows: +mexico+: chiapas: acacoyagua, usnm 114754-61; *2 km. w acacoyagua, ummz 87843 (28), 87844 (50), 87845 (50), 87846 (45), 87847 (27), 87848 (3); 32 km. n arriaga, ku 57619-24, 59917-8 (skeletons); asunción, fmnh 100413, 100501-4, uimnh 26989-90, usnm 134267; *la esperanza, usnm 114737-48, 114750-3, 17 km. s las cruces, ku 57625-49, 59997 (eggs); 8.5 km. n puerto madero, ummz 119981 (2); *11.7 km. n puerto madero, ummz 119982; tapachula, fmnh 100096, uimnh 26987; *11 km. s tapachula, ku 57605-18, 59916 (skeleton); tonolá, fmnh 27073, 100505-10, uimnh 26988. oaxaca: tapanatepec, ummz 115245 (2), *1.6 km. e tapanatepec, ummz 115244 (14); *4.3 km. e tapanatepec, uimnh 38368-9; *7.5 km. w tapanatepec, ummz 115246 (39); 12.8 km. w tapanatepec, ku 65007-14; 7.2 km. wnw zanatepec, ummz 115243 (77); *13.6 km. wnw zanatepec, tnhc 25213-22; 22.7 km. wnw zanatepec, tnhc 25203-9. +guatemala+: jutiapa: jutiapa, ummz 106848; la trinidad, ummz 107733 (23). retalhueleu: casa blanca, ummz 107732. +el salvador+: la libertad: 16 km. nw santa tecla, ku 44112. san salvador: 21.9 km. n san salvador, ummz 119983 (6). +hyla phlebodes+ stejneger _hyla phlebodes_ stejneger, proc. u. s. natl. mus., 30:817, june 4, 1906 [holotype.--usnm 2997 from "san carlos," costa rica; burgdorf and schild collectors]. taylor, proc. biol. soc. washington, 50:44, april 21, 1937; univ. kansas sci. bull., 35:888, july 1, 1952; univ. kansas sci. bull., 39:25, november 18, 1958. fouquette, evolution, 14:484, december 16, 1960. duellman and trueb, univ. kansas publ., mus. nat. hist., 17:348, july 14, 1966. _hyla underwoodi_, dunn, occas. papers boston soc. nat. hist., 5:413, october 10, 1931; occas. papers boston soc. nat. hist. 8:72, june 7, 1933; amer. mus. novitiates, 747.2, september 17, 1934, gaige, hartweg, and stuart, occas. papers mus. zool., univ. michigan, 357:5, october 26, 1937. breder, 1946, bull. amer. mus. nat. hist., 86:416, august 22, 1946. _diagnosis._--dark brown lateral stripe, if present, usually extending only to insertion of forearm, never posteriorly to sacral region; white line above brown stripe absent or faint; dorsal pattern weak, usually consisting of irregular dashes or interconnected lines; interorbital dark mark present; shanks having weakly defined transverse bars. _description and variation._--in the majority of specimens (70%) the lateral dark stripe extends from the nostril to the eye and thence above the tympanum to a point above the insertion of the arm; in 17 per cent the stripe extends to the mid-flank, whereas in 13 per cent the stripe is absent. a narrow and faint white line is present on the canthus in some specimens, but no distinct white stripe is present above the lateral dark line posterior to the eye. an interorbital bar and transverse marks on the shanks are invariably present. the dorsal markings are variable, but in most specimens (92%) consist of either an xor )(-shaped mark in the scapular region; in the other specimens the markings are irregular short lines or absent. approximately equal numbers of specimens have a transverse bar, chevron, or broken lines in the sacral region, whereas about eight per cent of the specimens lack markings in the sacral region. when active at night, individuals are pale yellowish tan with faint brown dorsal markings. by day they are tan with more distinct brown markings (pl. 14). the thighs are pale yellow; the belly is white. the iris is pale creamy tan with brown flecks. in breeding males the vocal sac is yellow. _tadpoles._--tadpoles of this species have been found in an extensive grassy pond at puerto viejo, costa rica. the following description is based on ku 104099, a specimen in development stage 36 (gosner, 1960). total length, 21.0 mm.; body length, 6.7 mm.; body slightly wider than deep, snout pointed; nostrils large, directed anteriorly, situated near end of snout; eyes small, situated dorsolaterally, directed laterally; spiracle sinistral, located just posteroventral to eye; anal tube dextral. tail xiphicercal; caudal musculature moderately deep, extending far beyond posterior edge of fins; fins deepest at about midlength; dorsal fin extending onto body, slightly deeper than caudal musculature; ventral fin slightly shallower than musculature. mouth small, terminal, lacking teeth and fringing papillae, but having finely serrate beaks. in preservative top of head olive-tan with brown flecks; dark stripe from snout through eye to posterior edge of body; belly white, flecked with brown anteriorly; tail creamy tan with grayish brown blotches. in life, dorsum of body reddish tan mottled with darker brown; lateral stripe dark brown; belly white, mottled with brown and black; caudal musculature heavily pigmented with grayish tan; posterior tip of tail marked with dark gray; caudal fins heavily blotched with grayish tan; iris orange-tan peripherally, red centrally (pl. 15). _remarks._--this species has been confused with _hyla microcephala underwoodi_ by many workers. dunn (1931, 1933, 1934) and breder (1946) referred panamanian specimens of _h. phlebodes_ to _h. underwoodi_; likewise, gaige, hartweg, and stuart (1937) made the same error. cole and barbour (1906) and kellog (1932) used the name _h. phlebodes_ for mexican specimens of _h. microcephala underwoodi_. the similarity in color pattern of _h. microcephala underwoodi_ and _h. phlebodes_ easily accounts for the misapplication of names. although both species have nearly identical dorsal color patterns, that of _h. microcephala underwoodi_ usually is bolder. furthermore, in that species a narrow white line usually is present above the well-defined lateral dark stripe, whereas the lateral dark stripe is short and posterior to the eye is not bordered above by a white line in _h. phlebodes_. the type locality "san carlos, costa rica" given by stejneger (1906:817) apparently refers to a region, the llanuras de san carlos, in the northern part of alajuela province, costa rica. [illustration: fig. 3. map showing locality records for _hyla phlebodes_.] _distribution._--_hyla phlebodes_ inhabits humid tropical forests from southeastern nicaragua southeastward on the caribbean slopes and lowlands to the canal zone in panamá, thence eastward in the chucunaque basin of eastern panamá and onto the pacific lowlands of colombia (fig. 3). the species also reaches the pacific slopes in the arenal depression in northwestern costa rica and in the panamanian isthmus, where it occurs in humid forests on the pacific slope of el valle and cerro la campana. mostly the species is found at low elevations, but it occurs at 600 meters at turrialba and at 700 meters at finca san bosco in costa rica. _specimens examined._--410, as follows: +nicaragua+: zelaya: isla grande del maíz, mcz 14848; río mico, el recrero, ummz 79720 (6). +costa rica+: alajuela: 12.4 km. n florencia, mvz 76108-10, usc 2628; *las playuelas, 11 km. s los chiles, usc 7216; los chiles, usc 7217, 7219; 3 km. ne muelle de arenal, usc 2644 (2); *"san carlos," usnm 29970. cartago: chitaría, ku 103690; *1.6 km. e río reventazón bridge, east of turrialba, ummz 119978 (2); *tunnel camp, near peralta, ku 32456, 32458-69, 41098 (skeleton); turrialba, fmnh 101794, 103188-9, ku 25725-9, 32439-48, 41095-7 (skeletons), 64797-827, 68300-2 (skeletons), 68403 (eggs), 68404 (tadpoles), mcz 29224-5, 29310-2, ummz 119979 (6), usc 31, 256 (2), 458 (2), 580, 594, 599 (7), 7074 (2), usnm 29933. guanacaste: arenal, usc 6254; *finca san bosco, usc 62724, 6276 (3), guayabo de bagaces, usc 7022 (3), 7023; *laguna arenal, usc 6262 (4); 3 km. ne tilarán, usc 524; *5 km. ne tilarán, usc 6269; *6 km. ne tilarán, ummz 122653 (6), s-2680 (skeleton), usc 523 (8). heredia: puerto viejo, ku 64828-63, 68303-7 (skeletons), 68405-6 (tadpoles), 104099-100 (tadpoles); *1.5 km. n puerto viejo, ku 64871; *1 km. s puerto viejo, ku 86432-40; *4.2 km. w puerto viejo, ku 64864-5; *5.9 km. w puerto viejo, ku 64866-70; *7.5 km. w puerto viejo, ku 86431. limón: batán, ummz 119980 (2); la castilla, ansp 23707; puerto +limón+, ku 32449-55. +panama+: bocas del toro: 3.2 km. nw almirante, ku 96026; cayo de agua, ku 96027-31; fish creek, ku 96032-4. canal zone: barro colorado island, amnh 69790, ansp 23244-50; fmnh 13380, 22972-4; juan mina, amnh 55429, uu 3899; *8.6-13.8 km. n miraflores locks, tnhc 23439, 23477, 23484-8, 23491, 23494-9, 23501-2, 23504-8, 23510-17, 23519-30, 23532-8, 23541-54, 23561. *rio chagres, amnh 55431-4; río cocolí, 3.5 km. n miraflores locks, tnhc 23461, 23489-90, 23493, 23500, 23503, 23509, 23518, 23531, 23539-40; *summit, ansp 23361, ku 97788; *three rivers plantation, su 2130. coclé: el valle de antón, amnh 55435, 69786-9, ansp 23506-9. colón: achiote, ku 77215-78; ciricito, cas 71499-500, 71505-6. darién: río canclon at río chucunaque, ummz 126733; río chucunaque, near yavisa, amnh 51783. panamá: cero la campana, fmnh 67847-50. +colombia+: chocó: andagoya, fmnh 81856; boca de raspadura, amnh 13570-8. +hyla sartori+ smith _hyla underwoodi_ (in part), smith and taylor, bull. u. s. natl. mus., 194:85, june 17, 1948. _hyla microcephala sartori_ smith, herpetologica, 7:186, december 31, 1951 [holotype.--uimnh 20934 from 1 mile north of organos, south of el treinte, guerrero, méxico; h. m. smith and e. h. taylor collectors]. duellman, univ. kansas publ., mus. nat. hist., 15:124, december 20, 1961. porter, herpetologica, 18:168, october 17, 1962. davis and dixon, herpetologica, 20:230, january 25, 1965. duellman, univ. kansas publ. mus. nat. hist., 15:652, december 30, 1965. _diagnosis._--dorsum tan with broad dark brown chevrons or transverse bars; shanks marked with two or three broad transverse bars; dorsolateral stripes absent. _description and variation._--no noticeable geographic variation is apparent in either structural features or coloration in this species. all specimens lack a dorsolateral dark stripe and white line, although a dark line is present on the canthus and dissipates in the loreal region. a broad interorbital brown bar is present in all specimens. the color pattern on the dorsum invariably consists of a broad, dark, chevron-shaped mark in the scapular region and a broad dark chevron or transverse bar in the sacral region. the shanks invariably have two or three dark brown transverse bars. when active at night individuals are yellowish tan above with chocolate brown markings (pl. 14). the belly is white, and the thighs are pale yellowish tan. the iris is dark bronze-color. in breeding males the vocal sac is yellow. by day some individuals were observed to change to creamy gray with distinct darker markings. _remarks._--although tadpoles of this species have not been found, observations on the breeding sites indicate that the tadpoles probably develop in ponds. except for calling males observed around a pool in a stream-bed 11.8 kilometers west-northwest of tierra colorada, guerrero, all breeding congregations have been found at temporary ponds. smith (1951:186) named _hyla sartori_ as a subspecies of _hyla microcephala_. this subspecific relationship seemed reasonable until analysis of the mating calls showed that the call of _h. sartori_ is more nearly like that of _h. phlebodes_ than that of _h. microcephala_. the broad hiatus separating the ranges of _h. microcephala_ and _h. sartori_ is additional evidence for considering _h. sartori_ as a distinct species. [illustration: fig. 4. map showing locality records for _hyla sartori_.] _distribution._--_hyla sartori_ occurs in mesophytic forests to elevations of about 300 meters on the pacific slopes of southern méxico from southwestern jalisco to south-central oaxaca (fig. 4). the lack of specimens from colima and michoacán probably reflects inadequate collecting instead of the absence of the species there. on the basis of available habitat the species would be expected to occur in nayarit, but extensive collecting there has failed to reveal its presence. the semi-arid plains of tehuantepec apparently limit the distribution to the east. _specimens examined._--190, as follows: +méxico+: guerrero: 5 km. e acapulco, amnh 54611-2; 23.2 km. n acapulco, uimnh 26404-7; colonia buenas aires, 23 km. e tecpán de galeana, ummz 119223 (7); *el limoncito, fmnh 75785, 100390-402, 104631, 104633, ummz 117250, usnm 134266; el treinte, fmnh 100403, uimnh 20935-7; laguna coyuca, amnh 59686; la venta, mcz 29635; *morjonares, uimnh 26392-402; 1.6 km. n organos, fmnh 100404-5, uimnh 20933-4; 19.2 km. s petaquillas, uimnh 26408; 6.1 km. e. tecpán de galeana, tnhc 23396-408; *11.2 km. n tierra colorada, uimnh 26403; 11.8 km. wnw tierra colorada, ummz 119225 (51), s-2677-9 (skeletons); zacualpán, ummz 119224 (6). jalisco: 6.4 km. ne la resolana, ku 67853-69; 24 km ne la resolana, ku 67870-3. oaxaca: 3 km. n pochutla, ku 57539; 13.4 km. n pochutla, ummz 123495 (40). cranial osteology the frogs of the _hyla microcephala_ group have a minimal amount of cranial ossification as compared to more generalized hylid skulls, such as _smilisca_ (duellman and trueb, 1966). in the _hyla microcephala_ group the sphenethmoid is small and short, and a large frontoparietal fontanelle is present. the quadratojugal exists only as a small spur and is not in contact with the maxillary. the proötics are poorly developed. the anterior and posterior arms of the squamosal are short; the anterior arm extends no more than one-fourth of the distance to the maxillary, and the posterior arm does not have a bony connection with the proötic. the nasal lacks a maxillary process, and the medial ramus of the pterygoid lacks a bony connection to the proötic. teeth are absent on the parasphenoid and palatines, but present on the maxillaries, premaxillaries, and prevomers. the teeth are simple, pointed, and slightly curved. although the number of teeth varies (table 3), no consistent differences between the species are apparent. table 3.--variation in the number of teeth in the species of the hyla microcephala group. (n=number of jaws, or twice the number of individuals; means are given in parentheses after the observed ranges). ========================+====+=============+==============+========== species | n | maxillary | premaxillary | prevomer ------------------------+----+-------------+--------------+---------_h. microcephala_ | 32 | 31-47(37.8) | 4-13(8.9) | 2-4(3.2) | | | | _h. phlebodes_ | 10 | 38-45(40.1) | 8-13(10.3) | 2-5(3.9) | | | | _h. robertmertensi_ | 6 | 23-43(32.8) | 7-12(10.5) | 2-3(2.7) | | | | _h. sartori_ | 6 | 27-43(38.2) | 9-10(9.3) | 3-4(3.7) ------------------------+----+-------------+--------------+--------- [illustration: plate 13 upper figure, _hyla microcephala microcephala_ (ku 64593); middle figure, _h. microcephala underwoodi_ (ku 64565); lower figure, _h. microcephala underwoodi_ (ummz 115247). all approximately ×3.] [illustration: plate 14 upper figure, _hyla robertmertensi_ (ummz 115243); middle figure, _h. phlebodes_ (ku 64798); lower figure, _h. sartori_ (ummz 119225). all approximately ×3.] [illustration: plate 15 tadpoles of _hyla microcephala_ group: upper figure, _h. m. microcephala_ (ku 104097); lower figure, _h. phlebodes_ (ku 104099). both ×4.] [illustration: plate 16 audiospectrograms and sections of mating calls of _hyla microcephala_ group: (a) _h. m. microcephala_ (ku tape no. 19); (b) _h. robertmertensi_ (ku tape no. 41); (c) _h. phlebodes_ (ku tape no. 6); (d) _h. sartori_ (ku tape no. 190).] table 4.--comparative cranial osteology of hyla microcephala group ===============+=======================+========================+ character | _h. microcephala_ | _h. robertmertensi_ | ---------------+-----------------------+------------------------+ frontoparietal | minimally ossified | ossification extensive | | with large fontanelle | anteriorly with narrow | | extending from | medial separation; | | sphenethmoid to | fontanelle largest in | | occipital ridge. | parietal region. | | | | | | | nasals | moderately long and | moderate in size; | | slender; arcuate in | slightly wider | | dorsal view. | anteriorly than | | | posteriorly in dorsal | | | view. | | | | sphenethmoid | extremely short in | moderately short in | | dorsal view. | dorsal view. | | | | | | | | | | columella | distal and greatly | distal and slightly | | expanded. | expanded or not. | ---------------+-----------------------+------------------------+ table 4. (continued) ===============+========================+======================== character | _h. phlebodes_ | _h. sartori_ ---------------+------------------------+-----------------------frontoparietal | ossification extensive | ossification moderately | anteriorly with narrow | extensive anteriorly; | medial separation; | medial separation of | fontanelle largest in | about uniform width | parietal region. | throughout length of | | fontanelle. | | nasals | moderate in size; | long and broad; | slightly wider | arcuate in dorsal | anteriorly than | view. | posteriorly in dorsal | | view. | | | sphenethmoid | moderately short in | moderately short in | dorsal view. | dorsal view; ossified | | anteriorly between | | nasals. | | columella | distal and not | distal and not | expanded. | expanded. ---------------+------------------------+----------------------- [illustration: fig. 5. dorsal views of the skulls of (a) _hyla m. microcephala_ (ku 68293) and (b) _h. sartori_ (ummz s-2677). both × 12.] [illustration: fig. 6. dorsal views of skulls of (a) _hyla phlebodes_ (ku 68303) and (b) _h. robertmertensi_ (ku 59917). both × 12.] despite the great reduction in the ossification of the cranial elements, certain apparently consistent differences exist between the species seem to be consistent. the most notable differences are: 1) amount of ossification of the frontoparietals and consequent shape and size of the frontoparietal fontanelle, 2) shape of the nasals, 3) shape and extent of the sphenethmoid, and 4) shape of the columella (table 4, figs. 5-6). on the basis of these characters, _hyla microcephala_ can be set apart from the other species and characterized as having a poorly ossified frontoparietal and correspondingly large frontoparietal fontanelle; long, slender, arcuate nasals; extremely short sphenethmoid; and expanded distal end of the columella. the other species in the group (_phlebodes_, _robertmertensi_, and _sartori_) have more ossification of the frontoparietals, broader nasals, only a moderately short sphenethmoid, and an unexpanded distal end of the columella. among these three species, the skulls of _phlebodes_ and _robertmertensi_ are most nearly alike, whereas the skull of _sartori_ differs by having a differently shaped frontoparietal fontanelle, broader nasals, and an ossified anterior extension of the sphenethmoid between the nasals (compare fig. 5b with fig. 6 a-b). although all skulls examined belong to breeding adults, the extent of the ossification of the frontoparietals and the resulting shape of the frontoparietal fontanelle might be correlated with the age of the frog. nevertheless, in the 24 skulls of _hyla microcephala_ examined, the frontoparietals are less extensively ossified than in the skulls of the other species. the trivial differences among the other three species certainly are suggestive of close relationship, but on the basis of present knowledge of the evolutionary trends in hylid cranial osteology, the differences offer little evidence for determining phylogenetic lineage. analysis of mating calls calls of all five taxa were compared in several characteristics, of which three are deemed most significant systematically. these are 1) the pattern and duration of the notes of a call-group, 2) the fundamental frequency, and 3) the dominant frequency. air temperatures were noted at the time the calls were recorded, but no valid correlation could be determined between this factor and any of the parameters of the calls; consequently recordings made at all temperatures (21-29° c.) were grouped together. _pattern and duration of notes._--in all five taxa the basic pattern consists of a call-group made up of one primary note followed by a series of shorter secondary notes. in some species the secondary notes differ from the primary in other characteristics. both subspecies of _hyla microcephala_ have a long, unpaired primary note followed by 0 to 18 (usually about 4) somewhat shorter paired secondary notes. in calls of _hyla m. microcephala_ the mean duration of the primary is 0.131 (0.10-0.16) second and that of the secondaries is 0.101 (0.05-0.14) second, whereas in _h. m. underwoodi_ the mean duration of the primary is 0.018 (0.05-0.15) second and that of the secondaries is 0.086 (0.06-0.11) second. _hyla robertmertensi_ has a reverse of this pattern in that the primary note is paired and the secondaries are unpaired. in the sample studied a call-group contains 0-28 secondary notes (generally about 3). the mean duration of the primary is 0.091 (0.07-0.11) second and that of the secondaries is 0.040 (0.025-0.06) second. _hyla phlebodes_ and _sartori_ have call-groups composed of a rather short, unpaired primary and several short, unpaired secondaries (0-28 in _phlebodes_, 0-23 in _sartori_). the mean duration of the primary of _phlebodes_ is 0.105 (0.07-0.16) second and that of the secondaries is 0.067 (0.035-0.12) second. the mean duration of the primary of _sartori_ is 0.080 (0.07-0.09) second and that of the secondaries is 0.053 (0.035-0.07) second. the two subspecies of _h. microcephala_ are identical in call pattern and agree closely in duration of notes, although those of the nominate subspecies tend to be slightly longer. _hyla robertmertensi_ is distinctive in call pattern in that it is the only species having a paired primary; the duration of the primary is completely overlapped by that in the other species, but the secondaries tend to be the shortest in the group. the call patterns of _h. phlebodes_ and _h. sartori_ are identical and the range of duration of notes of _phlebodes_ completely overlaps that of _sartori_, although both the primary and secondary notes of the latter tend to be somewhat shorter (table 5, pl. 16). _fundamental frequency._--this parameter was analyzed for the primary notes. it was measured for the secondaries as well and was found to differ in magnitude in the same way as the primary note. in a few examples of both subspecies of _h. microcephala_ a high primary note, in which the fundamental frequency is exceptionally high, is sometimes emitted (fouquette, 1960b). none of these notes was used in this analysis; only the fundamental frequencies of normal primary notes are compared (table 5, fig. 7). table 5.--comparison of normal mating calls in the hyla microcephala group. (observed range given in parentheses below mean; unless otherwise noted data are for primary notes.). ----------------+--+---------+---------+-------------------+------------- | |dominant | funda |duration of notes | repetition | | | mental| (seconds) | rate of species |n |frequency|frequency+---------+---------+ secondaries | | (cps) | (cps) | primary |secondary|(notes/minute) ----------------+--+---------+---------+---------+---------+-------------_h. m. |44| 5637 | 205 | 0.13 | 0.10 | 268 microcephala_ | |(5150 |(184-244)|(0.11 |(0.05 | (192-353) | | -5962)| | -0.16)| -0.14)| | | | | | | _h. m. |47| 5772 | 220 | 0.11 | 0.09 | 283 underwoodi_ | |(5177 |(192-275)|(0.05 |(0.06 | (197-384) | | -6200)| | -0.15)| -0.11)| | | | | | | _h. |25| 5388 | 162 | 0.09 | 0.04 | 418 robertmertensi_| |(5150 |(140-178)|(0.07 |(0.03 | (368-570) | | -5785)| | -0.11)| -0.06)| | | | | | | _h. phlebodes_ |34| 3578 | 148 | 0.11 | 0.07 | 284 | |(3220 |(125-158)|(0.07 |(0.04 | (210-350) | | -4067)| | -0.16)| -0.12)| | | | | | | _h. sartori_ |10| 3217 | 126 | 0.08 | 0.05 | 434 | |(2950 |(116-135)|(0.07 |(0.04 | (396-477) | | -3600)| | -0.09)| -0.07)| ----------------+--+---------+---------+---------+---------+-------------the two subspecies of _h. microcephala_ agree closely in fundamental frequency. there is considerable overlap, but the difference between the means is significant at the 0.001 level of probability (t = 4.2406). the call of _h. robertmertensi_ does not overlap that of _h. sartori_ or either subspecies of _h. microcephala_ in this parameter; but it does overlap that of _h. phlebodes_, although again the difference between the means is significant at the 0.001 level (t = 9.360). _hyla phlebodes_ and _sartori_ have the lowest fundamental frequencies, and there is some overlap, but here too the difference between the means is significant at the 0.001 level (t = 4.923). _dominant frequency._--a dominant band of frequencies cuts across the harmonics of the fundamental, obscuring the harmonic pattern and generally shifting upward in frequency. the midpoint of this band is measured at the terminal border as the dominant frequency. as with the fundamental frequency, only the normal primary notes were utilized in the comparisons (table 5, fig 8). [illustration: fig. 7. variation in the fundamental frequency of the normal primary notes in the _hyla microcephala_ group. the horizontal lines = range of variation, vertical lines = mean, solid bars = twice the standard error of the mean, and open bars = one standard deviation. the number of specimens in each sample is indicated in parentheses after the name of the taxon.] the two subspecies of _h. microcephala_ agree more closely in this parameter than in fundamental frequency. the overlap is great, but the difference between the means is significant at the 0.001 level (t = 3.658). the calls of both subspecies completely overlap that of _robertmertensi_ in this parameter, but the difference between the means is significant at the 0.001 level. the calls of _h. phlebodes_ and _h. sartori_ overlap considerably in this characteristic, although the difference between the means is significant at the 0.001 level (t = 7.504) (fig. 9). the call of neither species overlaps those of _h. microcephala_ and _robertmertensi_. [illustration: fig. 8. variation in the mid-point of the dominant frequency band of the normal primary notes in the _hyla microcephala_ group. the horizontal lines = range of variation, vertical lines = mean, solid bars = twice the standard error of the mean, and open bars = one standard deviation. the number of specimens in each sample is indicated in parentheses after the name of the taxon.] [illustration: fig. 9. scatter diagram relating the dominant and fundamental frequencies of the normal primary notes in the _hyla microcephala_ group. each symbol represents a different individual.] _repetition rate._--the repetition rate of the secondary notes, in calls consisting of more than one secondary, was measured for each form. a considerable amount of variation in this parameter was found in all of the taxa (table 5). this variation probably is due in part to the effect of temperature differences. repetition rate is the only parameter analyzed for which there is a correlation with the air-temperature, but even here the correlation is weak, probably due to the microenvironmental effects of humidity, air-movement, and other factors in addition to the ambient air temperature that influences the body temperature of the frogs. these rates are nearly alike in both subspecies of _h. microcephala_ and in _phlebodes_. the repetition rates in _h. robertmertensi_ and _h. sartori_ are considerably faster than in the other three taxa. _hyla sartori_ has the fastest repetition rate of the group. in all characteristics of the mating calls the two subspecies of _h. microcephala_ agree closely, as might be expected, although the differences are statistically significant. _hyla robertmertensi_ is distinctive in call pattern and seems to be closer to _microcephala_ in dominant frequency but closer to _h. phlebodes_ in fundamental frequency. thus, it is somewhat intermediate between _microcephala_ and _phlebodes_. the identical pattern and similarity in fundamental and dominant frequencies of the calls of _h. phlebodes_ and _h. sartori_ possibly indicate close relationship. _geographic variation in call._--_hyla m. microcephala_ has higher fundamental and dominant frequencies in costa rica than in panamá. in costa rican _h. m. underwoodi_ the fundamental and dominant frequencies are lower than in other parts of the range. frogs of this subspecies recorded in nicaragua and honduras have slightly lower dominant frequencies and higher fundamental frequencies than those recorded in guatemala or oaxaca. the duration of both primary and secondary notes decreases to the south; samples from nicaragua and costa rica have the shortest notes. comparison of duration of notes in the two subspecies shows that the panamanian _h. m. microcephala_ have slightly longer notes than do any _h. m. underwoodi_; the more northern populations of _h. m. underwoodi_ from méxico most closely approach _h. m. microcephala_ in this characteristic. the calls of _h. robertmertensi_ in oaxaca have higher dominant and fundamental frequencies and longer secondary notes than do those in chiapas. the calls of _h. phlebodes_ recorded at puerto viejo, costa rica, have slightly lower dominant frequencies than do those recorded at turrialba, costa rica, and in panamá, whereas those recorded at turrialba have lower fundamental frequencies than in other samples. the duration of notes is slightly shorter in both costa rican samples than in those recorded in panamá. life history the frogs of the _hyla microcephala_ group breed in shallow grassy ponds. in some places they breed in permanent ponds, but usually congregate around temporary pools, such as depressions in forests, flooded fields, and roadside ditches. at the height of their breeding season, usually in the early part of the rainy season, the congregations are made up of large numbers of individuals. in april, 1961, and in june, 1966, the senior author noted nearly continuous choruses of _h. m. microcephala_ in roadside ditches along the 75 kilometers of road between villa neily and palmar sur, puntarenas province, cost rica; on june 20, 1966, at puerto viejo, heredia province, costa rica, he estimated approximately 900 _hyla phlebodes_ in one pond, and two nights later noticed that the number of individuals had increased substantially. other observations by the first author on size of breeding congregations include nearly continuous choruses of _h. m. underwoodi_ between villahermosa and teapa, tabasco, in july of 1958, an estimated 400 _hyla robertmertensi_ in a road side ditch 7.2 kilometers west-northwest of zanatepec, oaxaca, on july 13, 1956, and approximately 150 _hyla sartori_ around a rocky pool in a riverbed, 11.8 kilometers west-northwest of tierra colorada, guerrero, on june 28, 1958. the length of the breeding season seemingly is more dependent on climatic conditions in various parts of middle america than on behavioral differences in the various species. thus, fouquette (1960b) found in the canal zone that _h. m. microcephala_ formed breeding choruses from may through january, the entire rainy season in that area. in the wetter coastal region of puntarenas province, costa rica, the species breeds as early as mid-march, whereas in the drier region encompassing guanacaste province, costa rica, and southwestern nicaragua breeding activity is initiated by the first heavy rains of the season, usually in june. _hyla phlebodes_ inhabits regions having rainfall throughout the year. although large breeding congregations are most common in the early parts of the rainy season, males probably call throughout the year. at puerto viejo in costa rica the senior author has heard _hyla phlebodes_ in february, april, june, july, and august. charles w. myers noted calling males of this species in the area around almirante, bocas del toro province, panamá, in september, october, and february. an exception to the correlation between rainfall and breeding activity was noted by the junior author in _hyla phlebodes_ in the canal zone, where he noticed a decrease in activity of that species in october and november, when the rains are heaviest and most frequent. furthermore, independent observations made by both of us indicate that _h. phlebodes_ does not reach peaks of activity during or immediately after heavy rains, but instead builds up to peaks of activity two or three days after a heavy rain. this is in contrast to the other species, all of which characteristically inhabit drier environments than does _h. phlebodes_. peaks of breeding activity in the other species occur immediately after, or even during, heavy rains. the calling location of the males generally is on vegetation above, or at the edge of, the water. _hyla microcephala_ and _h. phlebodes_ call almost exclusively from grasses and sedges; _phlebodes_ usually calls from taller and more dense grasses than does _microcephala_. except for some minor differences in calling location observed by the junior author (fouquette, 1960b) in the canal zone, the differences in density and height of grasses utilized for calling-locations probably is dependent primarily on the nature of the available vegetation. although bushes and broad-leafed herbs are usually present at the breeding sites, males of these species seldom utilize them for calling locations. both _h. robertmertensi_ and _h. sartori_ have been observed calling from grasses, herbs, bushes, and low trees. calling males of _robertmertensi_ have been found two meters above the ground in small trees. daytime retreats in the breeding season sometimes are no more than shaded clumps of vegetation adjacent to a pond or in clumps of grass in a pond. individuals of _h. m. underwoodi_ were found by day under the outer sheaths of banana plants next to a water-filled ditch. dry season refuges are unknown. amplexus is axillary in all four species. egg deposition has been observed in _h. m. microcephala_, _m. underwoodi_, and _phlebodes_. in all three the eggs are deposited in small masses that float near the surface of the water and usually are at least partly attached to emergent vegetation. each clutch does not represent the entire egg complement of the female. tadpoles are definitely known of only _h. m. microcephala_ and _phlebodes_; these have been described in the preceding accounts of the species. the tadpoles of these two species can be distinguished readily (pl. 15). the tadpole of _h. microcephala_ has a uniformly white venter and nearly transparent tail, whereas in _h. phlebodes_ the venter is flecked anteriorly and the tail is mottled. in life, _h. microcephala_ is easily recognized by the orange posterior half of the tail, whereas the tail in _h. phlebodes_ is mottled tan and grayish brown. phylogenetic relationships the evidence already presented on osteology, external structure, coloration, mating call, and life history emphatically show that the four species under consideration are a closely related assemblage. now the question arises: to what other groups in the genus is the _hyla microcephala_ group related? furthermore, it is pertinent to this discussion to attempt a reconstruction of the phylogeny of the group as a whole and of the individual species in the _hyla microcephala_ group. with regard to the relationships of the group we must take into account certain species in south america. our endeavors there are hampered by the absence of data on the mating calls and life histories of most of the relevant species. as mentioned in the account of _hyla m. microcephala_, the species _microcephala_ possibly is subspecifically related to _hyla misera_, a frog widespread in the amazon basin. _hyla misera_ resembles _microcephala_ in coloration, external structure, and cranial characters. the frontoparietals are equally poorly ossified, and the frontoparietal fontanelle is extensive. our principal reason for not considering the two taxa conspecific at this time is our lack of knowledge concerning the color of living _h. misera_, the structure of the tadpoles, and the characteristics of the mating call. even with the absence of such data that we think essential to establish the nomenclature status of the taxa, we are confident that the two are sufficiently closely related that any discussion of the phylogenetic relationships of one species certainly must involve consideration of the other. _hyla misera_ possibly is allied to other small yellowish tan south american _hyla_ that lack dark pigmentation on the thighs. probable relatives are _hyla elongata_, _minuta_ (with _goughi_, _pallens_, _suturata_, _velata_, and possibly others as synonyms), _nana_, and _werneri_. the consideration of the interspecific relationships of these taxa is beyond the scope of this paper, but we can say that each of these species has a pale yellowish tan dorsum, relatively broad dorsolateral brown stripe, and narrow longitudinal brown lines or irregular marks on the dorsum. furthermore, examination of the skulls of _elongata_, _nana_, and _werneri_ reveals that they are like _misera_ and _microcephala_ in the nature of the frontoparietal fontanelle and in having a greatly reduced quadratojugal. thus, on the basis of cranial and external characters the _hyla microcephala_ group can be associated with _hyla misera_ and its apparent allies in south america. this association can be only tentative until the mating calls, tadpoles, and chromosome numbers of the south american species are known. among the middle american hylids, only the _hyla microcephala_ group and _h. ebraccata_ have a haploid number of 15 chromosomes (duellman and cole, 1965). all other new world _hyla_, for which the number is known, have a haploid number of 12; the only other _hyla_ having 15 is a papuan _hyla angiana_ (duellman, 1967). _hyla ebraccata_ occurs in the humid tropical lowlands of middle america and the pacific lowlands of northwestern south america. it is the northernmost, and only central american, representative of the _hyla leucophyllata_ group, which is diverse (about 10 species currently recognized) and widespread in tropical south america east of the andes. this group is characterized by having broad, flat skulls with larger nasals and more ossification of the frontoparietals than in the _hyla microcephala_ group. the quadratojugal is present as a small anteriorly projecting spur that does not connect with the maxillary. externally, the _hyla leucophyllata_ group is characterized by having a well-developed axillary membrane, uniformly yellow thighs, and a dorsal color pattern in many species consisting of a dark lateral band, a pale dorsolateral band or dorsal ground color, and a large middorsal dark mark. in some species, the dorsal pattern consists of small dark markings or is nearly uniformly pale. at least in the central american _hyla ebraccata_, the mating call consists of a single primary note followed by a series of shorter secondary notes, the tadpoles have xiphicercal tails and lack teeth, and the haploid number of chromosomes is 15. on the strength of these observations it seems imperative to consider the _hyla leucophyllata_ group as a close ally to the _hyla microcephala_ group. successful artificial hybridization supports the close relationship of _h. m. microcephala_ and _phlebodes_; partial success of artificial hybridization of these two with _ebraccata_ (fouquette, 1960b) provides further evidence for close relationship between the _hyla leucophyllata_ and _hyla microcephala_ groups. in méxico and northern central america two small species, _hyla picta_ and _hyla smithi_, comprise the _hyla picta_ group. these frogs resemble members of the _hyla microcephala_ group by having a yellowish tan dorsum with a dorsolateral white stripe and uniformly yellow thighs. furthermore the mating call is not unlike those of the species in the _hyla microcephala_ group. despite these similarities, the _hyla picta_ group differs from the _hyla microcephala_ group by having a well-developed quadratojugal that connects to the maxillary, tadpoles with teeth present and caudal fins completely enclosing the caudal musculature, and a haploid number of 12 chromosomes. in all of these characteristics the frogs of the _hyla picta_ group more closely resemble other middle american _hyla_ than they do the _hyla microcephala_ group. therefore, it can best be presumed that the superficial resemblances of coloration and the mating call are the result of convergence. since the _hyla microcephala_ and _leucophyllata_ groups apparently are related and since the greatest diversity of these frogs is in south america (if _hyla misera_ and its relatives are placed with the _hyla microcephala_ group), it seems appropriate to place the centers of origins of these groups in south america. therefore, the _hyla microcephala_ group and _hyla ebraccata_ of the _hyla leucophyllata_ group either have immigrated into central america, or they are representatives of those groups that were isolated in central america during most of the cenozoic when south america was separated from central america. the interspecific relationships of the species in the _hyla microcephala_ group are not clear. on the basis of coloration, _h. m. microcephala_ and _h. robertmertensi_ are close, and _h. m. underwoodi_ and _h. phlebodes_ are nearly identical. the mating calls of _h. phlebodes_ and _sartori_ closely resemble one another, whereas the call of _robertmertensi_ is intermediate between these and _microcephala_. in most respects _hyla microcephala_ is distinct from the other species, and with the exception of the amount of ossification of the frontoparietals, the other species can be easily derived from a _microcephala_-like ancestor. possibly the slightly increased ossification of the frontoparietals in _robertmertensi_, _phlebodes_, and _sartori_ is secondary, or possibly after differentiation of the species the amount of ossification was further reduced in _microcephala_. if so, the species fall into a reasonable phylogenetic scheme that has _microcephala_ as the extant species most like the ancestral stock. we visualize the evolutionary history of the group to have followed a course that began with the invasion of central america by a _microcephala_ ancestral stock that differentiated into two populations in lower central america--a _microcephala_-like frog on the pacific lowlands and a _phlebodes_-like frog on the caribbean lowlands. differentiation could have been brought about by isolation by montaine or marine barriers. the population on the pacific lowlands either was preadapted for subhumid conditions or became so adapted and dispersed northward onto the pacific lowlands of northern central america. simultaneously the frogs on the caribbean lowlands, which were adapted to humid environments, dispersed northward in the humid forested regions to southern méxico and crossed the isthmus of tehuantepec onto the pacific slopes of oaxaca and guerrero northward to jalisco. subsequent development of arid conditions, possibly in the pliocene, pleistocene, or even as late as the thermal maximum in post-wisconsin time, resulted in a restriction of the ranges in northern central america, thereby isolating part of the _phlebodes_-stock on the pacific slopes of méxico, where it adapted to drier conditions and evolved into _sartori_. the rest of the _phlebodes_-stock was restricted to the humid forests on the caribbean lowlands of lower central america. the increased aridity on the pacific lowlands eliminated the _microcephala_-stock from southern honduras and northwestern nicaragua and in so doing left an isolated population on the lowlands of chiapasand guatemala, which differentiated into _robertmertensi_. the original stock on the pacific lowlands of panamá and southeastern costa rica became _microcephala_. if the _microcephala_-stock was, as we believe, better adapted for existence under subhumid conditions than was the _phlebodes_-stock, the development of subhumid conditions in much of the lowland region of northern central america and southern méxico would have permitted the expansion of the range of _microcephala_ into the area now inhabited by _h. m. underwoodi_, while _phlebodes_ was being eliminated from this area by climatic conditions that were unsuited to its survival there. perhaps the similarity in coloration of _h. m. underwoodi_ and _phlebodes_ is the result of convergence or possibly hybridization occurred at the time the former was expanding its range and the latter's range was being restricted. if hybridization did occur, the differences in mating call subsequently were enhanced, thereby providing a valid isolating mechanism in sympatric populations. _hyla microcephala_ and _phlebodes_ range into northern south america. probably both species entered south america in relatively recent times after they had differentiated from one another in central america. literature cited boulenger, g. a. 1898. fourth report on additions to the batrachian collection in the natural-history museum. proc. zool. soc. london, 1898, pp. 373-482, pls. 38-39. october 1. 1899. descriptions of new batrachians in the collection of the british museum (natural history). ann. mag. nat. hist, ser. 7, 3:273-277, pls. 11-12. breder, c. m. jr. 1946. amphibians and reptiles of the rio chucunaque drainage, darien, panama, with notes on their life histories and habits. bull. amer. mus. nat. hist, 86:375-436, pls. 42-60, august 26. cole, l. j. and barbour, t. 1906. vertebrata from yucatan: reptilia; amphibia; pisces. bull. mus. comp. zool., 50:146-159. november. cope, e. d. 1886. thirteenth contribution to the herpetology of tropical america. proc. amer. philos. soc, 23:271-287. february 11. 1894. third addition to a knowledge of the batrachia and reptilia of costa rica. proc. acad. nat. sci. philadelphia, 1894, pp. 194-206. duellman, w. e. 1956. the frogs of the hylid genus _phrynohyas_ fitzinger, 1843. misc. publ. mus. zool., univ. michigan, 96:1-47, pls. 1-6. february 21. 1967. additional studies of chromosomes of anuran amphibians. syst. zool., 16:38-43, march 17. duellman, w. e. and cole, c. j. 1965. studies of chromosomes of some anuran amphibians (hylidae and centrolenidae). syst. zool., 14:139-143. july 9. duellman, w. e. and trueb, l. 1966. neotropical hylid frogs, genus smilisca. univ. kansas publ., mus. nat. hist., 17:281-375, pls. 1-12. july 14. dunn, e. r. 1931. the amphibians of barro colorado island. occas. papers boston soc. nat. hist., 5:403-421. october 10. 1933. amphibians and reptiles from el valle de anton, panamá. _ibid._, 8:65-79. june 7. 1934. two new frogs from darien. amer. mus. novit., 747:1-2. september 17. fouquette, m. j. jr. 1960a. call structure in frogs of the family leptodactylidae. texas jour. sci., 12:201-215. october. 1960b. isolating mechanisms in three sympatric tree frogs in the canal zone. evolution, 14:484-497. december 16. gaige, h. t., hartweg, n. and stuart, l. c. 1937. notes on a collection of amphibians and reptiles from eastern nicaragua. occas. papers mus. zool., univ. michigan, 357:1-18. october 26. gosner, k. l. 1960. a simplified table for staging anuran embryos and larvae with notes on identification. herpetologica, 16:183-190. september 23. kellogg, r. 1932. mexican tailless amphibians in the united states national museum. bull. u.s. natl. mus., 160:1-224. march 31. rivero, j. a. 1961. salientia of venezuela. bull. mus. comp. zool., 126:1-207. november. smith, h. m. 1951. the identity of _hyla underwoodi_ auctorum of mexico. herpetologica, 7:184-190. december 31. stejneger, l. 1906. a new tree toad from costa rica. proc. u. s. natl. mus., 30:817-818. june 4. stuart, l. c. 1935. a contribution to a knowledge of the herpetology of a portion of the savanna region of central petén, guatemala. misc. publ. mus. zool., univ. michigan, 29:1-56, pls. 1-4. october 4. taylor, e. h. 1952. the frogs and toads of costa rica. univ. kansas sci. bull., 35-577-942. july 1. _transmitted july 11, 1967._ transcriber's notes this file was derived from scanned images. with the exception of the list of typographical errors that were corrected below, the original text is presented. in the copy of the original, the plate text contains the notation "x 2" after the caption to let the reader know that the image was enlarged by a factor of two. emphasis notation _text_ = italic +text+ = bold typographical errors corrected: several minor typographical corrections were made (missing periods, commas, incomplete italicization, etc.); but are not indicated here. more substantial changes are listed below: page 533 umz => ummz page 534 diganosis => diagnosis page 544 fontanells => fontanelle page 545 prrimary => primary page 547 band of of frequencies => band of frequencies page 550 ad => had page 551 clumbs => clumps page 552 acount => account page 557 minchigan => michigan _occasional papers_ of the museum of natural history the university of kansas lawrence, kansas -------------------------------------------------------------- number 1 april 29, 1971 -------------------------------------------------------------- a synopsis of neotropical hylid frogs, genus _osteocephalus_ by linda trueb[1] and william e. duellman[2] [1] research associate, division of herpetology, museum of natural history, university of kansas. [2] curator, division of herpetology, museum of natural history, university of kansas. when we initiated a study of the herpetofauna at santa cecilia in amazonian ecuador in 1966, we were immediately confronted with many kinds of animals that we could not identify with the existing literature. comparisons of our specimens with those preserved in other museums resolved some of the problems, but many identifications could be made only after study of type specimens; even then some determinations remained questionable. we now find that in order to prepare a meaningful account of the herpetofauna of santa cecilia, we must complete several taxonomic studies, the limits of which extend far beyond eastern ecuador. because of our interests in hylids we have begun our studies on these frogs. one of us (trueb, 1970a) studied the cranial osteology of casque-headed hylid frogs and redefined the genus osteocephalus but did not determine the number of species in the genus. our work in amazonian ecuador resulted in the discovery of the sympatric occurrence of three species at each of two localities; one of these species was found with a fourth species at another locality. study of museum specimens confirmed the recognition of these four species in the amazon basin and lower amazonian slopes of the andes. a fifth species from bolivia and perú also is included in the genus. examination of museum specimens has provided data on the geographic variation in, and distribution of, the five species. however, our conclusions pertaining to some populations need substantiation, because we have been hampered by inadequate material from areas beyond ecuador. more than half of the 905 specimens of _osteocephalus_ are from ecuador, a relatively small part of the total range of the genus. in this paper we are presenting a taxonomic review of the genus _osteocephalus_; of necessity our study has been at the alpha level. we have utilized all of the usual external characters, as well as osteological features in our definitions of the species. tadpoles and mating calls are available for only one species, _o. verrucigerus_ (trueb and duellman, 1970); these and other important systematic characters, such as karyotypes, are not available for the group at this time. our tendency has been to take a conservative view of species; thus it is doubtful that any subsequent worker will recognize fewer species in the genus. our observations on these frogs in amazonian ecuador are presented in a final section of this paper. acknowledgments for the loan of specimens or for the provision of working space in their respective institutions, we are indebted to james e. böhlke, werner c. a. bokermann, f. w. braestrup, nelly carrillo de espinoza, osvaldo r. da cunha, josef eiselt, m. j. fouquette, jr., alice g. c. grandison, jean guibé, birgitta hansson, walter hellmich, m. j. hoogmoed, robert f. inger, konrad klemmer, jean lescure, alan e. leviton, clarence j. mccoy, robert h. mount, charles w. myers, umberto parenti, günther peters, james a. peters, william f. pyburn, juan a. rivero, dorothy m. smith, paulo e. vanzolini, greta vestergren, david b. wake, charles f. walker, ernest e. williams, and richard g. zweifel. study of specimens in european museums was made possible by a grant (no. 5063) from the penrose fund of the american philosophical society. field work in ecuador was partially supported by grants from the watkins fund of the museum of natural history, university of kansas. at our base camp at santa cecilia, ecuador, we enjoyed the hospitality of ing. ildefonso muñoz b. transportation in ecuador was generously provided by the texaco petroleum company. during the course of our field work stephen r. edwards and thomas h. fritts contributed directly to our study of _osteocephalus_. michael j. tyler of the south australian museum provided information on the vocal sac structure. we extend our sincere thanks to all of these persons for their contributions to our endeavors. materials and methods we have examined 893 preserved frogs, including the type specimens of all included nominal taxa, 8 skeletons, 1 lot of eggs, and 3 lots of tadpoles that we refer to the genus _osteocephalus_; in addition skulls were removed from five preserved specimens, and radiographs were made of 12 other preserved specimens. we have been fortunate in seeing living individuals of all species, except _o. pearsoni_, but we have colored photographs of a living specimen of that species. figures 1 and 2 were drawn from projected colored transparencies of living frogs. terminology follows that of duellman (1970b). on the distribution maps solid symbols indicate localities from which we have examined specimens; open symbols represent additional locality records based on the literature. throughout the text specimens are listed by their catalogue numbers preceded by the appropriate museum abbreviation, as follows: amnh american museum of natural history ansp academy of natural sciences of philadelphia asu arizona state university aum auburn university museum bmnh british museum (natural history) cas california academy of sciences cas-su stanford university collection (in california academy of sciences) cm carnegie museum fmnh field museum of natural history ku university of kansas museum of natural history mcz museum of comparative zoology, harvard university mizs museo ed istituto di zoologi sistematico, università di torino mjp museo javier prado, lima mnhn muséum national d'histoire naturelle, paris mpeg museu paraense emiliano goeldi, belém mvz museum of vertebrate zoology, university of california, berkeley mzusp museu de zoología, universidade da são paulo nhmg naturhistoriska museet göteborg nhmw naturhistorisches museum, wien nhrm naturhistoriska riksmuseet, stockholm rmnh rijksmuseum van natuurlijke histoire, leiden smf senckenbergische museum, frankfurt uimnh university of illinois, museum of natural history ummz university of michigan museum of zoology up université de paris upr-m university of puerto rico, mayagüez uta university of texas, arlington usnm united states national museum uzm universitets zoologiske museum, copenhagen wcab werner c. a. bokermann, são paulo, brasil zmb zoologisches museum berlin zsm zoologisches sammlung münchen historical resumé because of the taxonomic confusion that has surrounded the generic name _osteocephalus_ and two of the species (and their synonyms), we present a brief resumé of the taxonomic history of the group. among the amphibians sent to the muséum national d'histoire naturelle in paris by a monsieur leprieur in french guiana was a single female specimen of a moderately large hylid having a well-ossified skull and smooth dorsal skin. this specimen escaped from the covetous eyes of johann tschudi, who prematurely named several species on the basis of specimens in paris, and survived without an epithet until duméril and bibron (1841) proposed for it the name _hyla leprieurii_. the description of the species is fairly detailed, but the specimen was not illustrated. this is the earliest trivial name now associated with _osteocephalus_. fitzinger (1843) in his generic synopsis of amphibians and reptiles proposed the generic name _osteocephalus_ but did not associate a specific name with the genus. consequently, _osteocephalus_ fitzinger, 1843, is a _nomen nudum_. franz steindachner followed leopoldo fitzinger at the naturhistorisches museum in vienna, where he had access to fitzinger's notes and, of course, the important collections housed in that museum. steindachner (1862) named two species of _osteocephalus_ on the basis of brasilian specimens collected by johann natterer. both species were named in the same publication; _o. taurinus_ appeared on page 77, and _o. favolineatus_, on p. 80. this is the earliest association of the generic name _osteocephalus_ with a specific name and a description, both of which satisfy the code of zoological nomenclature for generic availability. therefore, steindachner is the authority for the generic name _osteocephalus_, which has _o. taurinus_ as the type species by original designation. it is not possible to determine whether or not steindachner's usage of _osteocephalus_ was the same as that intended by fitzinger 19 years earlier. steindachner (1862) gave reasonably good descriptions of his two new species and provided excellent illustrations of the two specimens, both large females. apparently impressed by the similarities between _trachycephalus nigromaculatus_ tschudi, 1838, and _osteocephalus taurinus_, steindachner (1867) used the combination _trachycephalus (osteocephalus) taurinus_. this ambiguous usage for the 1860's precludes our determining if steindachner was in effect synonymizing _osteocephalus_ with _trachycephalus_ or whether he was placing _osteocephalus_ in a subgeneric status. steindachner (1867) did not mention _o. flavolineatus_; perhaps by that time he had concluded that _flavolineatus_ was only a color morph of _taurinus_. cope (1867) placed _hyla leprieurii_ in the genus _hypsiboas_ wagler, 1830. cope (1874) named _osteocephalus planiceps_ from nauta, perú. the single specimen was among the collections made by the orton expedition to the upper amazon basin and was deposited in the academy of natural sciences in philadelphia. boulenger (1882) placed both _osteocephalus_ and _trachycephalus_ the synonymy of _hyla_; he recognized _hyla taurina_ (with _o. flavolineatus_ as a synonym), _h. leprieurii_, and _h. planiceps_. in the same publication boulenger named _hyla buckleyi_ on the basis of 10 specimens in the british museum from ecuador; in the description he stated that _buckleyi_ was like _leprieurii_ and _taurinus_ in having paired lateral vocal sacs. boulenger held a lasting influence on taxonomic herpetology, and his generic synonymy of _osteocephalus_ was unchallenged until only a decade ago. goin (1961) presented a generic synopsis of the genera of hylid frogs, in which he recognized _osteocephalus_ and stated: "there are perhaps eight or ten species of this genus in south america. certainly _taurinus_, _britti_, _leprieuri_, _buckleyi_ and _pearsoni_ belong here. _o. planiceps_ is surely a synonym of _leprieuri_ and i believe that _garbei_ is as well. the status of such forms as _macrotis_, _riopastazae_, and _depressa_ has not yet been settled." goin defined _osteocephalus_ as follows: "males with paired vocal pouches, one at each angle of the jaw; derm of head not co-ossified with skull but roof of skull exostosed." trueb (1970a) elaborated on goin's definition and assuredly included only _o. taurinus_ and _o. leprieurii_ in the genus. goin's inclusion of _buckleyi_, _britti_, and _pearsoni_ in _osteocephalus_ was the first association of any of these names with that genus. duellman (1970a) demonstrated that _garbeana garbei_ miranda-ribeiro, 1926, was a member of the _hyla rubra_ group. _hyla macrotis_ andersson, 1945, is a _phrynohyas_. trueb and duellman (1970) determined that _hyla verrucigera_ werner, 1901, is the earliest name for an _osteocephalus_ displaying striking sexual dimorphism in coloration and texture of the dorsal skin; _hyla riopastazae_ andersson, 1945 (female holotype), and _hyla orcesi_ funkhouser, 1956 (male holotype), were placed in the synonymy of _osteocephalus verrucigerus_. _hyla pearsoni_ gaige, 1929, is a small species of _osteocephalus_. our findings substantiate goin's suggestions relative to two other taxa. _hyla leprieurii britti_ melin, 1941, from the rio uaupés, brasil, and _hyla depressa_ andersson, 1945, from the río pastaza watershed, ecuador, are members of the genus _osteocephalus_, but both are synonyms of earlier names--_leprieurii_ and _taurinus_, respectively. another name proposed by melin (1941), _hyla (trachycephalus) vilarsi_ from taracuá, brasil, also is placed in the synonymy of _o. taurinus_. cochran and goin (1970) were unaware of the identities of _hyla verrucigera_ and _riopastazae_; they used the later name _osteocephalus orcesi_ for colombian frogs that are correctly referred to _o. verrucigerus_. although goin (1961) placed _hyla buckleyi_ and _h. pearsoni_ in _osteocephalus_, cochran and goin (1970) recognized a "_buckleyi_ group" in _hyla_ that included these two species plus a new species, _hyla cabrerai_ from amazonian colombia and brasil (total of three specimens). also, these authors named _hyla carri_ from a single colombian specimen. study of the types of _hyla cabrerai_, _h. carri_, and _h. festae_ peracca, 1904, from ecuador, reveal that all of these names are synonyms of _osteocephalus buckleyi_. much of the taxonomic confusion and multiplicity of trivial names is due to the great amount of color variation in _taurinus_ and to the sexual dimorphism in the texture of the dorsal skin in all of the species. the details of variation in these and other characters and our justifications for the synonymies are given in the accounts of the species. all of the trivial names that apply to species herein recognized as members of the genus _osteocephalus_ are listed in table 1. =osteocephalus= steindachner, 1862 _osteocephalus_ steindachner, 1862:77 [type species.--_osteocephalus taurinus_ steindachner, 1862, by original designation]. not _osteocephalus_ fitzinger, 1843:50 (_nomen nudum_). _diagnostic definition._--1) skull broader than long; 2) dermal roofing bones of skull well ossified, exostosed, and/or co-ossified in some species; 3) prenasal and internasal bones absent; 4) parasphenoid alae posterolaterally oriented; 5) dentigerous processes of prevomers angular (/--\); 6) vocal sacs paired, posterior, and when inflated protruding posteroventral or posterolateral to angles of jaws; 7) submentalis muscle moderate in size and araphic; 8) intermandibularis muscle undifferentiated and bearing an elongate median aponeurosis; 9) parotoid glands absent or poorly developed, skin not producing viscous secretion characteristic of _phrynohyas_; 10) skin on dorsum tuberculate in males, smooth in females; 11) tympanum large, 60 percent or more of diameter of eye; 12) fingers about one-third, toes more than three-fourths webbed; 13) discs large, round; 14) nuptial excrescences present in breeding males; 15) inner metatarsal tubercle not modified for digging; 16) outer metatarsal tubercle absent; 17) tarsal fold weak or absent; 18) pupil horizontal; 19) palpebrum clear; 20) known tadpoles having two upper and five lower rows of teeth. table 1.--alphabetical synonymy of the species of _osteocephalus_. trivial name, original generic name, author, date current name --------------------------------------------------------------------- _britti (hyla leprieurii)_ melin, 1941 _o. leprieurii_ _buckleyi (hyla)_ boulenger, 1882 _o. buckleyi_ _cabrerai (hyla)_ cochran and goin, 1970 _o. buckleyi_ _carri (hyla)_ cochran and goin, 1970 _o. buckleyi_ _depressa (hyla)_ andersson, 1945 _o. taurinus_ _festae (hyla)_ peracca, 1904 _o. buckleyi_ _flavolineatus (osteocephalus)_ steindachner, 1862 _o. taurinus_ _leprieurii (hyla)_ duméril and bibron, 1841 _o. leprieurii_ _orcesi (hyla)_ funkhouser, 1956 _o. verrucigerus_ _pearsoni (hyla)_ gaige, 1929 _o. pearsoni_ _planiceps (osteocephalus)_ cope, 1874 _o. taurinus_ _riopastazae (hyla)_ andersson, 1945 _o. verrucigerus_ _taurinus (osteocephalus)_ steindachner, 1862 _o. taurinus_ _verrucigera (hyla)_ werner, 1901 _o. verrucigerus_ _vilarsi (hyla)_ melin, 1941 _o. taurinus_ _content._--as defined here, the genus contains five known species: _o. buckleyi_ (boulenger), _o. leprieurii_ (duméril and bibron), _o. pearsoni_ (gaige), _o. taurinus_ steindachner, and _o. verrucigerus_ (werner). _distribution._--the guianas and amazon basin; also in the upper orinoco and magdalena drainages. most localities are at elevations below 500 m, but the genus ascends the amazonian slopes of the andes to elevations of about 1800 m. analysis of characters _size and proportions._--frogs of the genus _osteocephalus_ are moderate to large hylids. the largest species is _taurinus_, attaining a snout-vent length of 103.1 mm; the smallest is _pearsoni_, which attains a length of 54.7 mm. considerable intraspecific geographic variation occurs in adult size, especially in _taurinus_. females of all species attain a noticeably larger size than males, but no significant differences are apparent in proportions (table 2). table 2.--comparison of size and proportions in the species of _osteocephalus_. (means are given in parentheses below observed ranges) ======================================================================================================== species n snout-vent tibia length/ foot length/ head length/ head width/ tympanum/ length s-v l s-v l s-v l s-v l eye ------------------------------------------------------------------------------------------------------- _o. buckleyi_ [m] 30 37.9-48.1 0.478-0.580 0.375-0.444 0.319-0.357 0.329-0.368 0.608-0.820 (43.3) (0.520) (0.408) (0.343) (0.351) (0.711) [f] 31 48.6-75.1 0.476-0.599 0.363-0.469 0.310-0.358 0.318-0.367 0.574-0.905 (61.7) (0.553) (0.428) (0.333) (0.348) (0.734) _o. leprieurii_ [m] 21 41.2-48.4 0.514-0.571 0.383-0.430 0.308-0.357 0.326-0.368 0.652-0.884 (44.7) (0.538) (0.408) (0.335) (0.348) (0.777) [f] 21 46.6-61.5 0.516-0.592 0.382-0.453 0.314-0.343 0.328-0.363 0.698-0.909 (57.1) (0.539) (0.404) (0.329) (0.349) (0.785) _o. pearsoni_ [m] 2 45.3-46.2 0.481-0.504 0.404-0.437 0.322-0.335 0.327-0.342 0.660-0.673 (45.8) (0.493) (0.421) (0.329) (0.335) (0.666) [f] 1 54.7 0.521 0.405 0.318 0.346 0.862 _o. taurinus_ [m] 59 40.3-84.6 0.512-0.576 0.387-0.445 0.296-0.345 0.301-0.355 0.638-0.896 (66.3) (0.541) (0.416) (0.318) (0.324) (0.752) [f] 45 45.1-103.1 0.520-0.577 0.391-0.448 0.306-0.334 0.308-0.347 0.640-0.817 (75.8) (0.542) (0.420) (0.321) (0.327) (0.758) _o. verrucigerus_ [m] 11 50.4-54.3 0.494-0.552 0.409-0.442 0.322-0.346 0.328-0.344 0.623-0.804 (53.0) (0.519) (0.427) (0.333) (0.337) (0.730) [f] 3 63.1-65.8 0.532-0.561 0.435-0.463 0.345-0.347 0.348-0.379 0.692-0.808 (64.5) (0.545) (0.448) (0.346) (0.358) (0.731) -------------------------------------------------------------------------------------------------------_coloration._--all _osteocephalus_ are predominantly brown frogs usually with some darker dorsal markings (figs. 1 and 2). _osteocephalus verrucigerus_ has a nearly uniform dark brown dorsum and no distinct transverse bars on the limbs, whereas all of the other species have distinct bars on the limbs. the dorsal markings on the body consist of irregular blotches in _buckleyi_, _pearsoni_, and _taurinus_ but are narrow transverse marks in _leprieurii_. a narrow middorsal cream or yellow stripe is present in some individuals of _buckleyi_ and _taurinus_ but absent in all individuals of the other species. the flanks are uniform pale tan in _leprieurii_ and uniform reddish brown in _verrucigerus_; in the other species the flanks are cream to brown with dark brown or black spots (also dark diagonal marks in some _buckleyi_). a creamy white anal stripe is present in some specimens of _leprieurii_ but absent in all individuals of other species. the postocular region, encompassing the tympanum, is dark brown in most specimens. in adults of _pearsoni_ and _taurinus_ the upper lips are dark brown. a pale cream or tan suborbital spot is present in _pearsoni_ and in some _taurinus_; in some specimens of _taurinus_ the suborbital spot is expanded posteriorly forming a labial stripe on the posterior part of the lip. the labial markings of _verrucigerus_ are similar to the latter pattern, except that in females a distinct, light labial stripe extends the length of the lip. _osteocephalus leprieurii_ has a distinct, broad, pale labial stripe. the lips are barred cream and dark brown in _buckleyi_. the venter is uniform creamy white or pale tan in _leprieurii_, uniform white in some _buckleyi_ (most males), and uniform tan in some _taurinus_. the other species and some individuals of _taurinus_ and _buckleyi_ (most females) have dark ventral markings. these markings are most distinctive in _verrucigerus_, in which the venter is white with bold black mottling and spots (fig. 3c). those individuals of _taurinus_ having ventral markings usually have indistinct, diffuse brown spots on the throat and chest (fig. 3b). _osteocephalus pearsoni_ is characterized by a fine brown reticulation on the venter and on the anterior and posterior surfaces of the thighs in adults (fig. 3a). individuals of _buckleyi_ that have ventral markings vary between the patterns illustrated for _pearsoni_ and _taurinus_ (figs. 3b and c). ontogenetic change in coloration is slight or non-existent in _buckleyi_, _pearsoni_, and _taurinus_, except that juveniles lack ventral markings. a dark blotch on the back and distinct transverse bars on the limbs are evident in juveniles of _verrucigerus_; these markings are obscured in the adults. juveniles of _leprieurii_ are olive-brown with yellow dorsolateral stripes; the transverse dark marks, characteristic of the adults, appear before the stripes are lost. [illustration: fig. 1. species of _osteocephalus_: top. _o. pearsoni_, ku 136312, [m]; middle. _o. buckleyi_, ku 123172, [m]; bottom. _o. verrucigerus_, ku 123177, [m]. ×1.5.] [illustration: fig. 2. species of _osteocephalus_: top. _o. leprieurii_, ku 126611, [f]; bottom. _o. taurinus_, ku 126648, [m]. ×1.] [illustration: fig. 3. diagrammatic views of ventral color patterns in _osteocephalus_: a. _o. pearsoni_, ummz 57533, [m]; b. _o. taurinus_, usnm 166037, [m]; c. _o. verrucigerus_, ku 123185, [f].] _skin._--the dorsal skin of all males of _osteocephalus_ is tuberculate to varying degrees, whereas the dorsal skin of females is smooth, or nearly so (fig. 4). _osteocephalus verrucigerus_ differs from other members of the genus by the presence of numerous, large tubercles bearing keratinized tips. the tubercles of _leprieurii_ are numerous and spinous but much smaller than those of _verrucigerus_; those of _taurinus_ are spinous but less numerous than in _leprieurii_. _osteocephalus buckleyi_ has a mixture of large and small, non-spinous tubercles, and _pearsoni_ has only a few, small, scattered, non-spinous tubercles. fleshy tubercles occur on the eyelids and supratympanic fold in females of _buckleyi_; a few small tubercles are present on the back of females of _pearsoni_, whereas the dorsal skin in females of the other species is smooth. the skin on the flanks of both sexes of _buckleyi_ is weakly areolate; in the other species the flanks are smooth. the skin on the top of the head in _taurinus_ is rugose as a consequence of co-ossification. in all species the anal opening is directed posteriorly at the upper level of the thighs. _hands and feet._--the feet of _osteocephalus_ are fully webbed or nearly so. webbing between fingers one and two is basal in all species. webbing between fingers two, three, and four is most extensive in _taurinus_, in which the three fingers are about one-half webbed (fig. 5). _osteocephalus buckleyi_, _pearsoni_, and _verrucigerus_ have reduced webbing between fingers two and three, and _leprieurii_ has reduced webbing between fingers two, three, and four. all members of the genus have well-developed subconical subarticular tubercles on the fingers and toes; there is a tendency for the distal tubercle on the fourth finger to be weakly bifid. palmar and plantar supernumerary tubercles are well developed in _taurinus_, moderately developed in _buckleyi_, _leprieurii_, and _pearsoni_, and barely evident in _verrucigerus_. all of the species have a noticeable fold on the wrist and enlarged prepollices, bearing horny nuptial excrescences in breeding males. the prepollex is least enlarged in _buckleyi_. outer metatarsal tubercles are absent. the inner metatarsal tubercle is moderately well developed and ovoid in _leprieurii_ and _pearsoni_; it is elliptical and flat in the other species. tarsal folds are absent in all species except _verrucigerus_, in which the folds are barely evident. [illustration: fig. 4. segments of dorsal skin of males of _osteocephalus_ showing size and arrangement of tubercles: a. _o. verrucigerus_, ku 123183; b. _o. taurinus_, usnm 166033; c. _o. leprieurii_, ku 126616; d. _o. buckleyi_, usnm 165999. each square = 1 sq. cm.] _cranium._--as a genus, the cranial structure is remarkably uniform and quite generalized when viewed in the context of the family hylidae. the skulls are broad and relatively flat, each being only slightly more broad than long and about one-third as high as long. in dorsal aspect the snouts are broadly rounded; the snout of _buckleyi_ is somewhat less rounded and appears to be slightly longer than the snouts of other species. this subtle difference relates to the relative narrowness of the premaxillaries in _buckleyi_. [illustration: fig. 5. palmar views of hands of males of _osteocephalus_: a. _o. buckleyi_, ku 109506; b. _o. leprieurii_, ku 126627; c. _o. pearsoni_, mcz 15565; d. _o. taurinus_, ku 126653; e. _o. verrucigerus_, ku 123177. ×4.] [illustration: fig. 6. skulls of two species of _osteocephalus_: a and b. _o. leprieurii_, ku 125961; c and d. _o. pearsoni_, ummz 67465. ×3.] the genus is characterized by well-developed dermal roofing bones and an unusually large exposure of the sphenethmoid dorsally (fig. 6). the conformation of the sphenethmoid exposed dorsally is determined by the marginal positions of the adjacent, overlapping elements--the nasals and frontoparietals. medially the nasals overlap the lateral margins of the sphenethmoid. anteromedially, the nasals converge in _leprieurii_ and _taurinus_, are narrowly separated in _buckleyi_ and _pearsoni_, or are more widely separated in _verrucigerus_. in all species the nasals terminate at the anterodorsal corner of the orbit. the frontoparietals of _buckleyi_, _leprieurii_, and _taurinus_ have an anterolateral extension, which marginally overlaps the dorsolateral edge of the sphenethmoid and articulates with the posterodorsal corner of the nasal in _buckleyi_ and _taurinus_; the bones are narrowly separated in _leprieurii_. the frontoparietals of _pearsoni_ and _verrucigerus_ have more extensive median ossification and less extensive anterolateral ossification. thus, in those species the posteromedian portion of the sphenethmoid is obscured, and the lateral margins are partly exposed. the frontoparietal fontanelle is completely covered in all species, except _buckleyi_ and _leprieurii_, in which an irregular, median separation of the frontoparietals exposes a small portion of the fontanelle. the posterolateral margins of the frontoparietals lie medial to the epiotic eminences. dermal ornamentation, involving the nasals, frontoparietals, and sphenethmoid, occurs in _taurinus_ and, to a limited extent, in _pearsoni_. in the latter species marginal portions of the frontoparietals, the dorsal surfaces of the nasals, and the posteromedial part of the exposed sphenethmoid are slightly exostosed, resulting in an open, reticulate pattern of dermal sculpturing of exceedingly low relief (fig. 6c). _osteocephalus taurinus_ is characterized by casquing, co-ossification, and a rather intricate pattern of dermal sculpturing, which was described in detail and illustrated by trueb (1970a). the squamosals of all species are moderately large structures having otic plates that overlie the lateral portion of the cristae paroticae. the posterior rami are short; the zygomatic rami of all species, except _taurinus_, extend slightly more than one-half of the distance to the maxillary. in _taurinus_ the zygomatic ramus extends nearly to, or articulates with, the maxillary. the maxillary arches are complete and relatively robust. the alary processes of the premaxillaries are vertically oriented in _leprieurii_, _pearsoni_, and _taurinus_ and very slightly inclined posteriorly in _buckleyi_ and _verrucigerus_. the maxillaries are uniformly characterized by the absence of postorbital processes and by the presence of preorbital processes that articulate with the maxillary processes of the nasals. the partes facialae of the maxillaries are moderately developed in all species, except _taurinus_, in which the pars fascialis tends to articulate with the lateral margin of the nasal in well-ossified individuals. the partes palatinae are poorly developed in all species, except _buckleyi_, in which the pars palatina of the premaxillary is moderately robust. the pterygoids are uniformly tri-radiate structures. the anterior rami terminate at about the mid-level of the orbit, and the medial rami articulate firmly with the anterolateral corner of the otic capsule. the palatines are well-developed elements bearing ventral ridges; the ridges are somewhat irregular in _buckleyi_, _taurinus_, and _verrucigerus_ but smooth in _leprieurii_ and _pearsoni_. the parasphenoids are large elements characterized by acuminate cultriform processes and posterolaterally inclined alary processes. the basal areas of the cultriform processes bear small odontoid protuberances in _buckleyi_, _taurinus_, and _verrucigerus_, whereas they are smooth in _leprieurii_ and _pearsoni_. the prevomers are remarkably uniform, moderately well-developed structures. in each species the anterior ramus lies adjacent to the premaxillary, and the lateral wings form the anterior, medial, and posteromedial margins of the internal nares. the dentigerous processes are characteristically large and angular and bear numerous teeth. the sphenethmoid and otoccipitals are well ossified; a dermal sphenethmoid is present only in _taurinus_. [illustration: fig. 7. dorsal views of vertebral columns of two species of _osteocephalus_: a. _o. leprieurii_, ku 125962, [f]; b. _o. buckleyi_, usnm 165997, [f]. ×2.] _vertebral column._--the cervical cotyles are uniformly widely displaced. the neural arches are low and non-imbricate. the transverse processes of the third presacral vertebrae are approximately equal in width to the sacral diapophyses in all species, except _buckleyi_, in which the processes of the third presacral are slightly narrower than the diapophyses. _osteocephalus buckleyi_ is further distinguished by the presence of narrow transverse processes on presacrals five through eight (fig. 7b); males have narrower processes than do females. the processes are moderately wide but subequal in width in _pearsoni_, _taurinus_, and _verrucigerus_, whereas they are nearly equivalent in width to one another and to the sacral diapophyses in _leprieurii_ (fig. 7a). the sacral diapophyses of all species are moderately dilated and posterolaterally inclined. the coccyx bears a distinct dorsal ridge and has a bicondylar articulation with the sacrum. _pectoral girdle._--the pectoral girdles are fully arciferal and bear small, cartilaginous omosterna and moderately large cartilaginous sterna. the coracoids are robust, and the clavicles are strongly arched. procoracoid cartilage seems to be absent. the scapulae are large, longer than the clavicles, and bicapitate proximally. the suprascapulae are moderately large and well ossified in _leprieurii_ and _taurinus_. the suprascapula of _verrucigerus_ is poorly ossified, and that of _buckleyi_ is not ossified. _pelvic girdle._--the ilia of _buckleyi_, _taurinus_, and _verrucigerus_ lack any indication of a crest on the shaft, whereas _leprieurii_ has a low crest. the dorsal acetabular expansion of the ilia is moderately low in _taurinus_ and _verrucigerus_, but distinctly lower in _buckleyi_ and _leprieurii_. the ilia of all species bear low dorsal protuberances. the ischia of _leprieurii_, _taurinus_, and _verrucigerus_ are moderately expanded; that of _buckleyi_ is somewhat less expanded dorsally. the pubis of _leprieurii_, _taurinus_, and _verrucigerus_ are calcified, whereas that of _buckleyi_ remains cartilaginous. _throat musculature and vocal sac structure._--tyler (1971) described the throat myology of _osteocephalus_. the genus is characterized by a moderate-sized araphic submentalis muscle and an undifferentiated intermandibularis having an elongate median aponeurosis. the intermandibularis and submentalis are completely independent in _buckleyi_, whereas in the other species there is a small attachment between these muscles. according to tyler (pers. com.), _osteocephalus_ has three distinctive types of vocal sac structure which result from differences in the development of the interhyoideus muscle and the overlying skin. in _leprieurii_ and _verrucigerus_ the supramandibular portions of the interhyoideus form a simple tubular, posterolateral extension; there is no modification of the associated skin. _osteocephalus buckleyi_ and _pearsoni_ have more extensive development of the supramandibular portions of the interhyoideus; furthermore, the associated skin forms a broad, loose fold extending from the ventromedial surface of the throat dorsolaterally to the base of the supratympanic fold. like _buckleyi_ and _pearsoni_, the supramandibular portion of the interhyoideus is much expanded in _taurinus_. the vocal sac structure of _taurinus_ differs from that of other members of the genus in that the skin of _taurinus_ forms an everted pouch, which dangles loosely beneath the supratympanic fold. key to the species of _osteocephalus_ 1. inner edge of third finger webbed to mid-length of antepenultimate phalange; dorsum brown with dark brown spots or median blotch; skull in adults casqued and co-ossified with prominent supraorbital flanges _o. taurinus_ inner edge of third finger webbed to base of antepenultimate phalange; dorsum plain or marked with dark blotches or transverse bars; skull in adults smooth or slightly exostosed, lacking supraorbital flanges 2 2. skin on flanks areolate; dorsum in males bearing a mixture of large and small non-spinous tubercles; lips distinctly barred _o. buckleyi_ skin on flanks smooth; dorsum in males bearing tubercles of uniform size; lips not barred 3 3. dorsal pattern consisting of narrow transverse dark bars; dorsum in males bearing numerous small spinous tubercles _o. leprieurii_ dorsal pattern not consisting of transverse bars; dorsal tubercles large or few in number 4 4. dorsum uniformly dark brown; venter heavily mottled with black, especially in females; dorsum in males bearing large, keratinized tubercles _o. verrucigerus_ dorsum tan with irregular dark brown blotches; venter cream with fine brown reticulations; dorsum in males bearing few, small non-spinous tubercles _o. pearsoni_ accounts of species =osteocephalus buckleyi= (boulenger) _hyla buckleyi_ boulenger, 1882:362 [syntypes.--bmnh 1947.2.13.36-39 from sarayacu, provincia pastaza, ecuador; bmnh 1947.2.13.40-41, 1947.2.13. 43-45 from canelos, provincia pastaza, ecuador; bmnh 1947.2.13.46 from "paitanga" (= pallatanga), provincia chimborazo, ecuador (in error); mr. buckley collector; bmnh 1947.2.13.44 here designated as lectotype]. _hyla festae_ peracca, 1904:39 [holotype.--mizs 2950 from "valle de santiago" (= lower río zamora), provincia morona-santiago, ecuador; enrico festa collector]. new synonymy. _osteocephalus buckleyi_--goin, 1961:13. _hyla carri_ cochran and goin, 1970:211 [holotype.--fmnh 69702 from acevedo, río suaza, departamento huila, colombia; philip collector]. new synonymy. _hyla cabrerai_ cochran and goin, 1970:215 [holotype.--usnm 152759 from caño guacayá, tributary of lower río apoporis, comisaria amazonas, colombia; isadore cabrera collector]. new synonymy. _justification of synonymy._--boulenger (1882:362) listed 11 specimens in his description of _hyla buckleyi_. we have examined all of these and conclude that one (bmnh 1947.2.13.42) is _o. leprieurii_. cochran and goin (1970:213) restricted the type locality to canelos, provincia pastaza, ecuador; we here select bmnh 1947.2.13.44 from that locality as the lectotype. this specimen is a male having a snout-vent length of 37.9 mm; the diameter of the tympanum is 3.5 mm, 81.4 percent of the diameter of the eye. the type series, exclusive of bmnh 1947.2.13.42 (=_o. leprieurii_) consists of six males having snout-vent lengths of 37.9-44.6 (mean 40.4) mm, and four females having snout-vent lengths of 50.0-53.9 (mean 51.5) mm. the dorsum in the males bears a mixture of large and small tubercles, whereas the dorsum in females is nearly smooth. the skin on the flanks, especially the axilla, is areolate. the coloration consists of a creamy tan ground color with irregular reddish brown markings on the back and broad transverse bars on the limbs. the dorsal markings are narrowly bordered by creamy white; those on the back consist of an interorbital bar and a pair of longitudinal marks beginning in the scapular region and usually diverging posteriorly in the sacral region or converging into a broad median blotch. one specimen has a middorsal creamy white stripe from the tip of the snout to the vent. in all of the types large dark brown spots are present on the flanks and posterior surfaces of the thighs. the ventral surfaces are pale creamy tan with or without diffuse brown spots on the throat and chest. the holotype of _hyla festae_ is a female having a snout-vent length of 75.0 mm; the diameter of the tympanum is 3.9 mm, 57.4 percent of the diameter of the eye. the skin is smooth on the dorsum and areolate on the anterior part of the flanks. the dorsum is pale brown with a large median longitudinal dark brown blotch on the back and broad transverse bars, narrowly outlined by cream, on the limbs. dark brown spots are present on the flanks; the posterior surfaces of the thighs are dark brown. the throat and belly are grayish white with irregular dark brown spots. the holotype of _hyla carri_ is a female having a snout-vent length of 66.1 mm; the diameter of the tympanum is 4.7 mm, 81.0 percent of the diameter of the eye. the skin on the dorsum is smooth with scattered small tubercles and areolate on the anterior part of the flanks. the dorsum is tan with irregular dark brown blotches on the back and transverse bars on the limbs; all dorsal markings are narrowly outlined by creamy white. dark brown spots are present on the flanks; the venter and posterior surfaces of the thighs are tan without dark spots. the holotype of _hyla cabrerai_ is a female having a snout-vent length of 52.7 mm; the diameter of the tympanum is 4.0 mm, 76.9 percent of the diameter of the eye. the skin on the dorsum is weakly tuberculate and that on the anterior part of the flanks is areolate. the dorsum is creamy tan with dark brown markings (interorbital bar, reticulations on occiput, three longitudinal streaks on back, and broad transverse bars on limbs). irregular dark brown spots are present on the flanks. the venter is pinkish tan with small reddish brown spots on the throat and darker brown spots on the chest and belly. in their description of _hyla cabrerai_, cochran and goin (1970:217) stated: "this species, together with _buckleyi_ and _pearsoni_ certainly make a closely knit group.... both _buckleyi_ and _cabrerai_ have long hind legs, with the extended heel reaching to the tip of the snout, while in _pearsoni_ the extended heel reaches only to the eye. _h. buckleyi_ has the belly dusky, while it is heavily spotted in _cabrerai_ and is reticulated in _pearsoni_. _h. cabrerai_ seems to have the heaviest hands with the most webbing between the fingers; the other two species have the webbing reduced between the fingers." the description of _hyla cabrerai_ was based on three specimens. we have examined the holotype and one paratype (wcab 13284 from territorio do amapá, brasil). another paratype in the private collection of c. j. goin from caño tuí, between mitú and raudal de yurupari, comisaria de vaupés, colombia, was not examined. cochran and goin (1970:211) based their description of _hyla carri_ on one gravid female and stated: "a large _hyla_ with the vomerine teeth in two ^^ shaped patches between the somewhat squarish choanae; reduced webs between the fingers; and a pattern of dorsal dark blotches bordered by light margins. the species is not similar to any other species known in colombia. it is perhaps most closely related to _hyla claresignata_ of brazil, from which it can be differentiated by its more heavily spotted dorsum, larger tympanum, and lack of dark anal spots." except for the inclusion of the name in checklists, _hyla festae_ has not been mentioned in the literature since the original description. the wholesale synonymization of names, which, on the bases of their published diagnoses, seem to apply to distinctly different species, is possible with the application of uniform criteria to the types and series of other specimens. in measurements and proportions the type specimens of the nominal taxa all fall within the range of variation exhibited by a series of 18 males and 15 females from provincia pastaza, ecuador, except the ratio of the diameter of the tympanum to that of the eye in the female holotype of _hyla festae_. in that specimen the ratio is 0.574, whereas the ratio in the 15 females from provincia pastaza is 0.587-0.905 (mean 0.736). ventral coloration is the most variable character among the types. the venter in the type of _hyla festae_ is boldly spotted; it is distinctly spotted in _cabrerai_, uniform tan in _carri_, and tan, flecked, or spotted in the type series of _buckleyi_. the ventral coloration in series of specimens from amazonian ecuador encompasses that observed in all of the types, except that of _festae_, which has more ventral spotting than any other individual. the webbing on the hand usually excludes the penultimate phalanges of the fingers, but in some specimens from amazonian ecuador the webbing encompasses the proximal parts of the penultimate phalanges of the fingers. in a few of these specimens, the holotype of _festae_, and one paratype of _cabrerai_ the webbing extends to the middle of the penultimate phalanges of the third and fourth fingers. in the holotype of _cabrerai_ the webbing extends to the middle of the penultimate phalanges of the third and fourth fingers and to the base of the disc of the second finger. the types of the nominal taxa and series of specimens from guyana and amazonian ecuador display noticeable variation in dorsal coloration. the variety of dorsal patterns of all of the types is included in the variation displayed by the other specimens. all specimens have some amount of dark spotting on the flanks; all have vertically barred lips, on which a pale subocular spot usually is evident. probably the most unifying physical characteristic of all of the specimens is the nature of the skin on the anterior part of the flank. the skin is elevated amidst an irregular network of depressions. this areolate dermal condition is present in all specimens and does not occur in other species of _osteocephalus_. the degree of tubercularity of the skin on the dorsum is variable and sexually dimorphic. all males are tubercular, whereas small females are smooth or have only a few scattered tubercles. large females usually have pronounced tubercles on the eyelids and supratympanic fold. in their description of _hyla carri_, cochran and goin (1970:211) misrepresented the nature of the dentigerous processes of the prevomers, which are angular, not ^-shaped. their suggestion that the colombian _hyla carri_ is related to _hyla claresignata_ in southeastern brasil is unfounded. the latter species is smaller (40 mm), has a yellow dorsum and venter, dark brown spots dorsolaterally, oblique dentigerous processes of the prevomers, small tympanum, and smooth skin dorsally. the ventral coloration of the type of _hyla festae_ resembles that of _osteocephalus verrucigerus_, but the type differs from _verrucigerus_ by having areolate skin on the flanks and distinct dark markings on the dorsum. in _verrucigerus_ the skin on the flanks is smooth, and the dorsum is uniform dark brown, except for a tan snout in females. comparisons of the types of the nominal species with series of specimens from guyana, colombia, ecuador, and perú suggest strongly that the types are representative of one taxon, the oldest name for which is _hyla buckleyi_ boulenger, 1882. consequently, we place _hyla festae_ peracca, 1904, _hyla carri_ cochran and goin, 1970, and _hyla cabrerai_ cochran and goin, 1970, as junior synonyms of _hyla buckleyi_ boulenger, 1882. _diagnosis._--1) size moderate, sexual dimorphism extreme; maximum observed snout-vent length in males 48.1 mm, in females 75.1 mm; 2) skin on dorsum in males bearing a mixture of large and small non-spinous tubercles; 3) skin on flanks, especially anteriorly, areolate; 4) web usually extending only to base of antepenultimate phalange on inner edge of third finger; 5) dorsum pale tan or green with irregular, longitudinal, dark brown blotches, usually narrowly outlined with cream; 6) venter cream or tan, suffused with brown or marked with brown spots in some specimens; 7) lips marked with vertical brown and cream bars; 8) flanks creamy tan with irregular brown spots and/or diagonal marks; 9) dermal roofing bones of skull lacking exostosis; 10) dermal sphenethmoid absent; 11) nasals widely separated medially; 12) anteromedial margin of frontoparietal at mid-level of orbit; 13) frontoparietal fontanelle partially exposed; 14) palatine serrate; 15) parasphenoid bearing odontoids; 16) zygomatic ramus of squamosal extending approximately one-half of distance to maxillary arch; 17) transverse processes of third presacral vertebra narrower than sacral diapophyses; transverse processes of presacral vertebrae 3-8 subequal in width and narrower in males than in females; 18) intermandibularis and submentalis muscles independent; 19) supramandibular portion of interhyoideus extensively developed; associated skin forming broad loose fold. _osteocephalus buckleyi_ can be distinguished readily from all other species in the genus by the presence of areolate skin anteriorly on the flanks and by the rather boldly contrasting dorsal pattern. furthermore, females are distinctive in having tubercles on the eyelids and supratympanic folds. _distribution._--the periphery of the amazon basin, in the guianas and territorio do amapá in northeastern brasil; the upper amazon basin from southern colombia to east-central bolivia; one locality (acevedo) in upper río magdalena drainage in colombia (fig. 8). all localities are at elevations of less than 700 m. records for pallatanga and santiago in provincia chimborazo, ecuador (high on the pacific slopes of the andes), are considered to be erroneous. 78 specimens from 40 localities. [illustration: fig. 8. distribution of _osteocephalus buckleyi_ (circles) and _o. pearsoni_ (triangles).] _remarks._--in life the dorsum is green with dark markings. a male (ku 123171) from santa cecilia, ecuador, was: "dorsum green with dark brown blotches. anterior and posterior surfaces of thighs dull blue. venter brown, flecked with white. iris greenish bronze with brown horizontal triangles and ventromedian brown line." (w. e. duellman, field notes, 16 june 1968.) a female (ku 126646) from lago agrio, ecuador, was: "dorsum pale green with darker green blotches and creamy yellow middorsal stripe. lateral blotches bronze-tan. flanks tan with black blotches. anterior surfaces of thighs dark brown. dorsal and posterior surfaces of thighs and shanks tan with dark brown blotches. webbing brown. suborbital spot green. postorbital bar black. belly grayish brown in appearance--tips of granules white; intergranular spaces brown. iris golden bronze with black flecks peripherally and median, horizontal, reddish brown streak." (w. e. duellman, field notes, 12 may 1969.) no ontogenetic change in coloration has been noted. =osteocephalus leprieurii= (duméril and bibron) _hyla leprieurii_ duméril and bibron, 1841:553 [holotype.--mnhn 4629 from "cayenne"; mons. leprieur collector]. _hypsiboas leprieurii_--cope, 1867:200. _hyla leprieurii britti_ melin, 1941:42 [holotype.--nhmg 489 from the rio uaupés, north of the rio japu, territorio do amazonas, brasil; douglas melin collector]. new synonymy. _hyla leprieurii leprieurii_--melin, 1941:42. _osteocephalus britti_--goin, 1961:13. _osteocephalus leprieurii_--goin, 1961:13. _justification of synonymy._--the holotype of _hyla leprieurii_ is a female having a snout-vent length of 46.6 mm. the diameter of the tympanum is 3.7 mm, 69.8 percent of the diameter of the eye. the dorsal roofing bones are smooth, and the skin on the dorsum is smooth. the penultimate phalanges of the fingers are not included in the webbing. when we examined the specimen on 2 july 1969, it was slightly soft and somewhat faded to a peculiar grayish green color with faint darker transverse bars on the limbs. duméril and bibron (1841:554) described the coloration, as follows: "the loreal region is black. a stripe of the same color extends from the posterior border of the orbit to the corner of the mouth, passing through the tympanum. all of the dorsal parts are grayish white with large transverse brown bands, which are more expanded and less regularly outlined on the back than on the limbs. there is one of these on the occiput that is in a triangular shape. all of the venter is white." (free translation from french.) the holotype of _hyla leprieurii britti_ is a male having a snout-vent length of 48.1 mm. the diameter of the tympanum is 3.6 mm, 65.5 percent of the diameter of the eye. the skin on the dorsum is tubercular; the tubercles are small on head and on the dorsal surfaces of the limbs and slightly larger on the back. the penultimate phalanges of the fingers are not included in the webbing. melin (1941:43) stated: "above blackish brown with a very indistinct band between the eyes; iris with mottle of metallic lustre; hinder parts of upper jaw whitish; sides of body mottled with blackish brown; hind limbs (especially tibiae and tarsi) with narrow, diffuse cross bars; beneath whitish with slight brown mottle along jaw." we examined the type on 17 february 1969; at that time it was dull brown above with faint, narrow, dark brown, transverse bars on the back and dorsal surfaces of the limbs. a cream subocular spot was evident, and the venter was creamy white. melin (1941:42) stated that the holotype of _hyla leprieurii britti_ "... resembles a good deal _h. leprieurii_ dum. & bibr. as, however, it differs from the latter species by its very concave loreal region, small tympanum, and almost uniformly brownish colour, it may at least form a subspecies of _leprieurii_...." the pattern of narrow transverse bars on the backs of the holotypes of _h. leprieurii_ and _h. britti_ is a condition shared only by these two nominal taxa that are placed in _osteocephalus_. melin noted that _britti_ differed from _leprieurii_ in the depth of the loreal concavity and in the size of the tympanum. neither of these differences is noteworthy in comparison with series of specimens. the depth of the loreal concavity is a highly subjective character, and we note no differences between the types. the ratio of the diameter of the tympanum to the diameter of the eye is relatively smaller in both holotypes (0.698 in _leprieurii_--[f]; 0.655 in _britti_--[m]) than in series of fresh specimens from lago agrio, ecuador (0.652-0.884, mean 0.785 in 17 males; 0.700-0.909, mean 0.790 in 20 females). the smaller proportions in the types may be due to geographic variation or to shrinkage as a result of many years in preservative (130+ years for _leprieurii_; 45 for _britti_). comparisons of the holotypes with series of specimens from ecuador, guyana, and surinam indicate that one morphological species occurs throughout the upper amazon basin and the guianas and that both type specimens are representatives of one species. consequently, we consider _hyla leprieurii_ duméril and bibron, 1841, to be a monotypic species with _hyla leprieurii britti_ melin, 1941, as a junior synonym. in their account of _osteocephalus leprieurii_, cochran and goin (1970:323) stated: "the specimen described and illustrated (mcz 28042) has been directly compared with the types of _leprieurii_, _planiceps_, and _vilarsi_ by the junior author and there seems to be no doubt that all are conspecific. another specimen (cnhm 69716) has been directly compared with the types of _planiceps_ and _vilarsi_ and these, likewise, are considered conspecific." with this justification cochran and goin (1970:322) included _osteocephalus planiceps_ cope, 1874, and _hyla vilarsi_ melin, 1941, in the synonymy of _osteocephalus leprieurii_. we do not concur with cochran and goin's synonymy and contend that _planiceps_ and _vilarsi_ are synonyms of _osteocephalus taurinus_; we give our reasons in the account of that species. we have examined the specimens listed as _o. leprieurii_ by cochran and goin; several of them, including cnhm (= fmnh) 69716, are _taurinus_. thus, due to cochran and goin's confusion of two taxa, their comparisons of certain specimens with types has little meaning. cochran and goin did not include _hyla leprieurii britti_ in their synonymy of _osteocephalus leprieurii_ but did discuss the name in their account of _osteocephalus orcesi_ (= _o. verrucigerus_), as follows (1970:319): "when we first examined one of the specimens we felt sure that we had melin's _hyla britti_ at hand, but on direct comparison with the type of _britti_ the two proved to be different. after studying the type of _orcesi_ (sunhm 13150) we have no doubt that the specimens at hand are _orcesi_ and that _britti_ is a different, probably valid species." _diagnosis._--1) size moderate, sexual dimorphism evident; maximum observed snout-vent length in males 48.4 mm, in females, 61.5 mm; 2) skin on dorsum in males bearing numerous, minute, spinous tubercles; 3) skin on flanks smooth; 4) web extending to base of antepenultimate phalange on inner edge of third finger; 5) dorsum tan or olive-brown with transverse brown or olive bars; 6) venter creamy white or pale tan without markings; 7) lips marked with creamy tan labial stripe and suborbital spot; 8) flanks pale tan with no markings; 9) dermal roofing bones of skull lacking exostosis; 10) dermal sphenethmoid absent; 11) nasals juxtaposed medially; 12) anteromedial margin of frontoparietal between midand anterior levels of orbit; 13) frontoparietal fontanelle partially exposed; 14) palatine not serrate; 15) parasphenoid lacking odontoids; 16) zygomatic ramus of squamosal extending about one-half of distance to maxillary arch; 17) transverse processes of presacral vertebrae 3-8 about equal in width to one another and to sacral diapophyses; 18) intermandibularis and submentalis muscles connected; 19) supramandibular portion of interhyoideus forming simple tubular posterolateral extension; associated skin unmodified. _osteocephalus leprieurii_ differs from all other members of the genus by having transverse dark bars on the back. two other hylids (_hyla lanciformis_ and _multifasciata_) in the amazon basin have transverse dark marks on the dorsum. both of these differ from _leprieurii_ by having pointed snouts, much longer hind limbs, and smooth skin dorsally. _distribution._--the periphery of the amazon basin, in the guianas and the upper part of the basin in southern colombia, ecuador, perú, and extreme western brasil (fig. 9). most localities are at elevations of less than 500 m, but the species ascends the lower andean slopes to elevations of 1100 m. 265 specimens from 31 localities. [illustration: fig. 9. distribution of _osteocephalus leprieurii_ (circles) and _o. verrucigerus_ (triangles).] _remarks._--most adults of _leprieurii_ have distinct transverse markings on the back; these are variable in width, extent, and arrangement. in some specimens, such as usnm 166557, some of the transverse bars are fragmented into spots; in a few specimens the dorsal pattern consists solely of small dark spots arranged in transverse rows. such specimens have a dorsal pattern resembling that of some _taurinus_. the transverse nature of the dorsal markings is further modified in some specimens, such as usnm 166555, in which the dark bars are fragmented and oblique. extreme ontogenetic change in color pattern is exhibited by this species (fig. 10). juveniles having snout-vent lengths of less than 28 mm have an olive-brown dorsum with a pale cream stripe across the head and broad, cream, dorsolateral stripes; transverse dark bars are absent on the body and limbs. individuals having snout-vent lengths of 30-35 mm have dark brown transverse bars on the back and limbs but still retain the light dorsolateral stripes, whereas the stripes are lost in larger individuals. [illustration: fig. 10. ontogenetic change in color pattern in _osteocephalus leprieurii_: a. ku 126644; b. ku 126640; c. ku 126625. ×2.] coloration in life of specimens from lago agrio, ecuador: "in males the dorsal ground color varies from dark brown to ochre-tan; dorsal markings uniformly dark brown. most specimens have dark brown and cream anal stripes; labial area cream-colored. flanks vary from tan to white. ventral coloration varies from salmon to tan to white. the iris is bronze with a greenish cast and black reticulations. in females the dorsal coloration is the same as in males, except that dark marks tend to be outlined with cream; venter tannish salmon." (w. e. duellman, field notes, 12 may 1969). =osteocephalus pearsoni= (gaige) _hyla pearsoni_ gaige, 1929:3 [holotype.--ummz 57548 from the upper río beni, below mouth of río mapiri, departamento el beni, bolivia; n. e. pearson collector]. _osteocephalus pearsoni_--goin, 1961:13. _justification of synonymy._--goin (1961:13) suggested that _hyla pearsoni_ gaige was an _osteocephalus_, but cochran and goin (1970:217) considered _pearsoni_ to be a _hyla_. the presence of exostosed dermal roofing bones, angulate prevomerine dentigerous processes, and the structure of the vocal sacs are characters which place the species in _osteocephalus_. _diagnosis._--1) size moderate, sexual dimorphism evident; maximum observed snout-vent length in males 46.2 mm, in females 54.7 mm; 2) skin on dorsum in males bearing a few, small, scattered non-spinous tubercles; 3) skin on flanks smooth; 4) web extending to base of antepenultimate phalange on inner edge of third finger; 5) dorsum tan with irregular brown blotches; 6) venter cream with fine brown reticulations; 7) lips dark with pale vertical bar below eye; 8) flanks pale tan with round, brown spots; 9) dermal roofing bones of skull slightly exostosed; 10) dermal sphenethmoid absent; 11) nasals narrowly separated medially; 12) anteromedial margin of frontoparietal between midand anterior levels of orbit; 13) frontoparietal fontanelle covered; 14) palatine not serrate; 15) parasphenoid lacking odontoids; 16) zygomatic ramus of squamosal extending about one-half distance to maxillary arch; 17) transverse processes of third presacral vertebra approximately equal in width to sacral diapophyses; transverse processes of presacral vertebrae 3-8 subequal in width; 18) intermandibularis and submentalis muscles connected; 19) supramandibular portion of interhyoideus extensively developed; associated skin forming broad loose fold. _osteocephalus pearsoni_ can be distinguished most readily from other members of the genus by the brown reticulate pattern on the venter, round brown spots on the flanks, and smooth skin on the flanks. also, it is the least tuberculate species in the genus. _distribution._--upper amazon basin and amazonian slopes of the andes in central perú (1620 m in río ucayali drainage) and northern bolivia (less than 500 m in río beni drainage) (fig. 8). 6 specimens from 3 localities. _remarks._--the specimen from yaupi, perú (ku 136312) is a subadult female having a snout-vent length of 39.8 mm. in life the coloration was: "dorsum light pinkish brown with large rich chocolate brown blotch from eyes to anterior tips of ilia; numerous small chocolate blotches on flanks; dorsal surfaces of thighs and shanks, canthus, and supraorbital region to insertion of forearm chocolate brown; supralabial border and short bar from eye to lip bronze-white; venter bronze-white with numerous tiny chocolate brown flecks [tending to form reticulations on throat and chest]; anterior and posterior surfaces of thighs light olive-brown; iris largely black with gold flecks." (thomas h. fritts, field notes, 23 march 1970.) on the basis of this one subadult, it seems likely that reticulations on the venter develop with age. =osteocephalus taurinus= steindachner _osteocephalus taurinus_ steindachner, 1862:77 [holotype.--nhmw 16492 from barra do río negro, manáus, territorio do amazonas, brasil; johann natterer collector]. _osteocephalus flavolineatus_ steindachner, 1862:80 [holotype.--nhmw 16495 from cucuí, territorio do amazonas, brasil; johann natterer collector]. _trachycephalus (osteocephalus) taurinus_ steindachner, 1867:64. _osteocephalus planiceps_ cope. 1874:122 [holotype.--ansp 11399 from nauta, departamento de loreto, perú; james orton collector]. new synonymy. _hyla taurina_--boulenger, 1882:363 [synonymized _osteocephalus flavolineatus_ steindachner, 1862, with _o. taurinus_ steindachner, 1862]. _hyla planiceps_--boulenger, 1882:364. _hyla (trachycephalus) vilarsi_ melin, 1941:40 [holotype.--nhmg 488 from taracuá, río uaupés, territorio do amazonas, brasil; douglas melin collector]. (_fide_ bokermann, 1966:64.) _hyla depressa_ andersson, 1945:73 [holotype.--nhrm 1966 from the río pastaza watershed (? provincia pastaza), ecuador; william clarke-macintyre collector]. new synonymy. _justification of synonymy._--the holotype of _osteocephalus taurinus_ is a female having a snout-vent length of 103.9 mm. the diameter of the tympanum is 6.8 mm, 77.3 percent of the diameter of the eye. the skull is strongly exostosed, and the lateral edges of the frontoparietals are elevated so as to form distinct ridges. the skin on the dorsum is smooth. when we examined the type on 5 august 1969, the specimen was soft and badly faded to a pale creamy tan with pale brown transverse bars on the hind limbs and spots on the flanks. steindachner (1862:79) described the coloration of the type: "in the preserved specimen the dorsum of the entire body, including fore and hind limbs, is a light yellow-brown color, which becomes lighter towards the venter. the belly is whitish, as are the undersides of the arms and legs. the throat is indistinctly marbled with brown. roundish dark brown flecks are randomly distributed in a considerable number along the side of the body up to the eye; the tympanum is more or less fully surrounded by brown. a few discrete spots, always more or less drawn out in length, on the sides of the body, are also found on the posterior part of the back. the dorsal surfaces of the fore and hind feet are marked with somewhat obliquely arranged brown transverse bands, which are more intensively colored near the margin than in the middle of the band." (free translation from german.) the holotype of _osteocephalus flavolineatus_ is a female having a snout-vent length of 81.8 mm. the diameter of the tympanum is 6.0 mm, 71.4 percent of the diameter of the eye. the skull is strongly exostosed, and the lateral edges of the frontoparietals are elevated so as to form a ridge on each side. the skin on the dorsum is very weakly tuberculate. we examined the type on 9 august 1969 and found it to be in excellent condition. the color pattern is unchanged from that described by steindachner (1862:81). the dorsum is tan with irregular brown blotches on the back, spots on the flanks, and transverse bars on the limbs. a narrow creamy white, middorsal stripe extends from the snout to the vent. the subocular area is creamy tan, and the venter is tan. boulenger (1882:363) questionably synonymized _flavolineatus_ with _taurinus_. we have observed that a middorsal cream stripe occurs in about 10 percent of the specimens of _taurinus_ and in some specimens of _buckleyi_. this is a common color morph in many species of _eleutherodactylus_. in the absence of distinguishing morphological characteristics we can only conclude that the middorsal stripe is a pattern variant and that boulenger was correct in synonymizing _flavolineatus_ with _taurinus_. the holotype of _osteocephalus planiceps_ is a male having a snout-vent length of 58.5 mm. the diameter of the tympanum is 4.9 mm, 77.8 percent of the diameter of the eye. the skull is moderately exostosed, and the lateral edges of the frontoparietals are distinctly elevated. the skin on the dorsum is tuberculate. cope (1874:122) described the coloration of the type as follows: "color above uniform dark brown, concealed surfaces on the limbs similar and without any markings. sides a little varied with the white of the belt. a light border to the upper lip, and lighter line from the orbit to the angle of the mouth; dermal scapular fold pale edged. femur and tibia with dark crossbands on the exposed surfaces." we examined the holotype on 25 september 1969, and found it to be soft and rubbed. the coloration remains much the same as described by cope, who provided no means of distinguishing _planiceps_ from _taurinus_. the coloration and morphometric and structural characters of the type of _planiceps_ all fall within the range of variation displayed by series of _o. taurinus_ from the upper amazon basin. the type of _hyla vilarsi_ is a gravid female having a snout-vent length of 62.7 mm. the diameter of the tympanum is 4.8 mm, 73.8 percent of the diameter of the eye. the dorsal roofing bones of the skull are moderately exostosed, and the lateral edges of the frontoparietals are distinctly elevated. the skin on the dorsum is smooth. melin (1941:42) described the coloration of the holotype as follows: "above uniform reddish brown; upper eyelids and sides of head darkish brown; below the rostral edge a narrow dark band, continuing as a broader light-edged one through the eye and tympanum towards the base of the forelimb and then farther on continuing along the sides as a line of black spots; sides of upper jaw whitish with traces of dark cross bars (one distinct under the eye); sides of body darkish with black spots and marble, often on a whitish ground; thighs, tibiae, and tarsi each with two broad light-edged, dark cross bars on a brownish ground (less distinct on thighs); sides of thighs finely mottled with brown; beneath whitish with small, sparse spots along jaw, on the chest and sides." we examined the type on 17 february 1969, at which time the specimen was somewhat desiccated, especially the hands and feet. the coloration remains much the same as described by melin, except that he failed to note the presence of four elongate spots on the back. the status of the names _osteocephalus planiceps_ cope and _hyla vilarsi_ melin was confused by cochran and goin (1970:322), who assigned these names to the synonymy of _o. leprieurii_. bokermann (1966:64) placed _hyla vilarsi_ in the synonymy of _osteocephalus taurinus_ without justification. the type specimens of both _planiceps_ and _vilarsi_ have moderately exostosed dermal roofing bones and distinct cranial ridges. the type of _planiceps_ has moderately large tubercles on the dorsum, and the type of _vilarsi_ has spots on the throat, chest, and flanks and longitudinal markings on the back. all of these features are characteristic of _taurinus_ and not of _leprieurii_, which lacks exostosis and cranial ridges and has transverse markings on the back, no spots on the throat, chest, and flanks, and in males has small dorsal tubercles. the type of _hyla depressa_ is a male having a snout-vent length of 69.8 mm. the diameter of the tympanum is 5.2 mm, 77.6 percent of the diameter of the eye. the dorsal roofing bones of the skull are moderately exostosed, and the lateral edges of the frontoparietals are elevated. the skin on the dorsum is tuberculate. the dorsum is dull brown with a broad darker brown longitudinal mark having indistinct lateral edges from the snout to the post-sacral area. a narrow cream middorsal line extends from the snout to the vent. the side of the head is dark brown, palest posteroventral to the orbit. the posterior surfaces of the thighs are dull brown; the flanks are pale brown, and the ventral surfaces are pale creamy tan. dark brown transverse bars are present on the limbs. when we examined the type on 3 january 1969, it was in excellent condition. andersson (1945:75) contrasted the type of _hyla depressa_ with _leprieurii_ and _buckleyi_, but he did not compare his specimen with _taurinus_, from which it exhibits no distinguishing features. _osteocephalus taurinus_ is a widespread and variable species, and it has received several specific names. we are convinced that _osteocephalus taurinus_ steindachner, 1862, is the oldest available name for this large amazonian species. the following names are junior synonyms: _osteocephalus flavolineatus_ steindachner, 1862; _osteocephalus planiceps_ cope, 1874; _hyla (trachycephalus) vilarsi_ melin, 1941; _hyla depressa_ andersson, 1945. _diagnosis._--1) size large; sexual dimorphism evident; maximum observed snout-vent length in males 84.6 mm, in females 104 mm; 2) skin on dorsum in males bearing many moderately large, spinous tubercles; 3) skin on flanks smooth; 4) web extending to middle of antepenultimate phalange on inner edge of third finger; 5) dorsum brown usually with a large medial dark brown blotch or, less frequently, several dark spots; narrow middorsal yellow line present in some; 6) venter cream or tan with or without small, irregular brown flecks; 7) lips brown with vertical cream bar below eye in some, expanded into pale labial stripe posteriorly in some females; 8) flanks tan or cream with or without small, irregular brown spots; 9) dermal roofing bones of skull exostosed, casqued, and co-ossified (in large adults); 10) dermal sphenethmoid present; 11) nasals juxtaposed medially; 12) anteromedial margin of frontoparietals at mid-level of orbit; 13) frontoparietal fontanelle covered; 14) palatine serrate; 15) parasphenoid bearing odontoids; 16) zygomatic ramus of squamosal usually articulating with maxillary arch; 17) transverse processes of third presacral vertebra approximately equal in width to sacral diapophyses; transverse processes of presacral vertebrae 3-8 subequal in width; 18) intermandibularis and submentalis muscles connected; 19) supramandibular portion of interhyoideus extensively developed; associated skin forming everted pouch. the moderately rugose dorsum (in males), large size, extensive webbing on the hand, and frontoparietal flanges in adults serve to distinguish _taurinus_ from other members of the genus. _distribution._--the amazon basin, the upper orinoco basin, and the guianas. most localities are below 500 m, but the species ascends the lower amazonian slopes of the andes to elevations of about 1000 m (fig. 11). a record from caracas, venezuela, and those from provincia carchi and provincia esmeraldas, ecuador, are considered to be erroneous. the latter specimens were included in a collection sold to the university of illinois; contained in the collection are many common amazonian species unknown from the pacific lowlands. 516 specimens from 151 localities. [illustration: fig. 11. distribution of _osteocephalus taurinus_.] _remarks._--this widespread species is highly variable in size and coloration. striking differences in snout-vent length are evident in series from various parts of the range. the smallest calling males (cas-su 12351-6 from rio tapirapé, brasil) have snout-vent lengths of 46.5-60.3 (mean 53.3) mm, whereas the largest (fmnh 140254, ku 92243-6, wcab 9997, 10001, 10003-4 from igarapé marmelo, brasil) have snout-vent lengths of 71.5-84.6 (mean 77.6) mm. mean values of snout-vent lengths of males from other localities are: río pastaza drainage, ecuador 73.8 mm, surinam 67.7 mm, río ucayali drainage, perú 57.6 mm, and guyana 55.5 mm. although the difference between the smallest and largest adults is highly significant, populations bridging the gap do exist. furthermore, the geographic arrangement of small versus large frogs is a confusing mosaic. we have entertained the thought that we have included more than one species in _taurinus_, but on the basis of preserved specimens we are unable to detect consistent differences distinguishing two or more taxa. the coloration and pattern of _taurinus_ are so variable that no one series of statements can describe samples drawn from the entire range of species. we have been unable to determine geographic trends in color pattern; instead the variation within a given sample can encompass the variety known in most other samples. two minor exceptions do exist. a narrow middorsal light stripe is present in some individuals from throughout the range, but striped specimens are most common in the upper amazon basin. the absence of dorsal markings is uncommon in the entire species, but it is most frequent in individuals from the guianas. a few individuals, such as ku 105230, have scattered white spots on the dorsum. the coloration of four males in life from lago agrio, ecuador (ku 126652-5) was: "dorsal ground color tan to dark brown with darker brown markings. flanks creamy tan to yellow with brown or black flecks or mottling. venter uniform creamy yellow or yellow with brown spots or reticulations. iris greenish yellow with radiating black streaks and a median, horizontal reddish brown streak." (w. e. duellman, field notes, 12 may 1969.) a female from santa cecilia, ecuador (ku 123173), was: "dorsum mottled olive-green and tan. flanks tan with brown spots. belly and throat creamy white, becoming tan posteriorly. edge of upper jaw olive-green." (w. e. duellman, field notes, 16 june 1968.) another female from santa cecilia (ku 123175), was: "brown dorsally with cream-colored mottling. transverse bars on legs darker brown with cream-colored edges. margin of upper lip creamy yellow. anterior and posterior surfaces of thighs tan. flanks white with brown spots. venter creamy white. iris greenish bronze with heavy radiating reticulations of black." (w. e. duellman, field notes, 22 july 1968.) the tendency for females to have a labial stripe posteriorly and the absence of dorsal tubercles in females has resulted in the identification of many such specimens as _o. leprieurii_. ontogenetic change in coloration is slight in _taurinus_. most juveniles (less than 40 mm in snout-vent length) can be identified readily. there is a tendency for the dorsal markings of juveniles to consist of several small spots. apparently with growth the spots usually coalesce, forming a large median blotch, but some adults retain the juvenile pattern. cochran and goin (1970:251) erroneously identified several juveniles from colombia as _hyla palpebrogranulata_ andersson. =osteocephalus verrucigerus= (werner) _hyla verrucigera_ werner, 1901:601 [holotype.--zmb 16589 from "ecuador"; richard haensch collector]. _hyla riopastazae_ andersson, 1945:72 [holotype.--nhrm 1960 from baños, río pastaza, provincia tungurahua, ecuador; william clarke-macintyre collector]. _hyla orcesi_ funkhouser, 1956:78 [holotype.--cas-su 13150 from río pacayacu, tributary of río cotapino, provincia napo, ecuador; collector unknown]. _osteocephalus orcesi_--cochran and goin, 1970:317. _osteocephalus verrucigerus_--trueb and duellman, 1970:601 [synonymized _hyla riopastazae_ andersson, 1945, and _hyla orcesi_ funkhouser, 1956, with _hyla verrucigera_ werner, 1901]. _justification of synonymy._--trueb and duellman (1970:605) discussed the assignment of the names in the synonymy of _o. verrucigerus_; only a brief resumé is given here. the extant type of _hyla verrucigera_ is a juvenile male having a snout-vent length of 32.0 mm. the dorsum is smooth except for tubercles on the eyelids; the skin is loose, and the body is soft. the specimen is faded to a pale brown; indistinct dark spots are present on the back, and transverse bars are evident on the limbs. the holotype of _hyla riopastazae_ is a gravid female having a snout-vent length of 64.7 mm. the dorsum is smooth. the dorsal ground color is pale brown with indistinct brown transverse bars on the limbs. the throat, chest, and belly are cream with brown spots and mottling. the holotype of _hyla orcesi_ is an adult male having a snout-vent length of 52.6 mm. the dorsum is heavily tuberculate. the dorsum is dark brown with faint transverse bars on the forearms and feet; the ventral surfaces are creamy brown. trueb and duellman (1970) provided conclusive evidence that the types of _h. verrucigera_, _riopastazae_, and _orcesi_ are a juvenile, adult female, and adult male, respectively, of one species, the earliest available name for which is _hyla verrucigera_ werner, 1901. _diagnosis._--1) size moderate, sexual dimorphism evident; maximum observed snout-vent length in males 54.3 mm, in females 65.8 mm; 2) skin on dorsum in males bearing large, keratinized tubercles; 3) skin on flanks smooth; 4) web extending to base of antepenultimate phalange on inner edge of third finger; 5) dorsum uniformly dark brown or black, with tan snout in females; 6) venter creamy white, heavily mottled with black or dark brown, especially in females; 7) lips marked with pale tan labial stripe and suborbital bar; 8) flanks dull reddish brown; 9) dermal roofing bones of skull lacking exostosis; 10) dermal sphenethmoid absent; 11) nasals widely separated medially; 12) anteromedial margin of frontoparietals at anterior border of orbit; 13) frontoparietal fontanelle covered; 14) palatine serrate; 15) parasphenoid bearing odontoids; 16) zygomatic ramus of squamosal extending approximately one-half of distance to maxillary arch; 17) transverse processes of third presacral vertebra approximately equal in width to sacral diapophyses; transverse processes of presacral vertebrae 3-8 subequal in width; 18) intermandibularis and submentalis muscles connected; 19) supramandibular portion of interhyoideus forming simple, tubular, posterolateral extension; associated skin unmodified. _osteocephalus verrucigerus_ can be distinguished from other members of the genus by its uniformly dark dorsum, heavily mottled venter, and large, spinous tubercles on the dorsum in males. _distribution._--lower amazonian slopes (500-1840 m) of the andes and on the western fringe of the amazon basin in ecuador and perú; one locality (acevedo) in upper río magdalena drainage in colombia (fig. 9). 40 specimens from 13 localities. _remarks._--in life the dorsum in males is dull olive-green; the groin, anterior and posterior surfaces of the thighs, inner surfaces of limbs, and upper arms are dark brown. the ventral surfaces of the limbs are pinkish tan; the other ventral surfaces are pale creamy tan with reddish brown spots. the suborbital spot is pale greenish tan, and the iris is deep reddish brown. in females the dorsum is dull olive-brown; the anterior part of the head is tan, and the suborbital spot is yellowish tan. the groin and hidden surfaces of the limbs are dark reddish brown. the ventral surfaces of the limbs are brown; the throat and chest are creamy white, and the belly is reddish tan, both with dark brown mottling. considerable ontogenetic change occurs in coloration. juveniles are pale above with a dark median dorsal blotch and dark transverse bars on the limbs. the venter is white. the change consists principally of an increase in dark pigment and subsequent obliteration of the juvenile pattern. tadpoles of this species have moderately long tails with low fins, robust bodies, two rows of labial papillae with median part of the upper lip bare, and two upper and five lower rows of teeth. trueb and duellman (1970) described the eggs, tadpoles, mating call, and variation in the adults. generic relationships among the 33 genera currently recognized in the family hylidae, there are two basic types of vocal sac structure (duellman, 1970b), namely the subgular type and the lateral type. only four hylid genera, all neotropical lowland groups, are known to possess paired lateral vocal sacs; these are _osteocephalus_, _argenteohyla_, _phrynohyas_, and _trachycephalus_. the geographical distributions and morphological characteristics of these four genera suggest that they are more closely related to one another than with any other hylid genera. of the four genera, _osteocephalus_ is the most generalized in morphology, and, like _phrynohyas_, has no specialized habits. _osteocephalus_ and _argenteohyla_ are similarly distinguished from _phrynohyas_ and _trachycephalus_ on the basis of vocal sac structure. the vocal sacs of _osteocephalus_ and _argenteohyla_ are posterior and protrude posterolateral to the angles of the jaws when they are inflated, whereas those of _phrynohyas_ and _trachycephalus_ are more lateral and protrude posterior to the angles of the jaws when inflated. although _osteocephalus_ and _argenteohyla_ have similar vocal sac structure, they are obviously distinct. the monotypic _argenteohyla_ is a rather specialized, semifossorial frog (trueb, 1970b), characterized by smooth skin, moderate-sized digital discs, and a large inner metatarsal tubercle. the general architecture of the skull is not unlike that of _osteocephalus_; the skulls of both are well roofed, broader than long, and characterized by posterolaterally oriented parasphenoid alae. _argenteohyla_ bears small, slightly curved prevomerine dentigerous processes in contrast to the large, angular processes of _osteocephalus_. the skull of _argenteohyla_ shows specializations, apparently adaptations to its semifossorial mode of existence, which further distinguish the genus from _osteocephalus_. in comparison with _osteocephalus_, the cranium of _argenteohyla_ is slightly depressed anteriorly, the roofing bones extensively casqued, and the palatines robust. osteologically, _osteocephalus_ more closely resembles _phrynohyas_ than either of the other two genera, but _osteocephalus_ and _phrynohyas_ are clearly distinct on the basis of their respective vocal sac structure. like _osteocephalus_, skulls of the members of the genus _phrynohyas_ are broader than long, have extensive dermal roofing bones, and have posterolaterally oriented parasphenoid alae. in contrast to _osteocephalus_, the dentigerous processes of the prevomers are curved, rather than angular in _phrynohyas_. furthermore, the latter genus is singularly distinguished from _osteocephalus_, _argenteohyla_, and _trachycephalus_ by having extensively developed parotoid glands that produce a viscous, milky volatile secretion. _trachycephalus_ is the most readily identifiable of the four genera under discussion. members of this genus are large frogs with heavily casqued and co-ossified skulls (trueb, 1970a). the dermal roofing bones bear ornate and characteristic patterns of sculpturing. the medial ramus of the pterygoid does not articulate with the otic capsule, and the parasphenoid alae are laterally, rather than posterolaterally, oriented. a dermal sphenethmoid is present, and the parasphenoid bears odontoids. the basic structure of the skull has many characters in common with both _osteocephalus_ and _phrynohyas_. the obvious modifications of dermal roofing bones and of palatal and suspensory elements seem to be specializations adapting members of the genus _trachycephalus_ to their peculiar phragmotic habits. the vocal sac structure of _trachycephalus_ is like that of _phrynohyas_ and therefore further distinguishes it from _osteocephalus_. morphologically, _osteocephalus_ seems to be sufficiently diverse and generalized so as to represent a modern derivative of an ancestral type which might have given rise to _phrynohyas_, _trachycephalus_, and _argenteohyla_. the specialized vocal sac structure in _phrynohyas_ and _trachycephalus_ suggests that these two genera may be rather closely allied and represent a single phyletic line from an ancestral stock similar to _osteocephalus_. _argenteohyla_ is quite distinct from _phrynohyas_ and _trachycephalus_ and apparently represents a distinct phyletic line from the ancestral stock. occurrence of _osteocephalus_ in amazonian ecuador all of our observations on members of this genus have been made at four localities: 1) santa cecilia at an elevation of 340 meters on the río aguarico, a tributary of the río napo, 2) lago agrio, 330 meters, about 14 kilometers east of santa cecilia, 3) puerto libre, 570 meters, on the río aguarico just east of its formation by the confluence of the río cofanes and río chingua, and 4) south slope of the cordillera del dué, above the río coca, 1150 meters. _osteocephalus leprieurii_ was found at all four localities, and _buckleyi_ was found at all but the last; _taurinus_ was found at santa cecilia and lago agrio, and _verrucigerus_ was found only in the cordillera del dué. our data are based on collections of 113 frogs and three lots of tadpoles, as well as observations on calling sites and young. the observations are summarized by species, as follows: _osteocephalus buckleyi._--no breeding activity was observed. males were found only at night in march, june, and july. one was perched on a _heliconia_ leaf in a swamp at puerto libre, and two were on bushes in the forest at santa cecilia. a gravid female was found on a recently felled tree at lago agrio on the night of 12 may 1969. _osteocephalus leprieurii._--males were heard calling sporadically at puerto libre in july 1968, and at santa cecilia in may 1969. a small chorus was found on the night of 12 may 1969 at lago agrio, where the frogs were perched on branches of fallen trees over a temporary pool. the call is a soft rattling chuckle. in late april and may many gravid females and males with well-developed nuptial excrescences were obtained from trees as they were felled at lago agrio. the reproductive condition of the frogs indicates that they probably breed in may. one individual called nearly every night from a large tree at puerto libre between 4-17 july 1968. the tree was felled on the latter date, but no frog was found. two nights later apparently the same individual called from a bromeliad at a height of about 10 m on a large bamboo adjacent to the felled tree; the frog was collected when the bamboo was cut down. throughout the rainy months that we have worked in ecuador (april-august) we have found occasional individuals perched on bushes or low trees at night. large numbers of adults were observed only during a clearing operation which resulted in the felling of many large trees. thus, it seems likely that _leprieurii_ is a tree-top inhabitant. a partially digested adult male was removed from the stomach of a _hemiphractus proboscideus_. at santa cecilia many recently metamorphosed young and juveniles were found in june and july 1968. most of these were on low bushes or herbs in swamp forest at night; some were found in unfolded _heliconia_ leaves by day, and one was observed on the forest floor by day. snout-vent lengths of 18 specimens are 12.3-17.0 (mean 15.1) mm. the smaller frogs were recently metamorphosed as evidenced by the melanophore deposits above the vent. the coloration of the young is strikingly different from that of the adults (see account of _o. leprieurii_), so the association of the young and adults was not made until individuals with intermediate patterns were obtained at lago agrio in may 1969. probably juveniles obtained in june and july are the offspring of an april or may breeding. we have been unable to associate tadpoles with this species. _osteocephalus taurinus._--a small chorus occurred at lago agrio on 12 may 1969. males were calling from the ground adjacent to a small pool amidst recently felled trees. the males were very wary and, when approached, jumped onto limbs and ran up branches; this behavior was noted by bokermann (1964). the call consists of a series of low-pitched, short notes--like a slow trill--four to six notes per call group. call groups are repeated two, three, or four times followed by a lapse of several minutes. although no amplectant pairs were found, several gravid females were collected at lago agrio in may, so it can be safely assumed that the species breeds in may. from april through july occasional individuals were observed on bushes and trees at night. during clearing operations at lago agrio several individuals were obtained from the tops of trees as they were felled. _osteocephalus verrucigerus._--observations were made in a broad, shallow ravine, in which there was a small stream. on 2-4 august 1968, males were observed calling from low bushes and rocks at the edge of a quiet pool in the stream. the call consists of a series of well-pulsed, low-pitched, guttural notes produced at the rate of 5-10 per minute. one amplectant pair was found at the base of a bush adjacent to the pool on 3 august. another female was found on a branch of a tree 2 m above the ground and 10 m from the stream. tadpoles of this species were found in the quiet silt-bottomed pool. specimens examined the localities for each of the specimens examined are given in the following paragraphs. the arrangement of the data is as follows: alphabetically by country, state (department or province), and locality; alphabetically by the first letter in the abbreviations for the museums, and numerically after each museum abbreviation. specimens lacking precise locality data are listed first in the most restricted political unit possible; localities which have not been found on maps or the positions of which are not known to us are given in quotation marks. where more than one specimen is included under one museum number, the number of specimens is given in parentheses after the museum number. unless noted otherwise, all specimens are alcoholics. _osteocephalus buckleyi_ bolivia: _el beni_: ivón, bmnh 1967.2070-1. _santa cruz_: buenavista, cm 4333, 4339, ummz 66563-5. brasil: _amapá_: no specific locality, wcab 13284. colombia: _amazonas_: río guacaya, usnm 152759. _huila_: acevedo, río suaza, fmnh 69702. _nariño_: rumiyacu, fmnh 54756. _meta_: río guejar, campamento la macarena, usnm 152199. ecuador: no specific locality, nhmw 6209, wcab 35499. _chimborazo_: pallatanga, bmnh 1947.2.13.46; santiago, fmnh 42529. _morona-santiago_: "río santiago" (= río zamora), mizs 2950. _napo_: lago agrio, ku 126646; puerto libre, río aguarico, ku 123172; santa cecilia, aum 8138, ku 105208-9, 109506, 123171. _pastaza_: alpayacu, bmnh 1912.11.1.64; canelos, bmnh 1947.2.13.40-1, 1947.2.13.43-5; colonia mena, río conambo, zsm 33/1962; don tomás, usnm 166014; guaché, río pastaza, amnh 79986; río bobonaza, usnm 166005; río capahuari, usnm 166554; río conambo at río shiona-yacu, usnm 166018; río copataza, upper río pastaza, usnm 166007-13; río pastaza, nhrm 1946; río pucyacu, usnm 165997 (skeleton), 165998-6001; río rutuno, usnm 166006; río villano, usnm 166002-4; sarayacu, bmnh 1947.2.13.36-9, mcz 26090, zmb 10166. guyana: _mazaruni-potaro_: kartabo, amnh 70971; membaru river, upper mazaruni river, ummz 85168; oko mountains, fmnh 26722-3. _north west_: amakura river, haulover, ummz 83558-9. _rupununi_: marudi river, amnh 46233; shudi-kar-wau, amnh 49252. _west demerara_: dunoon, ummz 52449, 52508. perú: _junín_: chanchamayo, bmnh 1911.12.13.79-80. _loreto_: andoas, amnh 79984-5; cashiboya, amnh 43454; san antonio, río itaya, amnh 43218. _puno_: yahuaramayo, bmnh 1913.2.25.7. surinam: _suriname_: powakka, cm 44217. south america: no specific locality, nhmw 6208. _osteocephalus leprieurii_ brasil: _acre_: tarauacá, fmnh 83247. _amazonas_: rio javarí, benjamin constant, cas-su 12620; río uaupés, north of rio japú, nhmg 489. colombia: _amazonas_: gino-goje, lower río apoporis, mcz 28038, 28040-2, 28044, usnm 152136-8. ecuador: no specific locality, wcab 35452-3; "napo-pastaza," usnm 166571. _napo_: avila, ummz 92093; south slope cordillera del dué, ku 123170; lago agrio, ku 125961-2 (skeletons), 126611-44, ummz 129326 (2); limón cocha, río napo, ku 99210-6, uimnh 63087-9, 63098, 63106-9, 63118-9, 64802-4, 64858, 87998-9, 88001-30, 88437-8, 88580, 88604-5, 89852-97, 89999-90000; loreto, cas-su 11439, wcab 36526; puerto libre, río aguarico, ku 123190-1; puerto napo, uimnh 55818-20; río cotapino, ummz 92094; río napo, ummz 92078; santa cecilia, aum 8099, 8102, 8113-5, 8127-9, 8131, 8137, 8139-46, 8148, ku 105210-20, 109509-11, 111971, 122964-87, 123169, 126645. _pastaza_: canelos, bmnh 1947.2.13.42, ku 120915; río alpayacu, ummz 92079; río arajuno, usnm 166560-2, wcab 40176; río oglán, usnm 16655203, 166558; río rutuno, usnm 166559; río shilcayacu, below puyo, usnm 166557; río villano, usnm 166551. french guiana: no specific locality, mnhn 4629. _inini_: lunier river, mnhn 98/217. guyana: _mazaruni-potaro_: kartabo, amnh 70967-8, 70972, 70976. _rupununi_: shudi-kar-wau, amnh 49255. _west demerara_: demerara falls, bmnh 72.10.16.23, 72.10.16.37-8. perú: _loreto_: estirón, río ampiyacu, mzusp 31033-4; pebas, cas-su 3158, 3160; roaboya, amnh 43064. surinam: no specific locality, mcz 2036, rmnh 11468. _marowijne_: camp 3, rmnh 13045-6; wane creek north, rmnh 11469-70. _saramacca_: right coppename river, rmnh 11467. _osteocephalus pearsoni_ bolivia: _el beni_: upper río beni, below mouth of río mapiri, mcz 15565, ummz 57548, 67464-5; rurrenbaque, ummz 57533. perú: _pasco_: yaupi, ku 136312. _osteocephalus taurinus_ bolivia: _el beni_: ivón, bmnh 1967.2040; reyes, ummz 57532. _la paz_: san ernesto, mapiri district, bmnh 1901.8.2.54. _santa cruz_: buenavista, amnh 33951-2, 33958, bmnh 1927.8.1.19, 1927.8.1.118, fmnh 27091, ummz 63319-21, 63959(2), 63961(2), 66566(2), 66567, 66568(2), 66569 (2), 66570, 66571(2), 66575-6, 68196; río mamore, 2 km n boca chaparé, amnh 79324; sara, cm 3840-1; surutu, cm 3814-5. brasil: no specific locality: "interior," bmnh 74.7.16.8-9. _acre_: plácido de castro, mzusp 6518; tarauacá, wcab 2496. _amazonas_: cucuí, nhmw 16495; manacapurú, zmb 28492, zsm 278/1925; manáus, mcz 56281, nhmw 16492; maués, amnh 69623, 76177; taracuá, nhmg 488, wcab 18463-4. _mato grosso_: mabuca, mzusp 4272; posto coluene, rio xingú, wcab 812; "puerto cabello," amnh 3154; tapirapé, amnh 73647-62, cas-su 12351-6, mnhn 46/324. _pará_: no specific locality, mpeg 623-6; belém, ku 129866; cachimbo, fmnh 175876, uimnh 42149, wcab 813; cametá, nhmw 15892; gurupá, bmnh 96.6.29.13; ilha de marajó, bmnh 1923.11.9.20-4; ilha mexicana, zsm 111/1911, 112/1912; "ponto dois indios," bmnh 1939.1.5.5; santarém, bmnh 75.10.22.1-4, mcz 354. _rondonia_: abuná, cas 49773-4, fmnh 64239; forte principe da beira, wcab 10230; igarapé marmelo, fmnh 140254, ku 84725 (skeleton), 92243-6, 92247-8 (skeletons), wcab 9997, 10001, 10003-4; porto velho, mzusp 16343. colombia: _amazonas_: gino-goje, lower río apoporis, usnm 152139; leticia, usnm 152010-1; raudal de la playa, lower río apoporis, mcz 28050; río apoporis, mcz 28060. _boyacá_: sutatenza, usnm 152054-6. _cundinamarca_: medina, mcz 16269-71, usnm 152089-90, 152092-7, 152757. _meta_: el mico, río guejar, usnm 152203; río duda, sierra de macarena, amnh 79914; río guapaya, sierra de macarena, fmnh 81332; río guaviari, casa de piedra, uta no number. _putumayo_: río mecaya, fmnh 69711-4, 69716. _vaupés_: gomogoje, lower río apoporis, mcz 28048. ecuador: no specific locality, wcab 35451, 35785; "oriente," ummz 90418. _carchí_: below salinas, usnm 166059. _esmeraldas_: carondelet, uimnh 53560-9; lagartera, río caoni, uimnh 53441, 53458-79. _morona-santiago_: macuma, uimnh 63142-3, 63145, 63147, 63151, 63154, 63157, usnm 166060. _napo_: avila, ummz 92077; cuyabeno, uimnh 63158, 90111; lago agrio, ku 126647-55; limón cocha, río napo, aum 8132-4, ku 99207-8, 99421-3, 99424 (skeleton), 99425, uimnh 64801, 87798, 87800, 88032-5, 88576, 90066, 90082, 90102, 90104, 90314, 90984; loreto, wcab 35352; río cotapino, ummz 92080; río napo, ummz 84120; san josé abajo, amnh 1295, 1449, 22180, 79990; santa cecilia, aum 8117, 8150, ku 105230-3; south slope volcán sumaco, usnm 166570. _pastaza_: no specific locality, zsm 31/1956; arajuno, usnm 165995; bufeo, lower río bobonaza, usnm 166046-8; canelos, bmnh 80.12.5.179, 1947.2.13.48, ummz 89066; don tomás, río bobonaza, usnm 166049-50; montalvo, cas-su 10320, usnm 165987-9, 166058, 166566; 2.5 km se puyo, usnm 166051; río arajuno, usnm 166043-5; río arajuno (headwaters), usnm 166053; río bobonaza, wcab 3613-4, 35504; río capahuari, usnm 165990, 166555-6; río capahuari (headwaters), usnm 166057; río conambo, usnm 166569, zsm 28/1962, 35/1962; río conambo at río ollaguanga, usnm 166568; río conambo at río shiona-yacu, usnm 166019, 166563-5; río corrientes, usnm 195994, 166020-38, wcab 3841-2; río huiyo-yacu, pico de conambo, usnm 166052; río pastaza, mcz 19697; río pastaza (drainage), nhrm 1966, usnm 165996; río pindo, usnm 166039-41; río pindo at río tigre (village), usnm 165992-3, 166042; río pucayacu, usnm 166054, 166056; río rutuno, usnm 166055; río solis, upper río bobonaza, wcab 39914; río villano, usnm 165991, 166567; sarayacu, bmnh 80.12.5.213, 80.12.5.239-40, mzusp 323; shell mera, ku 99420. _zamora-chinchipe_: "yani-inzari," amnh 43259, 43394; zamora, amnh 78928. french guiana: _cayenne_: crique grégoire, up 40; maripa, oyapok river, up 72; oyapok river, uzm 1473. _inini_: crique gabrielle, up 118-20. guyana: no specific locality: rmnh 1873(3), zmb 3102(2). _east demerara_: atkinson field, asu 11622. _mazaruni-potaro_: chinapora river, upper potaro river, bmnh 1905.11.1.20-1; kamakusa, amnh 21416, 21418-9, 21422; kartabo, amnh 11689, 11691, 11697-9, 11703, 11706-8, 23107, 39730, 70966, 70969-70, 70973-5, usnm 118057; moraballi creek, essequibo river, bmnh 1930.10.10.47-51; oko mountains, fmnh 26692-705; upper potaro river, tung district, bmnh 1905.11.1.40; rockstone, fmnh 26591. _north west_: amakura river, haulover, ummz 83735. _rupununi_: north of acaray river, west of new river, ku 69747-8; kuyuwini landing, amnh 46283; pakaraima mountains, bmnh 1933.6.19.49; shudi-kar-wau, amnh 10665, 39637, 49256(2). _west demerara_: demerara, cas 54773-4; demerara falls, bmnh 72.10.16.16-22, 72.10.16.25-32; dunoon, mcz 4834, ummz 46736, 52493-4, 52502, 52504-5, 57271; vryheid, bmnh 78.12.13.18. perú: _amazonas_: río cenepa, amnh 43400. _huanuco_: monte alegre, río pachitea, amnh 43014, 43019. _loreto_: achinamisa, río huallaga, amnh 42178, 42502; andoas, río pastaza, amnh 79991; cashiboya, amnh 43388, 43453; estirón, río ampiyacu, cas 93264-74, 93276, 93278-9, 93281, 93283-6, 93289, 93311, 93327; igarapé champuia, upper río curiuja, mzusp 10339; iquitos, amnh 42204, 42442, 43468, nhmw 6118; lago de miraño, mouth of río napo, amnh 42712, 43186; nauta, ansp 11399; ollanta, amnh 42865; pampa hermosa, río cushabatay, amnh 43124, 43146; pebas, cas-su 6375; pucallpa, mjp 101(2), 140(3); punga, río tapiche, amnh 43194; "rancho de indiana, iquitos district," mvz 16890; upper río abujao, amnh 42908; río itaya, amnh 42755; upper río pisqui, amnh 43536; río tapiche at río contaya, amnh 42983; río utoquinia at brasilian frontier, amnh 43137; sobral, río tamaya, amnh 43242; yurimaguas, bmnh 84.2.18.50. _san martín_: cainarachi, amnh 42763; moyobamba, zsm 19/1914. surinam: no specific locality, bmnh 70.3.10.67, nhmw 18433.3. _brokopondo_: afobaka, rmnh 16536; brownsweg, rmnh 16537; railway km. 121, rmnh 16534. _marowijne_: djai creek, rmnh 16513-4; maroni river, zmb 8240, 8531; nassaugebergte, rmnh 16517-33; paloemeu, usnm 159025; swamp camp, rmnh 16515. _nickerie_: sipaliwini, rmnh 16538. _saramacca_: left coppename river, rmnh 16535; tibiti, rmnh 16516. _suriname_: berlijn, rmnh 15064; powakka, cm 44226; zanderij, cm 50568. venezuela: _amazonas_: cerro duida, upr-m 2875; cerro marahuaca, upr-m 114-5; esmeralda, amnh 23174; iniridi, smf 2640; la culebra, mcz 28572, upr-m 117; laguna, between tama tama and esmeralda, upr-m 2760; río pescado, amnh 23177; tapara, upr-m 113. _distrito federal_: caracas, bmnh 51.7.17.182. _osteocephalus verrucigerus_ colombia: _huila_: acevedo, río suaza, fmnh 69709-10. ecuador: no specific locality, zmb 16589. _napo_: avila, ummz 90413; south slope cordillera del dué, ku 123176-88, 123189 (skeleton), 124208 (eggs), 124209-11 (tadpoles); l'alegria, usnm 167472-3; río pacayacu, tributary of río cotapino, cas-su 13150; southeast slope volcán sumaco, cas-su 11442. _pastaza_: abitagua, cas-su 5067, fmnh 25791, 27619, ummz 90414, 92092; alpayaca, río pastaza, bmnh 1912.11.1.64; mera, ummz 90412(4). _tungurahua_: baños, nhrm 1960. perú: _ayacucho_: la mar, sivia, río apurimac, fmnh 39853. _huanuco_: río pachitea, midway between puerto victoria and puerto inca, cas-su 17745. _junín_: satipo, mjp 38. literature cited andersson, l. g. 1945. batrachians from east ecuador collected 1937, 1938 by win. clarke-macintyre and rolf blomberg. arkiv zool., 37a(2):1-88. bokermann, w. c. a. 1964. field observations on the hylid frog _osteocephalus taurinus_ fitz. herpetologica, 20:252-255. 1966. lista anotada das localidades tipo de anfíbios brasileiros. são paulo, 183 pp. boulenger, g. a. 1882. catalogue of the batrachia salientia s. ecaudata in the collection of the british museum, ed. 2, london, xvi+503 pp. cochran, d. m. and c. j. goin 1970. frogs of colombia. bull. u.s. natl. mus., 288:xii+655 pp. cope, e. d. 1867. on the families of the raniform anura. jour. acad. nat. sci. philadelphia, 2:189-206. 1874. on some batrachia and nematognathi brought from the upper amazon by prof. orton. proc. acad. nat. sci. philadelphia, 25:120-137. duellman, w. e. 1970a. identity of the south american hylid frog _garbeana garbei_. copeia, (3):534-538. 1970b. the hylid frogs of middle america. monog. mus. nat. hist., univ. kansas, 1:xi+753 pp. duméril, a. m. c. and g. bibron 1841. erpétologie générale ou histoire naturelle compléte des reptiles, vol. 8. paris, 792 pp. fitzinger, l. 1843. systema reptilium. vienna, ix+106 pp. funkhouser, j. 1956. new frogs from ecuador and southwestern colombia. zoologica, 91:73-80. gaige, h. t. 1929. three new tree-frogs from panama and bolivia. occas. papers mus. zool. univ. michigan, 207:1-6. goin, c. j. 1961. synopsis of the genera of hylid frogs. ann. carnegie mus., 36:5-18. melin, d. 1941. contribution to the knowledge of amphibia of south america. göteborgs kungl. vetensk.-och vitterh.-sam. handl., ser. b, 1(4):1-71. peracca, m. g. 1904. viaggio del dr. enrico festa nell' ecuador e regioni vicine. reptile ed amfibii. boll. mus. zool. anat. comp., univ. torino, 19:1-41. steindachner, f. 1862. über zwei noch unbeschriebene batrachier. arch. zool. anat. fisiol., 2:77-82. 1867. amphibien. novara expedition. zool. theil, i, vienna, 70 pp. trueb, l. 1970a. the evolutionary relationships of casque-headed treefrogs with co-ossified skulls (family hylidae). univ. kansas publ. mus. nat. hist., 18:547-716. 1970b. the generic status of _hyla siemersi_ mertens. herpetologica, 26:254-267. trueb, l. and w. e. duellman 1970. the systematic status and life history of _hyla verrucigera_ werner. copeia (4):601-610. tyler, m. 1971. the phylogenetic significance of vocal sac structure in hylid frogs. univ. kansas publ. mus. nat. hist., 19:319-360. werner, f. 1901. ueber reptilien und batrachier aus ecuador und neu-guinea. verh. zool.-bot. gesell. wien, 50:593-614. university of kansas publications museum of natural history the university of kansas publications, museum of natural history, beginning with volume 1 in 1946, was discontinued with volume 20 in 1971. shorter research papers formerly published in the above series are now published as occasional papers, museum of natural history. the miscellaneous publications, museum of natural history, began with number 1 in 1946. longer research papers are published in that series. monographs of the museum of natural history were initiated in 1970. institutional libraries interested in exchanging publications may obtain the occasional papers and miscellaneous publications by addressing the exchange librarian, university of kansas library, lawrence, kansas 66044. individuals may purchase separate numbers of all series. prices may be obtained upon request addressed to publications secretary, museum of natural history, university of kansas, lawrence, kansas 66044. transcriber's notes except for the list of corrections below and minor corrections not listed, the text presented here is that of the original printed version. typographical corrections page correction ==== ================ 5 is => in 5 buckley => buckleyi 17 scaral => sacral 19 provicia => provincia 25 in => is 25 metalic => metallic 41 bromeiad => bromeliad text emphasis _text_ italics =text= bold billy bunny and uncle bull frog by david cory author of "billy bunny and daddy fox," "billy bunny and the friendly elephant," "billy bunny and uncle lucky lefthindfoot" illustrations by hugh spencer billy bunny books by david gory large 12 mo. illustrated 1. billy bunny and the friendly elephant 2. billy bunny and daddy fox 3. billy bunny and uncle bull frog 4. billy bunny and uncle lucky lefthindfoot other volumes in preparation 1920 contents i. billy bunny and mr. blacksnake ii. billy bunny and the freshwater crab iii. billy bunny and the sorrowful jay bird iv. billy bunny and the ting-a-ling telephone v. billy bunny and the runaway dog vi. billy bunny and mr. o'hare's escape vii. billy bunny and the policeman cat viii. billy bunny and the gray mouse ix. billy bunny and red rooster x. billy bunny and mrs. cow xi. billy bunny and the big bear xii. billy bunny and the rabbitville "gazette" xiii. billy bunny and mr. mole xiv. billy bunny and the water snake xv. billy bunny and the peacock xvi. billy bunny and the marble deer xvii. billy bunny and the forest dance xviii. billy bunny and ragged rabbit xix. billy bunny and tailor bird xx. billy bunny and parson crow xxi. billy bunny and jack-in-the-box xxii. billy bunny and mr. duck xxiii. billy bunny and the fretful porcupine xxiv. billy bunny and danny billygoat xxv. billy bunny and the whale xxvi. billy bunny and the mermaid. xxvii. billy bunny and the beanstalk xxviii. billy bunny and scatterbrains xxix. billy bunny and mrs. black cat xxx. billy bunny and big yellow dog xxxi. billy bunny and a happy birthday xxxii. billy bunny and the lost ring xxxiii. billy bunny and the great news xxxiv. billy bunny and jenny muskrat xxxv. billy bunny and the miller's dog xxxvi. billy bunny and the woodchuck xxxvii. billy bunny and little peewee xxxviii. billy bunny and old mother magpie story i. billy bunny and mr. blacksnake. rain, rain, go away, billy bunny wants to play. this is what willy wind sang one morning. oh, so early, as the raindrops pitter-pattered on the roof of the little rabbit's house in the old brier patch. and then of course he woke up and wiggled his little pink nose a million times less or more, and pretty soon he was wide awake, so he got up and looked into the mirror to see if his eyes were open, as he wasn't quite sure he was wide awake after all, for the raindrops made a drowsy noise on the old shingles and the alarm clock wouldn't go off, although it was 14 o'clock. well, after a little while, not so very long, his mother called to him, "billy bunny, the stewed lollypops are getting cold and the robin's eggs will be hard boiled if you don't hurry up, or hurry down, or something." "i'll be ready in a jiffy," answered the little rabbit, and then he brushed his whiskers and parted his hair in the middle with a little chip, and after that he was ready for breakfast and dinner and supper, for rabbits are always hungry, you know, and can eat all the time, so i've been told, and i guess it must be true, for why should an old rabbit have told me that if it isn't the truth, i should like to know, and so would you, i'm sure. "don't forget your rubber boots," said mrs. bunny after the morning meal was over, as billy bunny started to hop outdoors. so, like a good little bunny boy, he came back and put them on, and then before he went he polished the brass door knob on the front door and swept the leaves off the little stone walk. and after that he was ready to do whatever he liked, so out he went on the pleasant meadow to eat some clover tops so as not to feel hungry for the next ten minutes. and just then mrs. cow came along with her tinkle, tinkle bell that hung at her throat from a leather collar. "where are you going?" she asked, but the little rabbit didn't know. he was only looking around. he hadn't had time to make up his mind what to do, and just then, all of a sudden, just like that, mr. blacksnake rose out of the grass. "look out!" cried mrs. cow. "maybe he's going to eat you," but whether he was i'm sure i don't know, for billy bunny didn't wait to see. he didn't care whether mr. blacksnake wanted his breakfast, but hopped away as fast as he could and pretty soon, not so very far, he came to the babbling brook, and there sat the little fresh water crab on the sand, and when he saw billy bunny he said: "it's raining, billy bunny, but you and i don't care, for raindrops make the flowers grow and blossom fair." and this is what every little boy and girl should say on rainy days. story ii. billy bunny and the freshwater crab. let me see. it was raining in the last story when we left off, wasn't it? billy bunny and the little freshwater crab were talking together, weren't they? that's it, and now i know where to begin, for it's stopped raining since then and mr. happy sun is shining in the sky and the little clouds are chasing each other over the blue meadows like little lambs. "i like that little piece of poetry you just said," cried the little rabbit. "please say another." so the freshwater crab wrinkled his forehead, and then he began: "and when the sun is shining, and all is bright and gay, just keep a little sunshine to help a rainy day." "i will," said the little bunny, for he was a cheerful little fellow, and then he hopped away and by and by he came to the old mill pond. but uncle bullfrog was nowhere to be seen. there stood the old log, but there was nobody on it but a black snail. it seemed strange not to see the old gentleman frog sitting there, his eyes winking and blinking and his white waist-coat shining in the sun, and it made the little rabbit feel lonely. "where is uncle bullfrog?" he asked a big bluebottle fly, who was buzzing away at a great rate. but he didn't know, and neither did a big darning needle that was skimming over the quiet water. "i wonder if that dreadful miller's boy has taken uncle bullfrog away," thought billy bunny, and just then mrs. oriole flew down from her nest that swung in the weeping willow tree and said: "are you looking for uncle bullfrog, little rabbit?" "yes, ma'am. do you know where he is?" "he's down by the mill dam," answered the pretty little bird, and then she flew back to her nest that looked like an old white cotton stocking at christmas time because it was all bulgy and full, only, of course, hers had little birds inside and a christmas stocking has all sorts of toys, with an orange in the toe and a jack-in-the-box sticking out of the top. so off hopped the little rabbit, and pretty soon he saw the old gentleman bullfrog catching flies, and undoing his waistcoat one button every time a fly disappeared down his throat. "i thought at first that dreadful miller's boy had taken you away," said billy bunny, "and i was very sad, for i like you, uncle bullfrog, and i've never forgotten how you found the letter i lost a long time ago." "tut, tut," said the old gentleman frog. "how's your mother?" and then he swallowed another fly and unbuttoned the last button, and if he takes off his waistcoat i'll tell you so in the next story. story iii. billy bunny and the sorrowful jay bird. well, uncle bullfrog didn't take off his waistcoat, as i thought he might in the last story, so i'm not going to tell you anything more about him. we'll just leave him in the old mill pond and go along with billy bunny, who is hopping away toward the friendly forest. by and by, after he had gone into the shady depths for maybe a million and two or three hops, he came across his old friend the jay bird, who had sold him the airship, you remember, and then bought it back again. "i wish you'd kept your old flying machine," said the jay bird sorrowfully. "but you wanted to buy it back," said the little rabbit, "so it's not my fault." "perhaps not," replied the sorrowful jay bird, "but that doesn't make matters any better." "why, what's the trouble?" asked the little rabbit, sitting down and taking a lollypop out of his knapsack. "i had an accident," answered the jay bird. "i ran into a thunder cloud and spilled out all the lightning, and, oh dear, oh dear. i just hate to talk about it, but i will. the lightning jumped all around and then struck the old tower clock and broke the main spring, so that it wouldn't go any more, and now nobody in rabbitville can tell the day of the month, or when it will be thanksgiving or fourth of july." "let's go to the clock maker and ask him to fix it," suggested the little rabbit, and this so delighted the sorrowful jay bird that he smiled and flew after billy bunny, and pretty soon they came to the old clock maker, who was an old black spider. "certainly i'll fix it," he said, "but it will cost you nine million and some billion flies." "all right," said billy bunny. "i'll go down to the 3 and 1-cent store and buy a fly catcher." so off he went and pretty soon he came back with a great big fly catching box, and after he had set it down, they stood and watched the flies go in until it was so full that not another one could even poke in his nose. "now, mr. spider," said billy bunny, "there are maybe a trillion flies in that box, for the storekeeper told me it was guaranteed to hold that many, so please fix the town clock, for it would be too bad if the little boys and girls didn't know it was christmas when it really came." so the spider got out his little tool bag and climbed up the steeple and fixed that old town clock so well that it began to play a tune, which it had never done before, and all the people in rabbitville were so delighted that they gave the spider a little house to live in for the rest of his days. story iv. billy bunny and the ting-a-ling telephone. ting-a-ling went the telephone bell in uncle lucky lefthindfoot's house, the kind old gentleman rabbit who was the uncle of billy bunny, you know. and i only say this right here in case some little boy or girl should read this story without having seen all the million and one, or two, or three that have gone before. so uncle lucky jumped out of the hammock where he had been swinging up and down on the cool front porch of his little house in bunnytown, corner of lettuce avenue and carrot street, and hopped into the library and took down the receiver and said "helloa! this is mr. lucky lefthindfoot talking." "is that you, uncle lucky?" answered a voice at the other end of the wire. "this is billy bunny, and i'm lost in the friendly forest." "what!" cried the old gentleman rabbit, and he got so excited that he put the wrong end of the receiver to his left ear and got an awful electric shock that nearly wiggled his ear off. "where are you now?" "i don't know," replied his small nephew. "i'm lost, don't you understand?" "gracious, goodness mebus!" exclaimed the old gentleman rabbit, "then how am i to find you?" "i don't know, but please do," said billy bunny sorrowfully, "for i'm dreadfully hungry, and i haven't got a single lollypop or apple pie left in my knapsack." "well, you just stay where you are and i'll get into the luckmobile and find you," replied the old gentleman rabbit as cheerfully as he could, although he didn't know how he was going to do it, and neither do i, and neither do you, but let's wait and see. so pretty soon, in a few short seconds, uncle lucky was tearing along the dusty road toward the friendly forest, and by and by he came to the house where his cousin, mr. o'hare, lived. so he stopped the automobile and knocked on the door, and as soon as mr. o'hare opened it, he said: "jump in with me, for my little nephew is lost and i want you to help me find him." so away they went into the friendly forest, and they looked all around, but, of course, there was no little rabbit that looked like billy bunny anywhere in sight. so uncle lucky and mr. o'hare got out, and after tying the automobile to a tree, they set out in different directions to find the little bunny. and uncle lucky went along a little path and mr. o'hare followed a small brook, and after a while the old gentleman rabbit heard a bird singing: "i saw a little rabbit a-sitting by a tree, and i should say he'd lost his way- that's how he looked to me." "where did you see him?" asked uncle lucky excitedly. but what the little bird replied you must wait to hear in the next story. story v. billy bunny and the runaway dog. you remember in the last story just as uncle lucky asked the little bird to tell him where billy bunny was i had to leave off for there was no more room in the story for me to add another word? well, what the little bird said was: "follow the path, mr. lucky lefthindfoot, 'till you come to a bridge, and then turn to your right, and pretty soon, if the little bunny hasn't hopped away, you'll find your lost nephew." so uncle lucky started right off. he didn't wait to even dust off his old wedding stovepipe hat, and by and by he came to the bridge. but oh dear me! right in the middle of it stood a big dog, and when he saw the old gentleman rabbit he gave a loud bark and ran at him. and what do you think the dear old bunny did? he honked on his automobile horn, which he had in his paw, and this frightened the dog so dreadfully that he turned around and ran away so fast that he would have left his tail a thousand miles behind him if it hadn't been tied on the way dogs' tails are, you know. and after that uncle lucky crossed the bridge and turned to his right and pretty soon he saw billy bunny under a bush looking very miserable and unhappy. but when he heard his uncle lucky's voice, for the old gentleman rabbit gave a cry of delight as soon as he saw him, the little rabbit looked as happy as he had before he was lost. "here's an apple pie for you," said the dear, kind old gentleman rabbit, taking a lovely pie out of his pocket. "i knew you'd rather have something to eat than a million carrot cents." and of course the little rabbit would, for he was so hungry he could have eaten brass tacks, or maybe iron nails. "now come along with me," said uncle lucky. "we'll go back to the luckymobile. your cousin, mr. o'hare, went the other way to look for you, so i suppose we'll have a dreadful time to find him. but, never mind, i've found you." and dear, affectionate uncle lucky hugged his small nephew, he was so glad to be with him once more. well, after they reached the automobile they honked and honked on the horn hoping mr. o'hare would hear them. but i guess he didn't, for he never came back, although they waited until it was almost 13 o'clock. "we'll have to go home without him," said uncle lucky at last. and i guess he was wise not to wait any longer, for it was growing dark, and to drive an automobile through a forest is not an easy thing to do at night. and just then, all of a sudden, willie wind came blowing through the tree tops. when he saw the two little bunnies he said: "your cousin, mr. o'hare, has fallen into a deep hole over yonder." and willie wind pointed down the friendly forest trail. in the next story you shall hear how uncle lucky and billy bunny found their cousin, mr. o'hare. story vi. billy bunny and mr. o'hare's escape. you remember in the last story how willie wind whispered to billy bunny and uncle lucky that their cousin, mr. o'hare, had fallen into a deep hole? well, it didn't take the two little rabbits more than five short seconds and maybe five and a half hops to reach the spot, and then they looked over the edge, but very carefully, you know, for fear they might fall in, and there, sure enough, way down at the bottom was mr. o'hare looking very miserable indeed. "keep up your courage!" cried uncle lucky in as cheerful a voice as he could muster, and then he looked around to find a rope or a ladder. but of course there were not any ropes and ladders lying about, so that kind old gentleman rabbit peeped over the edge of the hole and called down again, "keep up your courage! we'll get you out!" although he didn't know how he was going to do it, and neither do you and neither do i and neither does the printer man. well, after a while, and it was quite a long while, too, billy bunny found a wild grapevine which he let down into the hole. "make a loop and put it around your waist and uncle lucky and i will haul you out," he called down, and then mr. o'hare did as he was told, and after the two little rabbits had pulled and pulled until their breath was almost gone, mr. o'hare's head appeared at the top of the hole. and then with one more big pull they brought him out safely, although his waist was dreadfully sore because the grapevine had cut into his fur and squeezed all the breath out of him. "i'm going to complain to the street cleaning department or the first policeman i see," said mr. o'hare. "it's a dreadful thing to have a hole like this right in the middle of the friendly forest trail." "never mind that," said billy bunny, "let's go back to the luckymobile. it will be late before we get out of the woods and maybe the electricity will all be gone and then we can't light the lamps, and maybe we'll be arrested." and this is just what happened. they had only gone a little ways when they heard a voice say: "stop your motor car, i say, you have no lamps to light the way. come, stop your car and get right out! listen, don't you hear me shout? stop your car or i will shoot. don't try away from me to scoot!" "we don't intend to," said uncle lucky, and he put on the brake and the luckymobile came to a standstill. and there in the road stood a big policeman cat, with a club and gold buttons on his coat and a big helmet, and his number was two dozen and a half. "get out of your car," he commanded, which means to say something sternly, but before the two little rabbits obeyed, something happened, but what it was you must wait to hear in the next story. story vii. billy bunny and the policeman cat. well, i'm glad to say it was something nice that happened just as i left off in the last story. you remember the policeman cat had arrested billy bunny and his uncle lucky. well, just as that policeman cat lifted his club to tickle uncle lucky's left hind foot, a big elm tree began to bark and of course the policeman cat was nearly scared to death. he thought it was a dog, you see, and instead of tickling dear, kind uncle lucky with his club, he turned tail and ran off down the road. and he ran so fast that he left his number behind and uncle lucky picked it up and put it on the automobile, and after that they asked two little fireflies to sit inside the lamps and make them shine, for you remember the electricity had all burned up. well, after a while, they came to a turn in the road and, goodness gracious! before they could stop the automobile they ran into a milk wagon. and, oh, dear me! there was whipped cream all over the place, and billy bunny and uncle lucky looked like two little cream puffs. and i suppose you are wondering where the driver of the milk wagon was all this time. and so were uncle lucky and billy bunny, and if you'll wait a minute i'll tell you, as soon as my typewriter behaves itself, for it got so excited when luckymobile ran into the milk wagon that it caught my thumb and pinched it. well, pretty soon, after uncle lucky had looked behind the moon and billy bunny into all the empty milk cans and one full one, they found the driver up in a weeping willow tree. "i'll come down if you'll promise not to run over me," he said, for he was nearly frightened to death and looked dreadfully funny, for one of the milk can covers had fallen on his head. "i thought he would be mad as a hornet," whispered billy bunny to his rabbit uncle. "but where's my horse?" said the milkman when he reached the ground. so they all looked around and everywhere else, but they couldn't find him until they looked up into another weeping willow tree. and there was the poor horse high up in the branches. "oh, i'll come down from this willow tree, if you'll promise me just one thing, and that is never again to say: 'gid-ap' as you drive me along the way, for i always go the best i can; i'm a faithful friend to every man, so please don't hurry me so, for i'm not trying to go too slow." "all right, my good old horse," said kind uncle lucky. "your master shall give me his word." so the horse jumped down and the willow tree stopped weeping right away, for it was so glad that the poor old milk horse was never again to be hurried on his way. and in the next story i'll tell you why. story viii. billy bunny and the gray mouse. you remember in the last story how the luckymobile had run into a milk wagon? well, after billy bunny had helped the milkman hitch up his horse and uncle lucky had filled the milk cans with ice cream and soda water from a near-by candy store, so as not to have all the little boys and girls disappointed at breakfast when they didn't get their milk, our two little rabbit friends got into the luckymobile and started off again. well, it was still evening, you know, and the little fireflies who had crawled into the lamps made them as bright as possible, so it wasn't hard to steer the automobile. and, after a while, maybe a mile, they came to a house, where lived a gray mouse, all alone by herself in a hole near a shelf, where cake and mince pies made her open her eyes, for they looked, oh, so good, as a pie or cake should. now i didn't know i was going to write poetry or i should have let my hair grow long like a poet instead of going to the barber for a shave. well, anyway, the two little rabbits stopped the automobile right in front of mousie's door and when she heard the horn go honk, honk, she came to the window and looked out. "why, it's mr. lucky lefthindfoot," she squeaked, and then she opened the door and asked the two little rabbits in and gave them some pie and cake. "you can put the automobile in the barn if you like," she said, "and spend the night here, for it's getting very dark and maybe you'll run into something." so billy bunny took the luckymobile around to the barn, and just then an old owl began to toot: "i'm very fond of little gray mice, and little white rabbits, too, are nice." and down flew that old gray owl and made a grab for billy bunny. but he didn't catch him. no, sireemam! for the little rabbit hopped into the henhouse through the little round door, and the big red rooster began to crow: "look here, mr. owl, if you come inside i'll hurt you with my spur. don't you dare get funny with billy bunny, or muss his pretty white fur." and then he flew down from his perch and said, "cock-a-doodle-do" three times and a half, and after that the owl flew away. "that was very kind of you," said the little rabbit. "oh, don't mention it," said the red rooster, "but there is one thing you can do for me." "what's that?" asked billy bunny. "take me luckymobiling," laughed the red rooster. "all right. to-morrow uncle lucky and i will invite you for a nice drive," said the little rabbit, and if the luckymobile doesn't get sick maybe uncle lucky will ask some little boy or girl to go, too, and maybe it might be you. story ix. billy bunny and red rooster. well, the next morning when the little rabbits woke up the sun was shining brightly through their bedroom window and mrs. mousie was singing a song down in the kitchen below as she made hot muffins for breakfast. and this is what she sang: "upstairs in my nice guest room are two nice little rabbits in bed. as soon as i'm able i'll fix up the table and give them some honey and bread. and then a hot muffin to give them a stuffin', and then they'll be bountifully fed." and when billy bunny heard her he grew so hungry that he hurried faster than he had ever hurried before, and so did the old gentleman rabbit, and he buttoned his collar on backwards and put his left shoe on his right foot and tripped over his old wedding stovepipe hat. and after that they both hopped downstairs, and as soon as mrs. mousie heard them she brought in the bread and honey and the hot muffins and they all had breakfast. and after that billy bunny asked her to go automobiling with them. so she put on her old gray bonnet with a bit of ribbon on it, and tied the strings under her chin, and put on her black silk mitts and her gold locket breastpin with the picture of mr. mousie inside. "you don't mind if we invite the red rooster to go along, too, do you?" asked billy bunny, and then he told her how the rooster had scared away the old owl. and of course mrs. mousie didn't care, so the rooster got in and sat on the back seat with mrs. mousie. well, after they had gone for maybe a mile, and maybe some more, they came to a beautiful candy store, where the windows were full of peppermint sticks and a brown sugar monkey did all sorts of tricks. "stop right here," said the red rooster, "and i'll get out and buy you a bag of candy." and when he came back he had four bags of candy. just think of that! in one bag was sugar-coated carrots for billy bunny, and another bag was full of candied carrots for uncle lucky, and in the bag he gave to mrs. mousie were two little chocolate mice. "what have you got in your bag?" asked uncle lucky as he made the luckymobile jump over a high ditch and run along through a lovely green meadow spread all over with buttercups. "sugared peanuts," answered the red rooster. "i just love them. the last time i went to the circus i ate forty-nine bags and a half and drank twenty-three glasses of pink lemonade and a bushel of popcorn." "wait a minute," said the old gentleman rabbit. "i've got a stomach ache listening. how did you do it?" and in the next story i'll tell you what the rooster said, that is, if nothing happens to prevent it, for he certainly was a wonderful rooster, to be able to eat all that. story x. billy bunny and mrs. cow. well, something did happen to prevent the red rooster from telling billy bunny how he had been able to eat forty-nine bags and a half of peanuts at the circus, as i mentioned in the last story. you see, as the luckymobile galloped along over the meadow, all of a sudden, just like that, it ran right into the babbling brook, and then of course it stopped so suddenly that billy bunny and uncle lucky didn't stop at all, neither did mrs. mousie and the red rooster. they just kept right on going, and the first thing they knew and the first thing you know, they all landed in the long grass beside mrs. cow. "my, how you startled me!" she exclaimed, and she rang the little bell at her neck and up ran her little calf, who was only two weeks old, and had never seen billy bunny and his friends before. after that she walked down to the babbling brook--but oh, dear me! all the electricity oil had spilled out of the cabaret and she couldn't drink the water, and all the little fish were covered with it just like sardines, you know, and the watercress had salad dressing all over it, so of course she couldn't eat the watercress. "never mind," said kind little billy bunny, and he took out of his knapsack a big yellow lemon lollypop and gave it to her, and then she didn't care, for she just loved candy. "i'll help you get the automobile out," said mrs. cow gratefully, for she liked anybody who was kind to her little calf. so she put her horns under the front of the luckymobile and then she said, "heave ho, e-ho!" and pushed and shoved and lifted that big heavy automobile right out of the brook without even cracking her two long horns. "if you don't mind," said the red rooster, "i'll leave you two little rabbits and make a call on cocky docky up at the old farm." "and if you don't care," squeaked little mrs. mousie, "i'll call on dickey meadowmouse." so uncle lucky and billy bunny hopped into the automobile and drove off, while mrs. cow tinkled her bell and sang: "moo, moo, moo. i'm glad i helped you two. one good turn deserves another. when you see your bunny mother, tell her how your car i took safely from the babbling brook." "it's a puzzle to me," said uncle lucky, "why we are always having so many accidents. maybe i had better get a chauffeur." "you won't need any chauffeur after i'm done with you," said a deep growly voice, and out from behind a clump of bushes jumped a wicked wildcat and bit one of the front tires, she was so hungry. and what do you suppose happened then? why the tire burst with such a loud noise, just like a gun, you know, that the wildcat was frightened nearly to death and she turned around and ran away so fast that she got home an hour too early for supper. story xi. bill bunny and the big bear. near the friendly forest pool is the woodland singing school. little squirrel bushy tail sings the do, ray, mee, fa scale. uncle bullfrog sings "ker-chunk" from his floating elm tree trunk. and a big good-natured bear sings an old familiar air. "it's time for your singing lesson," said mrs. bunny to her little rabbit. so billy bunny started off, hoppity hop, down the friendly forest trail, and by and by he reached the pool where all the pupils came to take their singing lessons. mr. grasshopper was there with his fiddle and the tree toad with his drum, and the lark with her flute and little jenny wren with her piano. and what do you suppose billy bunny had tucked away in his knapsack? why, uncle lucky's automobile horn. you see, the kind old gentleman rabbit was making a visit at the old brier patch where he had taken his automobile after that dreadful wildcat had bitten the front tire, and this is how billy bunny came to get the horn. well, sir, after the music started, he pulled out his horn and gave a tre-men-dous honk on it, and everybody thought an automobile was going to run over him. some jumped into the pool and some ran up the trees, and, oh, dear me! everybody got all out of tune, and the bear lost the air and couldn't find it again! and just then who should come along but a peddler with a pack of tin cans, rattling away on his back, and of course he made more noise than all the singing school put together. and when the big bear saw him he was so angry that he jumped from behind a tree and said, "boo!" "do you want to buy a tin plate?" asked the peddler, trying hard not to be frightened, "or would rather have a dishpan?" "don't want either," said the bear with a terrible growl. "perhaps you'd like a nutmeg grater," said the poor old peddler, and he was so frightened by this time that his knees knocked into the tin pans and made a dreadful noise. "i've a dandy egg beater," went on the peddler, in a trembling voice, but after that he never said another word, for that great big bear jumped right at him and took the egg beater out of his hands and growled so terribly that the tin peddler turned away and ran down the forest path as fast as he could go. and then all the little and big forest folk began to sing: "hip, hip hurray, the peddler's gone away. no more he'll make his tin pans shake and spoil our singing school beside the forest pool." and in the next story, if the baby who lives in the house opposite doesn't shake his rattle at me all night so that i can't get to sleep and dream about the next story in time to write it for to-morrow night, i'll tell you more about the little rabbit's adventures. story xii. billy bunny and the rabbitville "gazette." there was once a little rabbit who was very fond of pie, apple pie, with sugar on the crust. and he had a little habit, when his mother wasn't nigh, of eating apple pie until he bust. this is what mr. william bunny, the little rabbit's father, you know, was singing one day, and the reason was because mrs. bunny had found little billy bunny in the pantry. and what happened to the little rabbit i'm not going to tell you, for it is so sad that it would make you weep to hear it. "all day he nibbled pie till at last i thought he'd die," said the doctor with a sigh. and then mr. william bunny looked at his small son and sighed, too, for he had just paid the doctor's bill. "please don't sing any more," said little billy bunny. "don't you remember the doctor said i was to be kept quiet?" so mr. william bunny went out on the porch to smoke a cigar and read the rabbitville "gazette" until after supper time. and while he was reading mrs. bunny looked over his shoulder and read: "wanted, a secondhand automobile in good condition." "ring up your uncle lucky on the telephone," she called to billy bunny. "here's a chance for him to sell his luckymobile." so the little rabbit rang up 000 lettuceville, and in a few minutes he heard the old gentleman's voice at the other end of the wire. "but i don't want to sell my luckymobile," he said. "it's the only one in ex-is-tence," which means the only one ever made, and i guess he was right, for i never rode in a luckymobile, did you? "but mother thinks you ought to sell it," said billy bunny, "and so does father, for they both say you'll have a terrible accident some day if you don't look out." "well then, i'll look out," said uncle lucky with a laugh. "but i won't sell my luckymobile." and then he asked billy bunny to make him a visit. so the little rabbit put on his knapsack and picked up his striped candy cane and started off, after first asking his mother's permission, of course. and after he had gone for maybe a million hops, he came to a big tree where old barney the owl had his next. but of course, he wasn't awake. oh, my, no. he had his eyes tightly closed, for owls don't like a bright light, you know. they can see in the dark but not in the daytime. but when billy bunny called out, "helloa, mr. barney," the old gentleman owl blinked his eyes and said, "who's calling me?" and then the little rabbit thought he'd play a joke, so he said, "mr. mouse!" and if there was anything that old barney loved to eat, it was mice. and in the next story i'll tell you what billy bunny did. story xiii. billy bunny and mr. mole. you remember in the last story i promised to tell you what billy bunny did when old barney the owl asked him, "who's there?" and the little rabbit replied, "mr. mouse," just to fool him, you know. well, after that old barney the owl gave a terrible scowl as he looked at little bill bunny. you thought you were wise, but my blinky old eyes can see you are not a bit funny. i can see from my house you are not mr. mouse. and then the old blinkerty, winkerty owl flopped down to the ground and tried to catch the little rabbit. but billy bunny was too quick for him. he jumped into a hollow stump before you could say "jack rabbit!" "come out of there," cried old barney, in a screechery, teachery voice, but you just bet the little bunny didn't. he knew what would happen if he did. well, by and by, after a long while, he looked around, and, would you believe it, he found a little pair of stairs. so down he hopped until he came to a door on which was painted in red letters: "mr. mole, subway contractor." then the little rabbit knocked on the door and pretty soon it was opened and there stood mr. mole himself. "what do you want?" he asked, trying to squint out of his little tiny eyes that were hidden all over with hair. "it's me--billy bunny," replied the little rabbit. "mr. owl tried to catch me and i hopped into your hollow stump entrance, but i haven't got a ticket for the subway." "well, you can come in anyway," said the kind old mole; "my subway isn't finished yet and the trains won't be running for some time. come in." so billy bunny hopped inside and sat down on a chair close to a little brass railing, behind which stood mr. mole's desk. then mr. mole sat down and looked at billy bunny as much as to say, "and now what can i do for you?" so billy bunny said, "i would like to get up on the ground again. can you show me a new way, because i don't want to go back the way i came?" then mr. mole pressed a little bell, and in came a mole with overalls on and a little pickaxe. "show my friend, mr. billy bunny, through the tunnel to the moss bank entrance." "thank you," said the little rabbit, and he hopped after the workman mole until they came to an opening. and when the little rabbit got outside once more he found himself on a mossy bank where blossomed a lovely bed of violets. so he picked a bouquet for himself and stuck it in his buttonhole, and after that he hopped away singing a song. and if robbie redbreast hadn't heard it i never would have been able to tell it to you. wasn't it lucky that the little robin sang it to me this morning while i was still in bed? because, if he hadn't, how would i have ever learned it? over the clover and over the grass hoppity, hop, i go; over the leaves from the autumn trees and over the soft white snow, with a whistle and song i go hopping along, i'm billy bunny, you know. story xiv. billy bunny and the water snake. "over the grass or over the snow, fast as a little white breeze i go. i'm billy bunny, billy bunny, you know." thus sang the little rabbit even after i left off in last night's story. isn't it strange? maybe i dreamed it. anyhow, that's what i think he did, and after a while, when he had stopped singing, you know, he came to a little hill on the top of which was a high white pole with an american flag flying from it. and underneath was a whole regiment of little boy bunny scouts, dressed in khaki, with guns and caps and brass buttons and guns and drums and a captain and a fife, and i guess there were three or four fifes, and as soon as they saw the little rabbit, they all shouted, "here comes billy bunny. let's get him to join our regiment." "i belong to the billy bunny boy scouts of old snake fence corner," replied the little rabbit. "i can't join your regiment." so he hopped along and by and by he came to a big white swan that was sailing up and down on a pond. "would you like to take a sail?" she asked, coming up close to the bank. "because if you would, just hop on my back and i'll take you around the pond two times and maybe a half if you'll give me a lollypop." so the little rabbit opened his knapsack and gave her one and then he hopped on her back and went for a lovely sail in and out among the pond lilies and little green grass islands. well, everything was going along beautifully when, all of a sudden, just like that, a big water snake came swimming by. "oh, don't let him swallow me," cried the little rabbit, and he took his popgun out of his knapsack and stuck the cork in the end. "i'll shoot you on the tail if you touch me," he cried just as bravely as he could, but he nearly slipped off the swan's back just the same, he was so frightened. "don't you come any nearer," said the swan with a fierce hiss, but the snake didn't care. he swam around and around until the little rabbit got so dizzy that he had to hold on to the swan's neck. "please swim around the other way," pleaded the little rabbit, "you make me dreadfully dizzy." but the bad water snake said he wouldn't, because that's just what he wanted billy bunny to be--so dizzy that he would fall into the water and then that dreadful water snake could swallow him and maybe a pond lily besides. "look here," said the swan, "if you don't stop making snakery circles all around me, i'll bite your head off with my big, strong beak." and then what do you think the little rabbit did? why, he managed somehow to lift up his gun and shoot it off, and the cork hit the water snake on the end of the tail and gave him such a headache that he swam over to the long grass and ate watercress salad and a piece of lemon pie. and while he was doing that the swan took the little rabbit to the other side of the pond and he hopped away so fast that he didn't tell me what he was going to do in to-morrow's story. story xv. billy bunny and the peacock. well, if it hadn't been for robbie redbreast who saw little billy bunny hopping away from the lily pond, as i told you in the last story, i never would have found out what he did after that, and so there would have been no story to-night. so the next time you see robbie redbreast, please thank him. and now this is what he told me. after the little rabbit had hopped along for maybe a mile or three, he came to a high stone wall. "i wonder what's on the other side?" he said to himself, and then a beautiful peacock looked over and said: "i'll tell you, little rabbit. "it's a beautiful garden where a fountain plays all day and the breezes sing all night and the flowers whisper and bow their heads." "how can i get in?" asked the little bunny, "for i love flowers and i never heard a fountain play. what does it play?" "oh, all sorts of waterfall music," said the peacock, and he spread his beautiful tail out like a fan and brushed a little green fly off his nose. "it plays trills and rills and cascades and ripples and dipples." and this made the little rabbit so curious that he hunted all around to find a gate in the high stone wall. and pretty soon, not so very long, he came to one, with big iron rods and curiously carved images of lions and dragons and animals with wings. so he squeezed through and hopped up to the beautiful fountain where lots of little gold and silver fish swam around and around and the water fell in diamonds and rubies and emeralds, but he didn't know that it was mr. happy sun who colored the water drops to make them look like precious stones. "please play me a tune," said the little rabbit. and then the beautiful peacock said, "what tune would you like?" and the little rabbit answered: "sprinkle, sprinkle, little star, just a water drop you are. twinkle, twinkle, drops of dew, with the sunlight shining through." so the beautiful fountain played this little song while billy bunny sat there listening and the beautiful peacock spread his tail to catch the sparkle from the glittering drops of water. and then all the roses began singing: roses white and roses red, and roses yellow too, instead, and pretty lilies white as snow, and every other flower you know. and after that billy bunny asked the peacock to sing a song, but when he started to sing, oh dear, oh dear. for you know just because a bird has beautiful feathers he may not have a beautiful voice, and the sounds the peacock made were dreadful. yes, indeed. and if the little rabbit hadn't skipped away he would have had to hold his paws over his ears, and then maybe he couldn't have stopped them up, for he had very large ears and very small feet. story xvi. billy bunny and the marble deer. in the story before this i told you how the beautiful peacock sang a song which was dreadful, so very dreadful that little billy bunny had to hold his ears and run away from the lovely fountain. well, after he had hopped along for maybe a million hops or less, he came to a little deer on a smooth lawn. so he stopped and spoke to him, but the pretty little animal never said a word. he didn't even look at the little rabbit, so billy bunny touched him on the nose, but, oh, dear me! it was cold and hard, not at all like the nose of a real little deer. but the little bunny didn't know it was a marble deer. he just thought it was alive, you see, and he was puzzled and didn't know what to do and then a lovely white dove flew down and said: "he can't speak. he's only a statue." "what is that?" asked the little rabbit, for he had never seen one before. "why, a statue is a figure carved out of marble or stone," answered the dove, and then she began to coo and comb her feathers with her bill. "well, i'll just hop along then," said billy bunny, and he said good-by. and after a while he came to a little house all covered with red rambler roses, so he looked inside to see who lived there, for he thought perhaps it might be a fairy who owned this beautiful garden with the lovely fountain and the wonderful peacock. but there was no one inside, so he hopped in and sat down on a small wicker chair and rocked back and forth. for it was a rocking chair, you know. and, by and by, he fell asleep and dreamed that the beautiful peacock was flying around the fountain and scattering the water drops all about with his mag-nif-i-cent tail. and then, all of a sudden, the little rabbit woke up, for somebody was saying: "isn't this a dear little bunny?" and billy bunny opened his eyes and saw a little girl with yellow curls leaning over him. "give him to me," said a boy's voice. and there stood a small boy dressed in a sailor suit and a big sailor hat on which was written, "battleship uncle sam." and then billy bunny knew it was time to be going. so he gave one big hop and maybe two million and a half little skips and jumps, and soon he was far away, and if he hadn't maybe that little boy would have put him in a cage or a big box and kept him shut up for a long time. "goodness!" said the little rabbit, "i must be more careful next time." and then something happened. a little hard ball hit him on the left hind foot, and a man's voice called out, "if it hadn't been for that pesky little rabbit i would have made that hole." and the big man put his golf stick in the bag and watched billy bunny limp away to hide in the woods close by. story xvii. billy bunny and the forest dance. when the moon is big and bright little bunnies dance at night. how they hop and skip and go on their lucky left hind toe. well, sir, that's what billy bunny was doing. it was a lovely moonlight night in august, and the big, round moon was gleaming down on the pleasant meadow just like an electric lamp, only it was up in the sky, you know, and not on the ceiling. and mrs. bunny was there, too, and so was cousin cottontail, and all the little rabbits for miles around. now it's a dangerous thing to be dancing, even if the moon is bright, for owls and hawks fly by night, and if they happen to see a bunny dance, they always fly down and break it up. they don't say a word; they just fly away with one of the little bunny dancers and he never dances any more. no, sireemam. well, on this particular night little billy bunny was doing the fox trot with a nice little lady bunny, when all of a sudden from out of the friendly forest came slyboots and bushy tail, the small sons of daddy fox, you remember. and the reason they were out so late at night was because their father had sprained his foot jumping over a stone fence to get away from a pack of hounds who had chased him for a thousand and one miles and fourteen feet. now billy bunny had forgotten all about daddy fox. he was thinking only about robber hawk or old barney the owl, and so he never saw the two foxes until they were so close to him that they almost stubbed their whiskers on his powder puff tail. and if it hadn't been for the lady bunny who was dancing with him maybe slyboots, or maybe bushy tail, would have caught the little bunny. but the lady rabbit saw them just in time and she gave a scream and hopped into a hollow stump and billy bunny after her, and then all that the two foxes could do was to stand close by and say: "isn't that a shame, to spoil their little game, to stop their dancing and their prancing, who do you think's to blame?" "you are, you two bad foxes," said billy bunny, but he didn't come out of that hollow stump. no, sireemam, he staid inside and so did the little lady rabbit, and by and by the two bad foxes went away and told their father, daddy fox, all about it, and he said, "don't make any excuse. "you are very poor hunters if you can't catch a rabbit when he's dancing the fox trot." and i guess he was right, for slyboots and bushy tail were so ashamed that they didn't dare look in their mother's looking-glass for two days and three nights. and in the next story if billy bunny gets out of that hollow stump before i see him, i'll ask robbie redbreast to tell me what he does so that i can write to-morrow's story for you to read. story xviii. billy bunny and ragged rabbit. robbie redbreast told me this morning he saw billy bunny hop out of the hollow stump where he had hidden with the little lady bunny, you remember in the last story, to escape from the two bad foxes. well, after he had looked all around to make sure they were gone, he said good-by to miss rabbit. and then, so robbie redbreast told me, he looked at his gold watch and chain, which his dear, kind uncle lucky had given him for a birthday present, and it was just thirteen o'clock. "that's my lucky number," exclaimed the little rabbit; "maybe i'll find my fortune to-day." and he looked all about him, under a stone and behind a bush, but there wasn't any fortune in sight, not even a twenty-dollar gold piece. so he wound his watch and started off again; and by and by, not so very far, he came to a castle where lived a giant bunny whose name was "ragged rabbit" because he always wore torn and tattered clothes. and when he saw billy bunny hopping along, he said, "ha, ha. ho, hum, i'll eat that little bunny as sure as i'm a foot high!" and as he was twenty-one feet high less or more, he surely thought he would. "what did you say?" asked billy bunny, for his quick ears had caught the sound of the ragged rabbit's voice, but not the words. "oh, never mind," answered the ragged giant rabbit. "come and i'll show you my castle." and, oh, dear me. billy hopped in and the big giant rabbit closed the door with a bang, and all the pictures on the walls almost fell down and the chandelier rattled like a milk wagon full of empty cans. but the little rabbit wasn't frightened. and could you guess what he did if i let you guess until to-morrow night? well, sir, that brave little bunny took his popgun out of his knapsack and shot it off, and it made a dreadful loud pop, and the big ragged rabbit said, "oh, my! was that a cannon?" and then he laughed so loud that he broke a window pane and had to telephone right away to the plumber to have one put in. "that's my pop-gun, mr. giant," said billy bunny, "and if you try to hurt me i'll shoot you." and then the ragged giant rabbit laughed again, and this time the picture of his grandfather fell down and made a big dent in the floor. "if you don't stop laughing," said the little rabbit, "you'll deafen me. please only giggle." so the giant rabbit grew very polite indeed and only smiled, and then of course nothing was broken. "tell me who you are and where you are going and what time it is," he said, "and then i'll give you something to eat." but before the little rabbit could reply a loud knocking came at the door, and so you'll have to wait to hear who was there until to-morrow, for i've no more room in this story. story xix. billy bunny and tailor bird. you remember in the last story somebody was knocking at the door of the ragged rabbit's castle, don't you? the giant rabbit, who always wore torn and tattered clothes because he had no wife to mend them and wouldn't pay his tailor's bills? well, who do you suppose was on the other side of that door? just wait until the giant rabbit opens it and you shall see. now open your eyes, if you have shut them, and see uncle lucky, as sure as i am writing this story and you are reading it. yes, sir. there stood the dear old gentleman rabbit, and oh, dear me, didn't he look worried? i suppose he thought he'd find billy bunny inside the giant. but when he saw billy bunny standing there, safe and sound and happy, with his popgun in his hand and a smile on his face, he began to laugh. "whew!" exclaimed the old gentleman rabbit, greatly relieved, which means to feel much better. "i'm glad to see you, my dear nephew. and also to make your acquaintance, mr. ragged rabbit giant. my name is mr. lucky lefthindfoot. howdy!" and he put out his right front paw and shook hands with the giant, who had to lean way down to reach uncle lucky's paw. "but, goodness me!" said the old gentleman rabbit after looking at the giant for some moments, "you need a tailor. let me call the tailor bird to mend your clothes. you are too nice a rabbit not to be well dressed." and kind uncle lucky went to the telephone and told the tailor bird to bring a spool of thread a mile long and a needle as big as a spear for he had a giant customer for him with holes in his clothes as big as a circus ring. the tailor bird said he'd try to, but wouldn't promise unless he could send in a bill as big as a newspaper spread out flat. "will that be all right?" asked uncle lucky after he had explained matters to the ragged giant rabbit. "certainly," said the giant rabbit with a grin, "and tell him i'll pay him with a dollar bill as big as a turkish rug or a crex carpet." and then they all sat down and told funny stories, and billy bunny sang a song that went something like this, only much nicer, but i can't quite remember it all: "oh, you're a raggerty, taggerty man, in a castle big and old, and i'm a billy bunny boy with a heart that's brave and bold. you can't scare me with your thunder laugh or your club like a telegraph pole, so you'd better allow the tailor bird to sew up each raggerty hole." and then the tailor bird commenced and it took him until half-past fourteen o'clock to mend that giant rabbit's clothes. "i might just as well have made you a new suit," he said, as the last inch of the mile-long spool of thread was used up. "i declare i never had such a job before." and i guess he spoke the truth, for i never met a giant rabbit in my tailor's shop, although i once had a giant bill from my tailor. story xx. billy bunny and parson crow. well, after the tailor bird got his money from the ragged giant rabbit for mending his clothes, he thanked billy bunny and uncle lucky and said he must be going for he had to make a suit of clothes right away for parson crow. "if you'll wait a minute you can go with us," said kind uncle lucky; "we'll take you home in the automobile." of course the tailor bird was only too anxious to get a ride, although he did have a good pair of wings. but the needle was pretty heavy and, anyway, tailor birds don't often have the opportunity to ride in automobiles. well, after a little ways, not so very far, the luckymobile came to a stop and, of course, billy bunny had to get out to see what was the matter, and he hunted and hunted all over the machine, but couldn't find out what was wrong. by and by he saw one of the numbers had dropped off the little license plate that hung down from the rear axle. so he hopped back, and by and by, just as he was going to give up looking for it, parson crow flew by, and when he saw billy bunny he stopped and said: "what are you looking for, little rabbit?" and when billy bunny told him, he took the number 7 out of his pocket and handed it to the little bunny. "here's your number," cawed the black crow, although i never heard of a white one except once, and that was a bad bird who had been whitewashed by a colored painter because he ate up all the corn. "that's my lucky number," said billy bunny. and then the crow said in a mournful voice: "it's mine, too, and i just hate to give it up." "well, if you can get me another number, i don't care if you keep it," said the little rabbit. and then what do you think that crow did? why, he got a nice smooth little chip and made a lovely number 3 on it with a red pencil and handed it to the little rabbit. and as soon as he had tied it on the luckymobile, would you believe it if i didn't say so, that luckymobile started to go all by itself. and if billy bunny hadn't been mighty quick he would have been left behind. "where are you two rabbits going?" asked the crow as he flew alongside of the luckymobile. "because if you are not in a hurry, why don't you come with me to the meeting house to-night and hear me preach?" "we will," said kind uncle lucky, "and i'll drop a carrot cent in the collection box if you want me to." so after a while they stopped near a tall pine tree and parson crow sat on a limb and waited for all the little people of the forest to come to the meeting. well, after they were all there, he began: "now, listen to the words i say, and do your duty every day. be always good and most polite and do the things you know are right. oh, never say an angry word to any animal or bird, so when the night comes 'twill be good to feel you've done the best you could." and after that uncle lucky dropped a carrot dollar in the collection box and drove home with billy bunny. story xxi. billy bunny and jack-in-the-box. oh, i'm a rollicking jack-in-the-box, and i'm not afraid of a bear or a fox, for every one's scared when up i pop, and the little girl cries, "oh, stop! oh, stop!" i'm the bravest thing you ever saw, i'm not afraid of my mother-in-law! well, sir, i suppose you'll think billy bunny was frightened and that uncle lucky lost his breath and the automobile a tire. but nothing of the sort happened. instead, the old gentleman rabbit laughed so hard that his collar button fell out and it took him fifteen minutes and half an hour to find it. and then he never would have if the jack-in-the-box hadn't seen it first. and where do you suppose that ex-as-per-a-ting, which means teasing, button was? you'd never guess, so i'll have to tell you without asking you again. it was in the old gentleman rabbit's waistcoat pocket where he kept his gold watch and chain and pocket knife and pencil with a rubber on the end and a toothpick. "how did you see it pop into my pocket?" he asked the jack-in-the-box. "i'll never tell you," said the jack-in-the-box, "but what does that matter? you've found your collar button, and that's enough." "if i come across your cousin jack-in-the-pulpit," said uncle lucky, after he had buttoned up his collar and wound his watch, "i'll tell him how kind you were to find my collar button for me," and then the old gentleman rabbit took off his old wedding stovepipe hat and bowed to the jack-in-the-box and drove away in the luckmobile down the road, and when he came to a bridge he said to his little nephew, "do you think we're on the right road?" "i don't remember this bridge, do you?" and then a voice cried out, "don't be anxious, mr. lucky lefthindfoot. this is the road to lettuceville. "keep right on after you cross the bridge until you come to a little red schoolhouse and then turn to your left and then turn to your right and if you don't get home until morning you've made a mistake." "thank you," said uncle lucky. "and if i make a mistake i'll come back and give you a scolding," and after that they crossed the bridge, and just as they came to the first turn in the road they heard a dreadful loud noise in the woods close by. "what's that?" asked billy bunny, and he turned up his left ear and his coat collar so that he could hear better. "it's an old friend of yours," answered a deep growly kind of a voice, and before the two rabbits could wonder who it was their friend, the good-natured bear jumped out of the bushes. "take me with you, please," he said, "for i've run a splinter in my foot and it hurts me to walk." and in the next story you shall hear of another adventure which the two little rabbits had. story xxii. billy bunny and dr. duck. you remember in the last story how the good-natured bear asked billy bunny and uncle lucky to give him a ride in the luckymobile because he had run a splinter in his foot. well, as soon as he had climbed into the automobile, and it took him almost 23 1/2 seconds to do it, for the splinter was so long that it caught on the door, uncle lucky started off and by and by they came to the house where the good duck doctor lived.--dr. quack, you remember. "now, i'll go in and get him to come out and look at your splinter," said billy bunny, as he hopped out of the luckymobile and rang the front door bell, and in a minute, less or more, a nice looking lady duck came out and said, "the doctor is away on his vacation. he's gone to the lily pond for two weeks. but you can call him up on the telephone if you like. the number is waterville, 2 3 umpty eleven." so the little rabbit called up the number and when the doctor heard what was the matter, he said, "you had better come to see me. "you have the automobile right there, and it's a dangerous thing to have so large a splinter as that. tell mr. bear he'll have a dreadful corn if it isn't taken out at once." so they all hurried away and pretty soon they came to lily pond, and there was dr. duck swimming around among the pond lilies and the frogs, having a lovely time. and wasn't he sunburnt? well, i should say he was. his bill was as dark as a little brown berry and his nose was as red as a little choke cherry. "that looks very serious to me," said he, putting on his glasses and looking at mr. bear's injured feet. "i'll have to get a saw and cut off your foot." and then mr. bear gave a dreadful howl. "oh, please don't saw off my foot. it's sore enough already." "i didn't mean to saw off your foot," said dr. duck. "did i say that? i mean to saw off the splinter and then put on a poultice and draw out the pain." well, it took a long time to do all that, and the poor bear cried several times, for it hurt the splinter dreadfully, you know, to be sawed off that way. but by and by the poultice began to draw, and pretty soon out came the splinter, and mr. bear felt ever so much better. that is, until the doctor said, "it will cost you a million dollars, for that was a very serious operation." "i've never even seen a million dollars," said the bear. "nor even a million cents. you'll have to mail me a corrected bill," and then he jumped into the automobile and asked uncle lucky to drive away. "stop, stop!" cried the duck doctor, but uncle lucky paid no attention to him, any more than the bear paid the bill. "you send a corrected bill to my friend," said the old gentleman rabbit. "and, mind you, you had better correct it three times and a half if you ever want it paid." and in the next story you shall hear of an exciting adventure which the two little rabbits had with a fretful porcupine. story xxiii. bunny and the fretful porcupine. oh, never tease a porcupine, for reasons i'll relate, he's like a cushion full of pins that stand out stiff and straight. and if you stand too close i know he'll stick one in your little toe. well, that's just what uncle lucky did, and of course he got stuck with one of those prickly, stickery porcupine needles and it was an awful bother to get it out. and the fretful porcupine laughed and this made billy bunny very angry, and he took his popgun out of his knapsack and hit the porcupine on the end of the nose with the cork bullet, and this made the prickly animal run away. and after that the two rabbits started off again in the luckymobile and by and by they came to a little village where they made lollypops by the million. and the first thing uncle lucky did was to buy a big box full of them and put it in the back of the luckymobile, "for," said the kind old gentleman rabbit, "we may run across some boys and girls and then we'll have something nice to give them." wasn't that kind of him? but he was always doing nice things, was dear, kind, generous uncle lucky. well, after a while they came to some woods where a picnic was being held. there were lots and lots of children playing under the trees and the women were sitting around talking and telling their troubles, and the men were making whistles and bows and arrows for the boys and telling how they used to shoot with them when they were little boys. "helloa there, children!" cried uncle lucky, while billy bunny honked the horn. "don't you want some lollypops?" and in about five hundred short seconds there wasn't a lollypop left in that big box, and uncle lucky was a hero, or a santa claus, i don't remember which. and then one big boy said, "let's give three cheers for the two rabbits and one more for the luckymobile." and you never heard such a noise in your life. one little boy got so excited that he swallowed a raspberry lollypop and his mother had to reach down his throat and pull it out by the stick. "now be good until i see you again," said the kind old gentleman rabbit as he drove off, and by and by billy bunny saw something moving among the trees. "what's that?" he said to his rabbit uncle. but before the old gentleman rabbit could reply, a big stone hit one of the lamps on the automobile and broke it to splintereens. "stop that whoever you are!" shouted billy bunny. "if you do it again i'll shoot!" and he held his popgun up to his shoulder just like a soldier boy in battle. and if the little canary in my room doesn't wink at me all night so that i can't hear the alarm clock in the morning, i'll tell you another story. story xxiv. billy bunny and danny billygoat. well, my little canary bird didn't wink at me all night, as i feared it might in the last story, and my alarm clock said "good morning" to me at half-past fourteen o'clock, so i got up in time, and here is the story i wrote before i went out into the garden to eat raspberries with robbie redbreast. one evening as uncle lucky and billy bunny were driving along in the luckymobile, who should they come across but a little billygoat named danny. he had a little beard that hung down from his chin and two little horns that stuck up from his head, and he was playing on a flute while he sat cross-legged on a stone by the roadside. and when he saw our two small friends in their machine, he began to play: it's not so far to the twinkle star in the little white boat of sleep. so list to my tune, like a breeze in june, where the honeysuckles creep. over the sky, way up high, in the little white boat of sleep. ever so far to the twinkle star way up in the sky blue deep. "where did you learn that lullaby," asked kind uncle lucky, brushing a tear from his eye, for he remembered just a little song his mother used to sing when he was a little boy rabbit, you know. "i don't know," answered danny goat. he pulled on his goatee and smiled, and then he began again: "up in the sky when the sun is high the white cloud boats go sailing by, and the summer breeze in the tall, tall trees is singing a song the whole day long. and this is the song they sing: we ring the bell in the cool damp dell that grows on the lily's stalk, we bend the ferns in the river's turns and the tail of the great gray hawk; and the foamy spray in the big deep bay we blow on the great boardwalk." "that reminds me of atlantic city," said uncle lucky. "let's drive down there and go for a swim." "just the thing," said the little rabbit; "i've got my bathing suit in my knapsack. i'm ready." so off they went, and by and by they came to the seashore. but there wasn't a hotel in sight, so of course they knew they had made a mistake. they didn't care, especially billy bunny, for not very far from land was the big good-natured whale who had taken him for a sail a long, long time ago. "there's my friend the whaleship!" cried the little rabbit. and in the next story, if that whale doesn't swim away, i'll tell you something more about billy bunny and his kind uncle lucky. story xxv. billy bunny and the whale. you remember in the story before this that billy bunny and uncle lucky were at the seashore, and out a little ways from the land was the good-natured whale. well, as soon as he saw the little rabbit he swam up to the beach and said "hello." and then billy bunny introduced him to uncle lucky, and after that the whale said: "don't you both want to go for a sail?" and as the old gentleman rabbit had never been on a whaleship in his life, he said yes right away, and so did the little rabbit. then the whale pushed his tail up on the sand and the two little rabbits hopped over it just like a bridge, and then they sat down, and away went the whale with a swish of his tail that spattered the spray all over the bay. "goodness me!" cried the old gentleman rabbit, "i'll have to wipe off my spectacles," and he took his polka-dot handkerchief from his pocket, and after that he tied it over his old wedding stovepipe hat, for he wasn't going to lose that hat, no siree, and a no sireemam, not even if he had to tie the anchor to it. by and by, not so very long, they heard a sweet voice singing, so they looked everywhere, but the only thing they saw was the big green ocean. "i wonder who is singing?" said uncle lucky, and he took his spyglass out of his waistcoat pocket and twisted it around and around until he could see distinctly, which means plainly, you know. "there she is!" cried the old gentleman rabbit, and he got so excited that he looked through the wrong end of the spyglass and then he said, "no, she isn't!" for he couldn't see anything at all that way, you know. "what did you see?" asked the little rabbit, and he pushed forward uncle lucky's old wedding stovepipe hat to keep it from falling over his left ear. "a mermaid!" cried the old gentleman rabbit, and before he could turn the spyglass the other way a lovely mermaid swam up and handed him her card, and on it was written in lovely purple ink: miss coral seafoam, oceanville, u. s. a. "pleased to meet you," cried the old gentleman rabbit most politely. "this is my nephew, william bunny, brier patch, old snake fence corner, and my name is mr. lucky lefthindfoot and i live in lettuceville, corner of carrot and lettuce streets," and then he tried to take off his hat, but he couldn't, for it was tied down tight, you remember, with his blue polka-dot handkerchief. and after that the mermaid asked them to visit her coral island, where she and her sisters sold coral beads and scarfpins. and in the next story you shall hear--well, i guess i won't tell you now, but let you wait and see. story xxvi. billy bunny and the mermaid. well, now we'll commence by saying that as soon as billy bunny and uncle lucky reached the coral island, where the lovely mermaid lived, for she had asked them to call, you remember, they got off the whale, and, after asking him to wait for them while they made a little visit, sat down on the sand, and pretty soon the mermaid brought them each a lovely coral scarfpin, and the one she gave to uncle lucky was a little image of herself and the one she gave to billy bunny was a little fish. then the little rabbit opened his knapsack and took out a lovely apple pie and gave it to her. and she was so pleased that she ate it all up, and then she said, "i'll give you a lovely breast-pin made of beautiful coral for your mother, mr. billy bunny, if you'll give me another pie." so the little rabbit opened his knapsack and took out another fresh, juicy apple pie and placed the beautiful present for his mother carefully in the knapsack, and after that he ate a lollypop and uncle lucky drank a bottle of ginger ale, and then they said good-by and got aboard the whaleship and sailed away. and would you believe it? dear, kind uncle lucky almost cried! you see, he had never seen a mermaid before, and he thought she was lovely, and i guess she was, for uncle lucky couldn't make a mistake, i'm sure, for he had travelled abroad and had seen lots and lots of beautiful lady bunnies. "and now where are we going?" asked the little rabbit, but uncle lucky was too busy trying to find his other blue polka-dot handkerchief with which to wipe his eyes to answer. and then he couldn't find it, and the reason was because he had given it to a chinaman the day before, but he didn't remember that, for he was so miserable at leaving the beautiful mermaid. "oh, dear! oh, dear!" sighed the old gentleman rabbit, "'tis sad to part. my poor old heart is nearly, nearly breaking; alas! alas! that mermaid lass has set my head a-shaking!" and after that his old wedding stovepipe hat almost fell off his head, and it would have, i'm sure, if it hadn't been for the blue polka-dot handkerchief which he had tied over the top of it. and just then, all of a sudden, the whaleship bumped into a motor boat, and nearly upset it. "what's the matter with your pilot?" screamed the man who was in the motor boat, and when uncle lucky looked over the side of the whale he saw it wasn't a man at all, but the old billygoat who owned the ferryboat i told you about some umpty-leven stones ago. "excuse us, please," said the kind old gentleman rabbit, but what the billygoat said i'll have to tell you in the next story, for there's no more room in this one. story xxvii. billy bunny and the beanstalk. seeing it's you," answered the billygoat, who, you remember in the last story, had gotten very angry because billy bunny and uncle lucky had bumped into his motor boat with their whaleship. "i'll forgive you," and then he raced the whale all the way to the shore and would have beaten him, too, if he had gone faster. and as soon as the whaleship ran up on the beach, the two little rabbits hopped off and got into their automobile and drove away, and the whale went back and told the mermaid that the two little rabbits had a beautiful luckymobile, and she felt dreadfully sorry that she hadn't gone with them. well, after a little while, not so very far, they came across a wonderful beanstalk, which was growing up so high that you couldn't see the top, and if billy bunny had only known the story about "jack and the beanstalk," i guess he would have thought that the story had come true. "my gracious!" exclaimed uncle lucky. "my lima beans at home grow pretty high but never as high as this," and he took out of his waistcoat pocket his spyglass and tried to find the top of the beanstalk; but he couldn't, for it was hidden in the clouds. just think of that! "i'm going to climb up that beanstalk," said the little bunny. "maybe i'll find my fortune at the top." "and i'll go with you," said the old gentleman rabbit, for he wasn't going to let his small nephew go up a strange beanstalk and perhaps get lost in the clouds, you know. not good, kind uncle lucky. no, sireemam; so they hopped out of the luckymobile and started up the beanstalk, and by and by, after a pretty long time, they came to the top and the first thing they saw was their friend american eagle and his wife, and she was sitting on her nest hatching out the big eggs which she had laid. "we'll need lots of eagles now that we've gone to war," said the big bird, and he flapped his wings and sang "yankee doodle dandy" three times over and then once more. and this made the old gentleman rabbit so excited that he stood up and made a speech, and then he threw his old wedding stovepipe hat up into the air and gave three cheers and half a dozen tigers and two or three bears. and after that billy bunny opened his knapsack and took out an american flag and put it on the top of the beanstalk so that all the people in the aeroplane could see it and say "hip-hur-ray for the u. s. a.!" "when the little eagles come out of their shells you must bring them to call on me," said good, kind uncle lucky to mrs. eagle. "i have some popcorn and lollypops at home, and i know how children like those things." and this made mrs. eagle very happy and mr. eagle very proud, and he helped the two little rabbits to climb down the beanstalk in time for me to write what they did in the next story, which will be about an adventure in the friendly forest. story xxviii. billy bunny and scatterbrains. after billy bunny and uncle lucky reached the ground, for they had climbed down the beanstalk, you remember, as i told you in the last story, they jumped into the luckymobile and drove off toward the friendly forest, and when they had gone maybe a mile in and out among the trees, for there wasn't really any automobile road to go on, you know, they came across scatterbrains, the gray squirrel. now uncle lucky knew old squirrel nutcracker very well, and as the old gentleman squirrel was very nice and well behaved it made uncle lucky provoked to think that his son should be such a scatterbrains. so uncle lucky stopped the automobile and said: "well, young squirrel, have you been troubling your father lately?" and scatterbrains answered, "no, mr. lucky lefthindfoot, not lately. not since yesterday." "what!" exclaimed the old gentleman rabbit, "do you mean to say you troubled him yesterday? why didn't you wait until to-morrow?" and then uncle lucky winked at billy bunny and then scowled at scatterbrains. and just then they heard a dreadful noise. it sounded just as if the trees were snapping to pieces and, all of a sudden, a tornado struck them and up in the air went the luckymobile with the two little rabbits, but what happened to the little squirrel i really don't know, unless it took him up, too, and hid him in a cloud. and perhaps it did, for i've often seen clouds that looked exactly like squirrels, haven't you, and other animals, too, like bears and cats? "gracious me!" cried uncle billy. "hang on, billy bunny, and don't let the cushions slip or the electricity run out of the cabaret, for if we ever get back to earth, i'd like to get home and stay home forever. oh, home, sweet home," and the old gentleman rabbit took off his automobile goggles, for they were full of tears and he couldn't see anything. well, by and by, the tornado let go and the automobile fell on top of a clothesline and balanced there as nicely as a tight-rope dancer, and when the two little rabbits looked about them, they found they were in mrs. bunny's backyard in the old brier patch. wasn't that lucky? well, i guess it was! and just then mrs. bunny came out of the kitchen door to hang up some of billy bunny's little shirts on the line, for it was monday morning, you know. and when she saw the luckymobile on her clothesline she gave a scream, and then she began to laugh, and after that she ran back into the house and brought out her scissors and cut the rope and the automobile came down with a bang, and out tumbled the two little rabbits. "well, well, well," said mrs. bunny, and she sat down on the clothespin basket and laughed, but, of course, there weren't any clothespins, or any other kind of pins, in it, you see, for then she wouldn't have laughed. and in the next story, if my umbrella doesn't open and stand over my bed to keep off the mosquitoes, i'll tell you another story to-morrow night. story xxix. billy bunny and mrs. black cat. awake, awake, 'tis early morn. the cow is climbing the stalks of corn, the little bird is beating an egg, and the rooster is dancing about on one leg, and the pig is trying on her new bonnet, with a little blue bow and a red cherry on it. uncle lucky rolled over in bed and then he got up and wiggled his nose and his left ear, and after that he was so wide awake that he didn't want to get back into bed, as i did, when i woke up this morning. and just then the breakfast bell rang and mrs. bunny put on the coffee and the baked lollypops and the stewed prunes, and, oh, dear me! i really can't remember what rabbits eat every day, for i'm sure they don't eat the same old thing, for if they did they wouldn't be jolly and gay and hop about merrily all through the day, but would sit in a corner and sulk and be sad, and maybe get angry and maybe get mad. so always remember to have something new, for no one can always enjoy a prune stew. there! i've gone and written another piece of poetry and my typewriter wouldn't print it properly. isn't that too bad? well, after breakfast the old gentleman rabbit went out for a walk in the pleasant meadow, and he went all alone, too, for billy bunny had to stay home and polish the front door knob and sweep the piazza and feed the canary and bring in the wood, for mrs. bunny had to hurry up with the breakfast dishes so as to be able to go over and see cousin cottontail, who had just had a new baby rabbit. well, as i was saying, uncle lucky hopped along the pleasant meadow until he came to the old farm yard where cocky docky and henny jenny and all the other barn yard folk lived with the good-natured farmer. and just as he was going through the gate, who should bounce out at him but a big black cat. and, oh, dear me. her claws were sticking out of her feet like pins and her eyes were yellow as fire and her teeth glittered and her whiskers stood out like bayonets, and her tail was as big as a rolling pin and her back was humped up worse than a camel's. if you can think of anything worse than the way that cat looked i wish you would write me a letter and tell me so that i can scare uncle lucky, for, would you believe it, he wasn't the least big frightened. no, sireemam. he just took off his old wedding stovepipe hat and bowed most politely to mrs. black cat, and she was so surprised that she turned around and went back to her three little kittens who never wore mittens because they didn't have any. and after that the old gentleman rabbit hopped into the barn and ate some corn and had a talk with mr. sharptooth rat. and maybe he would have been talking there yet if something hadn't happened. and when you don't expect it, something very often, and sometimes most always, does happen. the miller's dog ran into the barn and made a grab for the old gentleman rabbit, but uncle lucky was too quick for him. he hopped to one side and then out of that barn so that he hopped right into to-morrow night's story. wasn't that wonderful? story xxx. billy bunny and big yellow dog. let me see. didn't i say that billy bunny hopped out of the old barn so fast in last night's story that he jumped right into this one? well, he did, and here he is saying, "i'm ready for another adventure!" and no sooner had he said this than along came a big yellow dog with a muzzle on his nose, and when the little rabbit saw him he laughed out loud, "oh, ho! mr. yellow dog! did you put your nose into a mouse trap?" "no, i didn't," replied the yellow dog. "it's a muzzle to keep me from biting little rabbits," and then he gave a dreadful growl and tried to pull off the muzzle with his front paws. "i won't wait until you get it off," said billy bunny, and he hopped away as fast as he could, for he wasn't the least bit curious to see whether that muzzle was tied on tight! and by and by he came to a hollow stump where lived an old rabbit named hoppity-hop. "helloa, my little friend," said the old rabbit, and then he wriggled his nose a million times or less, for i guess he smelt the lettuce sandwich which billy bunny had in his knapsack. "good morning," said billy bunny, but he didn't open his knapsack. no, sir! it wasn't fourteen o'clock, which is the luncheon hour in rabbitville, so i've been told. and this, of course, made the old rabbit very sad. "oh, dear me," he cried, "i'm so hungry, and if there is anything i love more than a lettuce sandwich it's apple pie!" "how do you know i've got an apple pie?" asked billy bunny, and he took out his gold watch and chain to see what time it was, for he began to feel hungry all of a sudden. but, oh, dear me! it wasn't fourteen o'clock, or anywhere near it, so he twisted the stem of his watch until the hands pointed at the luncheon time, and then he took out the lettuce sandwich and the apple pie and he and the old rabbit ate them up right then and there, and after that they felt ever so much better. "now i'll tell you a secret," said the old rabbit. "there's a carrot candy shop not very far from here, and if you've got any money in your knapsack i'll take you there." wasn't that kind of that old rabbit? so off they hopped and pretty soon, not so very far, they came to the candy shop, and the old lady woodchuck who kept it was awfully kind and generous, for she filled up a paper bag right to the top for a lettuce dollar bill, which i think was a very cheap price to pay for all that candy, don't you? and when it was all gone, billy bunny said good-by and hopped away singing at the top of his voice: "oh, who is so merry and who is so gay as a rabbit who always has money to pay for candy and popcorn and nice apple pie and other sweet things that you're longing to buy." and in the next story, if billy bunny does eat any more carrot candy and get so dizzy he can't hop in a circle, i'll tell you some more about the little rabbit. story xxxi. billy bunny and a happy birthday. it very often happens you don't know what to do, and then's the time the mischief man comes smiling round to you. he whispers something in your ear you know you shouldn't stop to hear, and then's the time for you to say, "oh, mischief man, please go away!" this is what dear good uncle lucky wrote in billy bunny's album, for it was the little rabbit's birthday, you know, and uncle lucky thought he ought to warn him against the mischief man. well, as soon as the ink was dry so that the little rabbit could put the album away in uncle lucky's desk, the kind old gentleman rabbit said: "let us take a ride in the luckymobile. maybe we can go some place where we will have a good time." so they got into the automobile and started off, and by and by they came to a shady spot in the woods. and there right under a big spreading chestnut tree, was a little table covered with a clean white cloth and in the middle was a lovely birthday cake with candles and big frosted letters, which read, "a happy birthday to billy bunny!" and oh, my, wasn't he delighted and so were all the little forest folk, for they were all there, let me tell you, from old squirrel nutcracker to the big brown bear. and so were the little people from the pleasant meadow, dicky meadow mouse and robbie redbreast and many others. and pretty soon along came the barnyard folk, cocky docky, henny jenny and duckey daddies. even mrs. cow wasn't too busy to be there, and if you'll wait a minute i'll tell you the names of some more of billy bunny's friends: turkey purky, danny beaver, old mother magpie, timmy chipmunk, scatterbrains, the gray squirrel, and shadow tail, his brother. daddy fox would like to have been there, only uncle lucky hadn't sent him an invitation. the only friend who wasn't there was uncle bullfrog. he couldn't leave his log in the old mill pond, so he sent his regrets by little mrs. oriole, who lived in the willow tree by the old mill. "now we'll cut the cake," said kind uncle lucky, and he went over to the luckymobile to get the big carving knife which he had hidden under the cushions. "there's a little gold ring hidden away somewhere," he said as he cut the cake very carefully so as not to topple over the pretty candles and get the pink and green melted wax all over the white frosting. and then everybody ate up his piece of cake as fast as he could to find the little gold ring. "i've got it! i've got it!" screamed timmy chipmunk. but, oh, dear me. it wasn't the ring at all. it was only a hard nut. and the little chipmunk was so disappointed that he ran home to tell his mother all about it, and she gave him one she had found when she was a little girl in the toe of her stocking one happy christmas morning. and in the next story you'll be surprised to hear who got the ring after all. story xxxii. billy bunny and the lost ring. something's going to happen; i feel it in the air. but what it is you soon shall know, so hold your breath and stare. you remember in the last story i told you about billy bunny's birthday party and promised to tell you who found the little gold ring in the frosted cake. well, just as the little rabbit said, "i've found it!" daddy fox sprang from behind a bush and grabbed the piece of cake right out of the little rabbit's paw. and then he jumped over the luckymobile and ran off to his den to give it to slyboots or bushy tail, his two little sons, you know, but which one got it i can't remember, for everybody was so excited that they forgot to ask the naughty old fox before he got away. "that's too bad," said kind uncle lucky; "i'll have to get you another one," so he said good-by to everybody and took billy bunny down to the 3 and 10 cents store, where they bought a lovely gold ring with a big ruby in it. wasn't that nice? and then they came back to the woods, but everybody had gone home and there was no more birthday cake anywhere to be seen, not even a little piece of candle. "well, what shall we do now?" said the kind old gentleman rabbit, and he poured some lettuce oil into the cabaret and took out his blue polka-dot handkerchief and wiped his ear, and then he dusted off his old wedding stovepipe hat and honked the automobile horn and blew up a tire and turned a cushion upside down to hide a grease spot. and after that he put on his goggles and started off again, and by and by, not so very long, they came to a signpost on which was written: "which road shall i take?" "goodness, gracious me!" exclaimed the old gentleman rabbit, "what's the matter with my goggles?" and he took them off and looked at the signpost again. "it says the same old thing," he said with a sigh, and he took off his old wedding stovepipe hat and dusted the top, and after he had put it on his head again he heard a voice saying: "take the road that leads to the left, and not the one to the right, for if you don't you will get left and you won't get home till night." "who's speaking?" said billy bunny. and the reason he hadn't said anything before was because he had been sound asleep. and then who should come out from behind that funny signpost but a great roaring bull with two horns and about ten feet long and big red, snorting nostrils. "don't let us disturb you," which means bother or something like that, said uncle lucky, and he honked the horn with all his might, and, would you believe it, the bull was so frightened that he ran away and never stopped till he got home and covered himself with the crazy quilt on his old four-poster bed. story xxxiii. billy bunny and the great news. once upon a time, so i've heard tell, there lived a little rabbit in a shady dell. and on one side a clover patch, where red-topped clovers grew, and 'tother side was lollypops of red and white and blue. this is the song mrs. bunny sang one morning as she set to work to wash her little rabbit's white duck trousers, for it was monday, and that is washday in rabbitville, so they tell me. and just as she was hanging them out on the line who should fly up but old mother magpie, and, my! wasn't she excited. why, she was so disturbed that her bonnet had fallen off her head and was hanging by the strings. "have you heard the news?" she asked, and she rolled off one of her black silk mitts and turned her wedding ring around three times and a half. "heard what?" asked mrs. bunny, putting the clothespin in her mouth instead of on the clothesline. "why, the miller's boy has gone off to the war." "hurray!" shouted little billy bunny, who was polishing the brass door knob on the back door. "hurray!" "you ought to be ashamed of yourself," said old mother mischief. "his poor mother is nearly crazy with grief." "i'm sorry for her," said mrs. bunny, and she thought how thankful she ought to be that her little rabbit didn't have to shoulder a musket. "well, i'm glad he's going," said billy bunny. "he can shoot at something else now besides little rabbits." old mother magpie ruffled her feathers. "well, if i had a boy like you i'd teach him not to glory over another person's grief," and then she flew away. "i'm sorry for his mother," said mrs. bunny, "but the miller boy will never be missed," and the clothespin fell out of her mouth and stood up in the grass like a little wooden soldier. "do you want anything at the store?" asked the little rabbit, after he had finished cleaning the door knob. "if you do, tell me, for i'm going by there." "you can order a pound of carrot tea and some lollypops," answered his mother, and then billy bunny picked up his striped candy cane and set off for the village, and by and by he came to the post office and the nice lady postmistress called to him that there was a letter there addressed to billy bunny, old brier patch, but what was written in it i'm not going to tell you now, for i must stop and play a game of pinochle with dear, kind uncle lucky, who just telephoned me to come over to his house and have a game with him this evening, and i mustn't keep him waiting another minute. story xxxiv. billy bunny and jenny muskrat. well, i played pinochle with uncle lucky lefthindfoot last evening and it was so late when i got home that i overslept myself this morning. and maybe i'd have slept all day if robbie redbreast hadn't come to my window and told me that billy bunny was reading a letter which i told you about in yesterday's story and that every time he turned a page he laughed harder than ever. well, i was so curious to know what he was laughing at that i told robbie redbreast to fly back to him and look over his shoulder and see what was in the letter while i hurried and dressed as fast as i could, and when i was all ready to go into the friendly forest where the little rabbit was, i saw him coming toward me with the letter in his hand and the little robin perched upon his knapsack. "good morning," he said and handed me the letter, and now you shall hear what was written to mr. william bunny, brier patch, old snake fence corner, u. s. a., care of uncle sam! "my dear billy bunny: "just a few lines from your old friend the circus elephant to tell you that he is coming to see you as soon as he gets over the measles. if you've never had the measles, dear billy bunny, don't get them, for they are dreadful things for there's so many of them. "please give my love to mr. lucky lefthindfoot and tell him as soon as i'm well, i'll be back in his circus. "your friend, "elly." and as soon as i'd read the letter the little rabbit put it in his pocket and hopped away and by and by he came to a little stone house by a river. and before i go any farther i'll just whisper to you how i know all this. you see, the little robin told me all about it, for he and i are great friends and his nest is in the old apple tree just under my window. well, pretty soon, after looking all around, billy bunny knocked on the door of the little stone house and in a few minutes it was opened by a nice lady muskrat, whose name was jenny eva. "how do you do, little rabbit," she said, and then she invited him in and gave him a cookie made out of carrot seeds and pumpkin flour. and after that he showed her the letter from his friend, the circus elephant, and just then, all of a sudden, the front door flew open and in came the miller's dog. and, oh, dear me! mrs. jenny eva muskrat forgot all about her society manners and ran down the back stairs into the river and the little rabbit forgot to say good-by and hid himself in a big hat box where she kept her last year's easter bonnet. and then, what do you suppose the miller's dog did? why, he began to sing: "old mrs. muskrat jumped into the river, splasherty, splasherty, splash! and little boy rabbit jumped into the box, that held her best bonnet and trampled upon it. masherty, masherty, mash!" and in the next story you shall know what the miller's dog did when he stopped singing, that is, if robbie redbreast isn't too frightened to look into the window and tell me all about it. story xxxv. billy bunny and the miller's dog. after the miller's dog stopped singing, as i told you in the story before this, he poked his nose into the hat box where billy bunny had hidden himself and said in a deep, growly voice: "come out of there or i will growl and bite the bonnet that mrs. muskrat wears for best and the purple flowers on it. and then she'll think it's you who did this dreadful unkind deed, and never speak to you again or you with cookies feed." "goodness me, but you are a very poor sort of a poet," said the little rabbit, peeping out of the hat box. "your poetry is dreadful," and this made the miller's dog so ashamed of himself that he couldn't wag his tail or even bark. no, sir. he couldn't do a thing but slink out of the door and close it so softly that it didn't pinch his tail hardly at all. "ha! ha!" laughed the little rabbit. "did you ever see such a silly dog?" and neither did i and neither did you, i know. well, after a little while, mrs. jenny eva muskrat carne up the back stairs from the river, where she had gone in the last story, you remember, and wasn't she glad that nothing more had happened? "if you had jumped into that other hat box," she said, "you would have spoilt my next year's easter bonnet, and that would have been too dreadful for anything." and wasn't the little rabbit glad? well, i guess he was twice over and maybe three times. and after that he said good-by and hopped away, and after he had traveled for a long, long ways he came to the field where his old friend the scarecrow lived. "how have you been?" asked the little rabbit, and he took a lollypop out of his knapsack and offered it to the scarecrow, but he didn't want it. "haven't you got a cigar?" he asked. "i haven't smoked for ever so long." "i'm sorry," said billy bunny. "i don't think i have any really and truly cigars. here's a chocolate one if that will do," and he handed it to his friend the old clothes man. but the old clothes man couldn't smoke it at all, although he tried the best he could, and pretty soon it began to rain and the chocolate became soft and sticky, and the little bunny all wet, so he said: "i guess i'll crawl into a hollow stump if i can find one." and it didn't take him long, for he hopped away to the woods nearby, and the first thing he saw was an old stump, so he hopped inside. and no sooner was he safely out of the rain than a voice said: "what are you doing in my hollow stump; who are you anyway? why didn't you knock on this old wood block if you really want to stay?" and in the next story i'll tell who it was that said this. story xxxvi. billy bunny and the woodchuck. you remember in the last story that just as billy bunny hopped into the hollow stump a voice said, "what are you doing in here?" "i came in to get out of the wet," answered the little rabbit, and then the voice replied: "what! is it raining? i'll lend you an umbrella!" and an old woodchuck opened a little door in the side of the stump and winked at billy bunny. "that's very kind of you," said the little rabbit, and he opened his knapsack and gave the woodchuck a nice lollypop, and after that the woodchuck said: "i think you'd better stay here with me until the rain is over. don't you think so?" and billy bunny said yes, for the woodchuck was very nice and had such good manners that the little rabbit felt quite at home. but oh, dear me! it began to rain so hard right then and there that the water just poured into the old hollow stump, and pretty soon it was very uncomfortable. so the woodchuck said: "now don't you ever tell anybody where i'm going to take you. for it's my very own house, and i never let anybody know just where i do live. you see, so many people are after me, some with guns and some with sharp teeth and claws, that i have to be very careful." so the little rabbit promised, and then he followed the woodchuck through the little door and down a long passage until they came to a nice, large, comfortable room. "now, this is where i live," said the woodchuck, and he went over to the cupboard and took out a carrot candy gumdrop and gave it to billy bunny, and then he lighted a big cigar and sat down in his old armchair and smoked. and all the time they could hear the rain pattering on the grass overhead, for it's wonderful how you can hear all sorts of sounds when you're under ground and have big ears like a rabbit, you know. "now, i'll tell you a story," said the old woodchuck after he had blown some lovely round rings of smoke into the air. "once upon a time, not so very long ago, a band of tiny fairies lived in the woodland near. and often i would hear them a-singing soft and low when all was dark and quiet and the moon shone bright and clear. so one evening i stole softly out of the hollow stump, and found them dancing merrily with tiny skip and jump; and just as i was going to say how do you do, the fairy queen began to scream. and then away she flew. and then her tiny subjects took fright and ran off, too, and now i never see them more a-dancing near my old stump door." "that's too bad," said the little rabbit, for he was so interested in what the old woodchuck was saying that he had forgotten all about his lollypop and had dropped it on the floor. and in the next story he'll pick up his lollypop and eat it, because i hate to have him lose it, don't you? story xxxvii. billy bunny and little peewee. let me stop for a moment and think where i left off last night. oh, now i remember. billy bunny was in the old woodchuck hollow stump, and it was raining. oh, my, yes. cats and dogs, as they say in grown-ups' stories, so we'll say kittens and puppies. well, after a while the rain stopped and the little rabbit said good-by and hopped away, and pretty soon, not very long, a little bird began to sing: "down the shady forest trail, o'er the hill and through the vale, billy bunny hops along with a whistle and a song. and if you have never heard a rabbit whistle like a bird, you must ask each little rabbit if he has the whistling habit." "who's singing?" asked billy bunny, and he took his silver policeman's whistle out of his knapsack and blew on it so hard that the little bird began to cry: "oh, dear! oh, dear! you will whistle my ear off!" and then, of course, the little rabbit stopped, for he didn't want to hurt that dear little bird. no sireemam. "who are you?" he asked, and the little bird replied: "i'm peewee, the littlest bird in the whole friendly forest." "what do you look like?" said the little rabbit, curiously, gazing here and there and everywhere and behind a tree and under a stone. "i've never seen a peewee." and then that little bird flew down from a tree and billy bunny saw the tiniest little bird he had ever seen. why, it wasn't much larger than a butterfly. "goodness, but you're small," said billy bunny. "are you so small that you don't like lollypops?" of course, the little bird said no, and so would you, no matter how small you were, but when she tried to fly away with the lollypop, she couldn't. no sireemam. wasn't that too bad? so the little rabbit gave her some sweet cracker crumbs instead, and after that he hopped away looking for another adventure. and it wasn't long before he had one. for, just as he was hopping across a fallen log that made a narrow bridge over a brook, a little fish swam up to the top of the water and said: "here is a letter from your friend, the whale," and he held up in his mouth a blue envelope. i guess it was made of some kind of waterproof paper, for it wasn't the least bit damp. and when billy bunny opened it, he found a small coral ring inside, and in the letter it said: "this ring is for you, billy bunny. "the pretty mermaid asked me to send it to you, so here it is. please tell the little fish that you have received it and that it fits you perfectly." and then the whale signed himself, "your great big-hearted friend, the whale." story xxxviii. billy bunny and old mother magpie. uncle bullfrog sings a song that is never very long. all he says is, "chunk, ker-chunk!" then he splashes in ker-plunk, and the little fishes swim, oh, so fast away from him! if they didn't, don't you think he would eat 'em in a wink? now who do you suppose was singing this song? why, a little tadpole named taddylegs. and it made uncle bullfrog quite cross, for he didn't like tadpoles anyway, and taddylegs wasn't very polite, as you can see. "now swim away," said the old gentleman frog, and he looked angrily at taddylegs. "now swim away or i'll swallow you and maybe your cousin and your aunt if they're around." so the little tadpole swam away and after a while old uncle bullfrog saw billy bunny not very far away. he was talking to mrs. cow about the clover patch. you see, mrs. cow was very fond of clover and so was the little rabbit, and he knew that mrs. cow could eat maybe three hundred and forty-seven times as much clover as he could, and so he was afraid she might eat up the whole patch and leave nothing for anybody else. "please don't eat all the clover tops; mother wants to preserve some for the winter." "don't you worry," replied mrs. cow, and she whisked a big horse fly off her side with her long tail. "don't you worry and don't you fret, there'll be some clover blossoms yet." so the little rabbit felt ever so much better and hopped away and by and by he came across old mother magpie. and he wasn't a bit pleased, for she was always finding fault with him, and everybody else, for that matter. yes, old mother magpie made lots of trouble and billy bunny had never liked her. but he couldn't get away without her seeing him, although he tried his best. "good morning, billy bunny," said the old lady magpie, and she raised her bonnet so she could see him better, for the brim was half over her left eye. "good morning," replied the little rabbit. "i'm sorry, but i'm in a dreadful hurry," and he hopped away so fast that he left his shadow a mile behind him. "gracious me!" exclaimed old mother magpie. "that bunny doesn't like me very much i guess." "yes, you don't have to guess again," cried a voice, and parson crow cawed and hawed, and this made the old lady magpie so angry that she flew away to tell barney owl that she was a very much abused person. but here we are at the end of this book, and so we will have to jump to the next, which i will call, "billy bunny and uncle lucky lefthindfoot." the end university of kansas publications museum of natural history volume 18, no. 6, pp. 505-545, 7 figs., 4 pls. december 2, 1969 the systematics of the frogs of the _hyla rubra_ group in middle america by juan r. león university of kansas lawrence 1969 university of kansas publications, museum of natural history editors: frank b. cross, philip s. humphrey, robert m. mengel. volume 18, no. 6, pp. 505-545, 7 figs., 4 pls. published december 2, 1969 university of kansas lawrence, kansas printed by robert r. (bob) sanders, state printer topeka, kansas 1969 the systematics of the frogs of the _hyla rubra_ group in middle america by juan r. león contents page introduction 508 acknowledgments 508 materials and methods 509 the hyla rubra group 509 key to species and subspecies 510 key to known tadpoles 511 accounts of species and subspecies 511 _hyla boulengeri_ (cope) 511 _hyla foliamorta_ fouquette 520 _hyla rubra_ laurenti 524 _hyla elaeochroa_ cope 525 _hyla staufferi_ cope 532 _hyla staufferi staufferi_ cope 537 _hyla staufferi altae_ dunn 540 evolutionary history 540 literature cited 543 introduction the tree frogs of the _hyla rubra_ group are abundant and form a conspicuous element of the neotropical frog fauna. representatives of the group occur from lowland méxico to argentina; the greatest diversity is reached in the lowlands of southeastern brazil (cochran, 1955). the group apparently originated in south america; the endemic central american species evolved from stocks that invaded middle america after the closure of the colombian portal in the late pliocene. dunn (1933) partially defined the _rubra_ group as it occurs in central america. cope (1865, 1876, 1887), brocchi (1881), boulenger (1882), günther (1901), noble (1918), kellogg (1932), dunn and emlen (1932), stuart (1935), and gaige (1936) dealt with the middle american species now considered to make up the _rubra_ group. more recently, taylor (1952, 1958), fouquette (1958), starrett (1960), and duellman (1960, 1963, 1966a) studied aspects of the taxonomy and biology of the species of this group. the five species of the _rubra_ group in central america have received ten different names. one species, _hyla staufferi_, has received five names (two subspecies are recognized herein). _hyla boulengeri_ was named in the genus _scytopis_, but the type species of _scytopis_ is a member of the genus _phrynohyas_ fitzinger, 1843 (duellman, 1956.) little has been published concerning the ecology, life history, osteology, and mating calls of the middle american species of this group. the purpose of the present report is to describe the species occurring in middle america and to comment on their distributions, ecology, cranial osteology, and mating calls, and in so doing provide evidence for the evolutionary history of the species inhabiting middle america. acknowledgments for permission to examine specimens in their care, i am grateful to drs. richard g. zweifel, american museum of natural history (amnh); robert f. inger, field museum of natural history (fmnh); ernest e. williams, museum of comparative zoology (mcz); hobart m. smith, university of illinois museum of natural history (uimnh); charles f. walker, university of michigan museum of zoology (ummz); jay m. savage, university of southern california (usc); james a. peters, united states national museum (usnm); richard j. baldauf, texas cooperative wildlife collection (tcwc); and w. frank blair, texas natural history collection (tnhc). ku refers to specimens in the museum of natural history, university of kansas. for the loan of tape-recordings of mating calls i thank drs. w. frank blair, university of texas, and richard g. zweifel, american museum of natural history. i am indebted to the ford foundation-universidad de oriente (venezuela) science project for a scholarship which enabled me to study for two years at the university of kansas, foster institution of the project. i have benefited by being able to work in the museum of natural history at the university of kansas and i am grateful to dr. e. raymond hall, director, for providing space and equipment. i gratefully acknowledge the assistance and advice of dr. william e. duellman, who suggested and directed this work, made available specimens under his care and gave much of his time in reading the manuscript and suggesting improvements. i am grateful to dr. frank b. cross who critically read the manuscript and made many editorial suggestions. i am indebted to linda trueb for assistance with the osteological aspects of this study; she helped to clarify many confusing points. i am grateful to charles w. myers for making available his field notes on these frogs in panamá, to arthur c. echternacht for reading part of the manuscript, and to john d. lynch for many suggestions and helpful criticisms. the illustrations were executed by david m. dennis. materials and methods for the purposes of the present study i examined 1383 preserved specimens, 50 skeletons, and 9 lots of tadpoles. external characteristics used in the analysis of variation are those currently employed in the study of anuran systematics. twelve measurements and six proportions were taken in the manner described by duellman (1956). only the most important references are given in the synonymies, except those of the two subspecies of _hyla staufferi,_ which are more nearly complete. the taxonomic history of each frog is discussed under _remarks_ in each account. the cranial osteology was studied by using skeletons and cleared and stained specimens of all species. developmental stages of tadpoles were determined from gosner's (1960) table. personal field work in central america in the summer of 1966 provided an opportunity to make observations on the ecology, calling sites, and color in life; these data were supplemented by field notes from, and discussions with, dr. william e. duellman and charles w. myers. the mating calls of the frogs were recorded in the field on magnemite and uher tape recorders by dr. duellman in the course of his work on the hylid frogs of middle america--supported by grants from the national science foundation (g-9827 and gb-1441). these recordings, plus those borrowed from other institutions, provided 50 tapes for analysis of the mating calls. the calls were analyzed on a vibralyzer (kay electric company). the hyla rubra group _definition._--the species forming the group are small to moderate-sized tree frogs (maximum snout-vent length of males of various species 20-49 mm.), distinguished from other groups in the genus _hyla_ as follows: brown, grayish brown, or yellowish tan above; thighs plain, marbled with dark brown, or having vertical bands; vocal sac single, median, subgular; snout flat, protruding, rounded or pointed; webbing between fingers reduced or absent; web between first and second toes reduced to fringe on second toe, rest of toes about half webbed; tarsal fold reduced or absent; shanks robust; inner metatarsal tubercle larger than outer; prevomerine teeth on transverse ridges between small to large sized choanae; skull generally longer than wide; nasals large (length more than 40 per cent total length of skull) and having pointed maxillary processes; maxillary bearing small ventromedial palatine process; quadratojugal slender, always joined to maxillary by bony suture; auditory region of proötic slender and short; delicate spatulate columella ventral to crista parotica, broad basally, compressed anterolaterally, slightly rounded distally; anterior arm of squamosal extending about half distance to maxillary; sphenethmoid wider than long; frontoparietal fontanelle present or absent; prevomerine, premaxillary, and maxillary teeth present; prevomer with two lateral processes forming incomplete bony margin to internal nares; tadpoles having pointed xiphicercal tail, snout short, rounded; 2/3 tooth rows; dorsal fin deeper than ventral fin; sinistral spiracle; short dextral anal tube not reaching edge of ventral fin; mating calls consisting of single long note or series of short notes. _composition._--this group contains about 24 currently recognized species, most of which occur in brazil. only five species--_boulengeri,_ _elaeochroa_, _foliamorta_, _rubra_, and _staufferi_ with two subspecies--occur in central america. _hyla boulengeri_ and _rubra_ are widespread in south america, and _foliamorta_ occurs in colombia, whereas the other species are known only from middle america. _distribution._--the species of the _hyla rubra_ group range from the lowlands of northern argentina and bolivia to southern tamaulipas and guerrero, méxico. _comments._--in central america two subgroups of species can be recognized. _hyla boulengeri_ and _h. foliamorta_ are distinctive in the large size of adults (snout-vent lengths 41-49 mm.); both have prominent bars on the thighs, a well-defined interorbital triangular mark, blotches or spots dorsally, and large choanae. _hyla elaeochroa,_ _h. rubra,_ and _h. staufferi_ are smaller (snout-vent lengths 29-40 mm.); they have the thighs weakly barred or vermiculate anteriorly and posteriorly or unmarked, an ill-defined interorbital triangular mark, linear markings dorsally, and small choanae. key to species and subspecies 1. larger frogs (males to 49 mm. snout-vent length); thighs strongly barred; supratympanic fold black; dorsum blotched or spotted 2 smaller frogs (males to 40 mm. snout-vent length); thighs weakly barred or plain; supratympanic fold pale brown; dorsum usually having linear pattern 3 2. dorsum tuberculate; snout subacuminate; vocal sac flecked with brown; tarsal fold rudimentary; web absent between fingers; black spots absent in scapular region _h. boulengeri_ dorsum smooth; snout pointed; vocal sac dark gray; tarsal fold absent; trace of web between fingers; two or more elongate dark spots in scapular region _h. foliamorta_ 3. snout-vent length more than 30 mm.; tympanum 2/3 to 3/4 diameter of eye; prevomerine elevations about size of choanae 4 snout-vent length less than 30 mm.; tympanum less than 1/2 diameter of eye; prevomerine elevations smaller than choanae 5 4. thighs mottled posteriorly; discs on fingers about 1/2 size of tympanum; faint dark line from nostril to eye _h. rubra_ thighs faintly barred or plain posteriorly; discs on fingers about size of tympanum; distinct dark line from nostril to eye _h. elaeochroa_ 5. dorsum brown with irregular dorsolateral stripes and interrupted paravertebral stripes; two transverse bars on shanks; interorbital bar present _h. staufferi staufferi_ dorsum gray with complete dorsolateral and paravertebral stripes; longitudinal stripe on shank; interorbital bar absent _h. staufferi altae_ key to known tadpoles 1. entire lower beak black; beaks moderate-sized, serrate; dorsal fin high, extending to middle of back 2 no more than half of lower beak black; beaks small, finely serrate; dorsal fin lower, barely extending onto body 3 2. papillae present only laterally _h. boulengeri_ papillae present laterally and ventrally _h. foliamorta_ 3. distinct brown stripe from nostril to eye; two stripes below eye, _h. elaeochroa_ faint stripe from nostril to eye; no stripe below eye _h. staufferi_ accounts of species and subspecies _hyla boulengeri_ (cope) _scytopis boulengeri_ cope, bull. u.s. natl. mus., 32:12, december 1, 1887 [holotype.--usnm 13974, from "nicaragua"; j. a. mcniel, collector]. _hyla boulengeri:_ günther, biologia centrali-americana, reptilia and batrachia, p. 267, june 1901. noble, bull. amer. mus. nat. hist, 38:339, june 1918. taylor, univ. kansas sci. bull., 35:856, july 1, 1952. _diagnosis._--size large (male to 49 mm., female to 53 mm.); skull as long as wide; frontoparietal fontanelle present; snout subacuminate; canthus not pronounced; choanae large; tongue cordiform, slightly longer than broad; interorbital triangle tubercular; skin on dorsum tuberculate; tarsal fold reduced or absent; thighs, shanks, and tarsi boldly barred with dark brown and pale yellow-green in life. _description._--head flattened, longer than wide; snout projecting beyond lower lip; loreal region oblique; canthus not pronounced; length of eye less than interorbital distance; tympanum large, about 70 per cent of diameter of eye; interorbital triangle distinct; arms short; fingers lacking web; palmar tubercle tripartite; subarticular tubercles distinct; long tubercle on base of first finger; discs truncate; legs long; tarsal fold reduced or absent; inner metatarsal tubercle rounded, larger than outer, both elevated; subarticular tubercles distinct; one phalanx free of web on second, third, and fifth toes, three free on fourth toe (fig. 1a and b); skin tuberculate on dorsum, less so on flanks; skin of belly granular, that on chest and throat weakly granular; tongue cordiform, longer than wide, free and notched behind; vocal slits large, lateral to tongue. [illustration: fig. 1. a and b.--hand and foot of _hyla boulengeri_ (ku 102173), × 3. c and d.--hand and foot of _hyla s. staufferi_× (ku 57790), × 6] in life, dorsum tan or brown with dark spots on snout, head, and scapular region; interorbital triangle and blotch posteriorly on dorsum dark brown; flanks pale green; groin pale green or orange, mottled with dark brown; thighs tan or brown above with dark transverse bars on anterior and posterior surfaces; spaces between bars green or orange; inner surfaces of shanks and outer surfaces of tarsi brown and orange; foot brown above; forelimbs brown and pale green above, weakly barred; belly creamy white with scattered brown spots; vocal sac creamy white flecked with brown; lower jaw brown with white spots on lips (pl. 1a). in preservative, head and dorsum dark brown with triangular spot between eyes; dark spots on head and scapular region and dark brown blotch posteriorly on dorsum; flanks creamy white with brown spots; groin creamy white mottled with dark brown; thighs brown above with dark brown transverse bars on anterior and posterior surfaces; inner surfaces of shanks and outer surfaces of tarsi barred with pale brown; dorsal surface of foot mottled brown and creamy white; ventral surface of foot and toes pale brown; forelimbs faintly barred with pale brown; belly white with a few pale brown spots; vocal sac flecked with pale brown; lower jaw marked with small white spots on lips. _variation._--geographic variation is evident in the snout-vent length, tibia length, and foot length, all in relation to snout-vent length, and the relative size of the tympanum to the eye (table 1). the largest specimens are from the humid pacific lowlands of costa rica; individuals from the caribbean lowlands of costa rica, canal zone, and south america are smaller. a general trend for increase in size extends from south america to the pacific lowlands of costa rica. most variation in color does not seem to be correlated with geography; color variation is nearly as great within a given population as between separated populations. however, most specimens from rincón de osa, puntarenas province, costa rica, are dusky brown, but a few are darker. in comparison with specimens from the caribbean lowlands of central america, specimens from the pacific slopes in costa rica have a darker interorbital triangle. in some specimens from the latter area rugosities are present on the borders of the interorbital triangle, on the snout, on the upper eyelid, and on the heel. specimens from the caribbean lowlands are less tuberculate, and most individuals from there lack rugosities on the tarsus. living individuals from puerto viejo, heredia province, costa rica, and from the canal zone, panamá, are brown above with a metallic green tint. rugosities are present on the posterior edges of the forelimbs in some specimens from throughout the range. in most respects, specimens from the canal zone resemble those from the caribbean lowlands of costa rica more than they resemble those from the pacific lowlands of costa rica, but some individuals from the canal zone are less metallic above and have small white spots dorsally on the body, head, and limbs. a moderate amount of color change from night to day has been noted. at night, a male from puerto viejo, heredia province, costa rica, was grayish tan above with slightly darker markings; the flanks were pale yellowish green. by day, the dorsum was brown with darker markings, and the throat was pale gray with white flecks; the rest of the venter was creamy white. the groin was pale green with black mottling; the anterior and posterior surfaces of the thighs and inner edges of the tarsi were greenish yellow with black bars. table 1.--geographic variation in size and proportions in males of _hyla boulengeri_. (means in parentheses below observed ranges.) ========================================================================= | | snout-vent| tibia | | | | length | length/ |tympanum/|foot length/ locality | n | (mm.) | snout-vent| eye | snout-vent --------------------+----+-----------+-----------+---------+------------costa rica: tilarán | 23 | 37.4-48.7 | 0.52-0.58 |0.62-0.76| 0.39-0.45 | | (43.8) | (0.55) | (0.71) | (0.42) | | | | | costa rica: rincón | 10 | 41.4-46.1 | 0.54-0.60 |0.68-0.80| 0.40-0.45 de osa | | (44.0) | (0.57) | (0.74) | (0.43) | | | | | costa rica: alajuela| 13 | 35.6-43.1 | 0.55-0.60 |0.63-0.78| 0.41-0.46 province | | (39.8) | (0.57) | (0.69) | (0.44) | | | | | costa rica: puerto | 25 | 37.5-42.9 | 0.51-0.62 |0.63-0.79| 0.38-0.46 viejo | | (41.6) | (0.55) | (0.71) | (0.43) | | | | | costa rica: suretka | 9 | 38.7-42.0 | 0.56-0.58 |0.53-0.72| 0.35-0.45 | | (41.0) | (0.56) | (0.62) | (0.42) | | | | | panamá: canal zone | 16 | 36.7-42.9 | 0.52-0.58 |0.70-0.78| 0.40-0.44 | | (39.0) | (0.54) | (0.74) | (0.42) | | | | | venezuela: santomé | 4 | 35.5-40.9 | 0.45-0.48 |0.63-0.67| 0.36-0.40 | | (38.5) | (0.46) | (0.65) | (0.38) table 2.--comparison of mating calls in the _hyla rubra_ group. (means in parentheses below observed ranges.) ============================================================================ | |notes| | | | major frequencies | | per |duration| pulses|fundamental| (cps) | |call | of note| per | frequency |--------------------species | n |group| (sec.) | second| (cps) | lower | upper ---------------+---+----+---------+-------+-----------+---------+----------h. boulengeri | 8 | 1 | 0.24 | 80-120| 70-74 |1400-1820|2520-3182 | | | 0.47 | (101)| (71) | (1611) | (2840) | | | (0.35) | | | | | | | | | | | h. foliamorta | 7 | 1 | 0.23 | 50-60 | 52-61 | 912-1026|2736-3477 | | | 1.86 | (51) | (56) | (918) | (3055) | | | (0.69) | | | | | | | | | | | h. elaeochroa |15 | 2-95| 0.12 | 40-50 | 48-65 |1254-1586|2562-3477 | | (19)| 0.24 | (42) | (57) | (1499) | (2911) | | | (0.17) | | | | | | | | | | | h. s. staufferi|18 | 2-77| 0.13 |100-130| 96-130 |1582-1872|1962-3744 | | (23)| 0.23 | (120) | (106) | (1743) | (3056) | | | (0.18) | | | | | | | | | | | h. s. altae | 7 | 2-22| 0.14 |110-130| 104-117 |1853-2106|3379-4056 | | (11)| 0.18 | (120) | (112) | (2008) | (3775) | | | (0.15) | | | | _cranial osteology._--the skull of _hyla boulengeri_ is as long as it is wide, and is flat; the premaxillary is small and bears 13 to 17 teeth (mean for 6 specimens, 14.9). the alary processes of the premaxillaries are widely separated, concave posteriorly, and vertical. ventrally, the premaxillary is connected to the prevomer by bony tissues. the maxillary is slender and bears 70 to 91 teeth (mean for 6 specimens 78.1). the pars facialis of the maxillary is laterally convex and about four times as high as the pars dentalis. the nasal is large (its length about 40 per cent of total length of skull), and pointed anteriorly and posteriorly in dorsal view. the nasals are separated anteromedially by the cartilaginous septum nasi. one or two protuberances are present on the midlateral concavity of the nasal. posteriorly, the nasal overlaps the sphenethmoid and articulates with the palatine. dorsally the sphenethmoid is large, pentagonal, and completely ossified. the frontoparietal is elongate, smooth, and bears a small supraorbital process on the anterior edge of the orbit. a keyhole-shaped frontoparietal fontanelle is present; the fontanelle is narrow anteriorly and wide posteriorly. the bony part of the proötic is separated dorsally from the squamosal by the cartilaginous crista parotica. the squamosal is small, its anterior arm slender and pointed. the posterior arm of the squamosal is pointed terminally and articulates with the proötic medially. the prevomer is large and elongate. anteriorly the prevomer is connected to the maxillary-premaxillary articulation; posteriorly, the prevomer is separated from the sphenethmoid by cartilage. each prevomer bears six to nine teeth. the palatine is present and edentate. the anterior end of the parasphenoid is broad (less pointed than in _hyla foliamorta_). the pterygoid is slender and well developed. _natural history._--_hyla boulengeri_ inhabits humid lowland tropical forests and breeds in temporary ponds. clasping pairs and gravid females were observed at puerto viejo, heredia province, costa rica, on june 21, 1966. males were calling from depressions in decaying logs and stumps, in forked stems, and from leaves of broad-leafed plants near the pond. males were observed in late july and early august calling from _calathea_ and _heliconia_ leaves at the edge of a pond in the wet forest of the osa peninsula. william e. duellman informed me that he collected calling males in january at el real, darién, and in march at almirante, bocas del toro, panamá. taylor (1952) found calling males in june at turrialba, cartago province, costa rica, and dunn (1931a) observed males calling in july, november, and december in panamá. gravid females have been found from april to august. breeding activities of _hyla boulengeri_ always seem to be associated with temporary ponds; in central america breeding apparently takes place throughout most of the year. the mating call of _hyla boulengeri_ consists of one short, moderately low-pitched note. each note has a duration of 0.24 to 0.47 second and is repeated at intervals of one second to several minutes. the notes have 80 to 120 pulses per second, a fundamental frequency of about 70 cycles per second, and a dominant frequency of 2,840 cycles per second (table 2, pl. 3a). the eggs are deposited in a mass in the water. no information is available concerning early development. tadpoles in advanced stages of development were found in a temporary pond at rincón de osa, puntarenas province, costa rica. the pond was about 10 cm. deep, had a muddy bottom and lacked vegetation. three recently metamorphosed young were found in mid-august, 1966, on grass at the edge of another temporary pond in the forest. _tadpoles_--twelve tadpoles are available. these were collected at rincón de osa, puntarenas province, costa rica. the maximum size represented is 34.0 mm., total length (stage 42 of development). a typical tadpole in stage 36 of development (ku 104295) has a body length of 12.0 mm., tail length of 20.0 mm., and total length of 32.0 mm. other characters are as follows: depth of tail equal to length of body; body deeper than wide; distance between eye and nostril equal to that between nostril and tip of snout; mouth anteroventral, upper and lower lips bare; papillae present laterally; tooth rows 2/3; upper rows about equal in length; first upper row slightly, and second upper row widely, interrupted medially; lower rows about equal in length, shorter than upper rows; third lower row containing 5-10 large teeth; beak strong, serrate; spiracle nearer anus than eye; anal aperture not extending to border of ventral fin; caudal musculature slender posteriorly, extending to tip of pointed tail; dorsal fin extending to middle of body, slightly deeper than ventral fin; posterior three fourths of tail spotted; rest of tail and body gray-brown or transparent; hindlimbs flecked or spotted with brown (table 3, fig. 2a and 3a). table 3.--sizes of tadpoles of _hyla boulengeri_ in relation to developmental stages. (means in parentheses below observed ranges; measurements in mm.) ======================================================= stage | n | body length | tail length | total length --------+---+-------------+-------------+------------- 30 | 1 | 11.0 | 22.2 | 33.2 | | | | 35 | 1 | 11.0 | 12.0 | 23.0 | | | | 36 | 3 | 9.5-12.0 | 20.0-21.5 | 31.0-32.0 | | (11.2) | (20.5) | (31.7) | | | | 38 | 2 | 11.5 | 22.0 | 33.5 | | | | 42 | 2 | 10.5-13.0 | 21.0-22.0 | 32.5-34.0 | | (11.8) | (21.5) | (33.3) | | | | 44 | 2 | 14.0-15.0 | 8.0-15.0 | 22.0-30.0 | | (14.5) | (12.5) | (26.0) | | | | 46 | 1 | 15.0 | 15.0 | a recently metamorphosed young has a snout-vent length of 15 mm.; the head is as long as wide, the eyes are prominent; the limbs are weakly barred; the skin is rugose above and granular below. the venter is immaculate; the dorsum and limbs are gray-brown in preservative (pale green in life). the interorbital space, supratympanic fold, and scapular region are darker than the rest of the body; the fingers lack webbing; the webbing on the foot is the same as in adults; small metatarsal tubercles are present, but the tarsal fold is absent. [illustration: fig. 2. tadpoles of (a) _hyla boulengeri_ (ku 104295) and (b) _hyla elaeochroa_ (ku 104134), × 3.] [illustration: fig. 3. mouthparts of tadpoles of (a) _hyla boulengeri_ (ku 104295) and (b) _hyla elaeochroa_ (ku 104134), × 25.] _remarks._--cope (1887:12) described _scytopis boulengeri_ from nicaragua. günther (1901:267) placed _boulengeri_ in the genus _hyla_, and stated that cope possibly placed _boulengeri_ in the genus _scytopis_ on the supposition that it had an accumulation of "sebaceous glands" above the tympanum. noble (1918:339) redescribed _hyla boulengeri_ on the basis of three specimens from zelaya province, nicaragua, and noted that the glands were not prominent in any of the specimens. duellman (1956:8) showed that _scytopis hebes_ (generotype of _scytopis_ by monotypy) is a phrynohyas, and thus placed _scytopis_ cope, 1862, in the synonymy of _phrynohyas_ fitzinger, 1843. dunn and emlen (1932:25) placed _hyla lancasteri_ barbour in the synonymy of _hyla boulengeri_; the former was known solely from one juvenile. they made no qualifying statements, but probably they were impressed by the strongly barred thighs, a coloration known among central american hylids at that time only in _hyla boulengeri_ (duellman, 1966a:271). taylor (1952:856) followed dunn and emlen with reservation and noted some differences. duellman (1966a:271) showed that the holotype of _lancasteri_ was a juvenile of a species subsequently named as _hyla moraviaensis_ by taylor (1952:865). in central america, _hyla boulengeri_ can be confused only with _hyla foliamorta;_ the latter is restricted to central and eastern panamá and northern colombia. the snout of _foliamorta_ is more pointed and protruding, and the vocal sac is darker than in _boulengeri_; the groin of _foliamorta_ usually is creamy white, whereas _boulengeri_ usually has a dark spot. the skulls differ in that _boulengeri_ has a frontoparietal fontanelle, the prevomer is larger and elongate, anteriorly connected to the premaxillary, and posteriorly separated from the sphenethmoid by cartilage; _foliamorta_ lacks a fontanelle, the prevomer is smaller, anteriorly separated from the premaxillary by cartilage, but connected by a bony suture to the sphenethmoid. the mating call of _boulengeri_ differs by having shorter notes, twice as many pulses per second, a higher fundamental frequency, and more closely approximated major frequencies than does that of _foliamorta_. _hyla boulengeri_ need not be compared in detail with the other central american members of the _hyla rubra_ group, because all of them are smaller and have shorter snouts, smoother skin, and dissimilar color patterns. _distribution._--in central america _hyla boulengeri_ inhabits the forested lowlands in locally humid areas in guanacaste province, costa rica, and in the humid golfo dulce region of costa rica; it occurs on the carribbean lowlands from central nicaragua to south america, where it ranges to guyana and ecuador. the highest elevations where _h. boulengeri_ has been found are 620 meters at turrialba, cartago province, and 700 meters at tilarán, guanacaste province, costa rica (fig. 4). _specimens examined._--costa rica: _alajuela_: 9 km n ciudad quesada, near la florencia, usc 8059 (4); 18 km n florencia, usc 2624; laguna monte alegre, ku 64334; las playuelas, 11 km s los chiles, usc 7216, 7217 (2), 7219; 3 km ne muelle del arenal, usc 2644 (5). _cartago_: turrialba, ku 24741. _guanacaste_: 7 km n liberia, usc 8096 (2), 8138 (6); 13.6 km n liberia, usc 8151, 8171 (2); 20.5 km s liberia, usc 8205; taboga, 20 km se las cañas, ku 102170, usc 7166; 4 km ne tilarán, usc 8023; 6 km ne tilarán, usc 523 (3), 6262, 7019. _heredia_: puerto viejo, ku 64323-7 (skeletons), 104351-3 (skeletons), 64330-3, 103592-620; 1 km ne puerto viejo, ummz 126042; 1 km s puerto viejo, ku 84983-4 (skeletons), 86317-22, 87774 (skeleton); 4.2 km w puerto viejo, ku 64329, 64328 (skeleton). _limón_: mountain cow creek, near banano, ku 37031, 41067 (skeleton); 3 km s río tortuguero, amnh 69057; suretka, ku 36482-8, 36699. _puntarenas_: 4.8 km s bahía rincón on nw side río rincón, usc 705; parrita, usc 6163; 4.5 km w rincón de osa, ku 102177-9, 104295-6 (tadpoles); 6 km sw rincón de osa, ku 102171-6; 4.4 km nw villa neilly, usc 8003; 10.5 km wnw villa neilly, ku 64321. _san josé_: 21 km wsw san isidro el general, ku 34104-6. [illustration: fig. 4. map showing locality records for _hyla boulengeri_ (circles) and _h. foliamorta_ (dots).] panamá: _bocas del toro_: 3.2 km w almirante, ku 95978. _canal zone_: barro colorado island, fmnh 13379; near clayton reservation, uimnh 42000; 2.6 km sw fort kobbe, ku 95977; miraflores locks, amnh 69764-5; summit, amnh 73445, ku 97777, 101540-9, 104350 (skeleton). _colón_: río gatuncillo, near nuevo san juan, ku 95976. _darién_: el real, ku 80451-3. _hyla foliamorta_ fouquette _hyla foliamorta_ fouquette, herpetologica, 14:125, april 25, 1958 [holotype.--tnhc 23109, 11 km. nw miraflores locks, canal zone, panamá; m. j. fouquette, jr. collector]. _diagnosis._--size medium (male to 43 mm., female to 41 mm.); skull longer than wide; frontoparietal fontanelle absent; snout acuminate, projecting; interorbital triangle bordered by white lines; scapular region having two or more elongate spots; dorsum smooth; vocal sac dark gray; groin creamy white; traces of web between fingers. _description._--head flattened, longer than wide; snout flat, pointed, protruding beyond lower lip; loreal region slightly concave; canthus moderately prominent; eyes smaller than interorbital space; tympanum distinct, 55 to 75 per cent of diameter of eye, smaller than internarial space; arms short; fingers having rudimentary webs; median palmar tubercle tripartite; inner palmar tubercle on base of first finger flat; subarticular tubercles distinct; discs of fingers smaller than diameter of tympanum; legs long; tarsal fold lacking; inner metatarsal tubercle larger than outer; one phalanx free on second, third, and fifth toes, two and one half phalanges free on fourth toe; narrow fringe continuing from web to discs of toes; discs of toes about the size of those on fingers; skin smooth on dorsum and flanks, that on belly and posterior part of thighs granular; tongue oval, longer than wide; vocal slits oblique, about one half length of tongue. in life, dorsum pale tan to pale reddish brown with irregular reddish brown markings; small dark spots on head; distinct dark brown triangular mark between eyes, bordered by thin white lines; apex of triangle always directed backward; supratympanic fold with black edge; scapular region having two to five small, elongate black spots; belly creamy tan with small brown spots; vocal sac uniformly dark brown with scattered creamy tan flecks; upper jaw dark brown; limbs creamy white below with scattered brown spots; groin marked with small brown spots in some specimens; anterior and posterior surfaces of thighs yellow-orange with three distinct black blotches; two dark bands on upper surface of shanks; webbing of feet yellowish tan with brown mottlings (pl. 1b). in preservative, dorsum brown or gray with darker markings; interorbital triangle distinct, bordered by white lines; supratympanic fold with black edge; two or more small elongate black spots in scapular region; belly white with numerous brown flecks; edge of upper lip dark brown; vocal sac dark gray; undersides of limbs creamy white; groin creamy white with or without brown spots; anterior and posterior surfaces of thighs having three black blotches separated by creamy white spaces; shanks having two brown bands; webbing of feet mottled with brown. _variation._--twenty-eight breeding males from the area between chepo and tocumen, panamá, have snout-vent lengths of 39.0 mm. to 46.0 mm. (mean 42.5 mm.). in these specimens, the ratio of the tibia length to the snout-vent length is 0.54 to 0.61 (mean, 0.57); the ratio of the diameter of the tympanum to that of the eye is 0.55 to 0.75 (mean, 0.67). one female has a snout-vent length of 41.0 mm., tibia/snout-vent length ratio of 0.57, and tympanum/eye ratio of 0.76. two to five (usually three) elongate black spots are present in the scapular region in different individuals. the flanks in some are spotted with brown; in others they are creamy white. a small black spot is present in the groin of some specimens. usually two to four blotches are present on the anterior and posterior surfaces of the thighs; in some specimens the blotches are reduced to small spots. one or two brown spots are present proximally on the shanks in most specimens. in some individuals tuberculations are scattered on the head and in the tympanic and scapular regions, but the dorsum is smooth in most specimens; the belly is creamy white flecked with brown. _cranial osteology._--the skull of _hyla foliamorta_ is flat and longer than it is wide. the premaxillary is small and bears 13 to 16 teeth (mean for 2 specimens, 14.8). the alary process of the premaxillary is vertical and concave posteriorly. ventrally, the premaxillary is completely separated from the prevomer by cartilage. the maxillary is slender; each bears 77 to 84 teeth (mean for 2 specimens, 81). the pars facialis of the maxillary is laterally convex and less than three times the height of the pars dentalis. the nasal is large and pointed anteriorly and posteriorly in dorsal view. the length of the nasal comprises about 40 per cent of the total length of the skull. the nasals are separated anteromedially by the cartilaginous septum nasi. one protuberance is present on the midlateral concavity of the nasal. posteriorly, the nasal overlaps the sphenethmoid; posterolaterally the nasal articulates with the palatine. the sphenethmoid is completely ossified and pentagonal in dorsal view. the frontoparietal is elongate, without a pronounced anterior supraorbital process. the frontoparientals are sutured medially throughout their lengths; the frontoparietal fontanelle is absent. the bony part of the proötic is narrowly separated dorsolaterally from the squamosal by the cartilaginous crista parotica. the squamosal is large; the anterior arm is pointed. the posterior arm of the squamosal is broad, rounded terminally, and articulates with the proötic medially. the prevomer is short and separated anteriorly from the premaxillary and maxillary by cartilage. the posterior margin of the prevomer has a bony articulation with the sphenethmoid. each prevomer bears five to seven teeth. the palatine is small and edentate. the anterior end of the parasphenoid is narrow (more pointed than in _hyla boulengeri_). the pterygoid is slender and well developed (fig. 5a). [illustration: fig. 5. dorsal views of the skulls of (a) _hyla foliamorta_ (ku 77687) and (b) _h. elaeochroa_ (ku 68289), × 3.] _natural history._--_hyla foliamorta_ inhabits lowland forests in eastern panamá and breeds in temporary ponds. males have been observed calling from grasses, bushes, and emergent vegetation near temporary ponds and ditches along roads. william e. duellman informed me that he found a breeding congregation of this species in june near chepo, panamá, where males were calling from spiny palms at the edge of a woodland pond. fouquette (1958) found calling males in may, august, and september near miraflores locks, canal zone. calling stations vary from one to two meters above ground. no clasping pairs have been found; only one female is known (ku 101589, from 8 km ne tocumen, panamá); this gravid individual was collected in early june. the mating call of _hyla foliamorta_ consists of one pulsed, low-pitched, moderate trill of about o.5 second duration. each note is repeated at intervals of 5 seconds to a few minutes. the notes have about 50 pulses per second, a fundamental frequency of 56 cycles per second and a dominant frequency of about 3000 cycles per second (table 2, pl. 3b). egg deposition sites are unknown. no information is available concerning early development, and little is known about the breeding season of _hyla foliamorta_. probably its breeding activities are restricted to the rainy months. _tadpoles._--eight tadpoles were collected from a weedy temporary pond near chepo, panamá, in early june. a typical tadpole in stage 35 of development (ku 104244) has a body length of 9.5 mm., tail length of 25.0 mm., and a total length of 34.5 mm. other characters are as follows: depth of tail equal to length of body; body deeper than wide; distance between eye and nostril equal to distance between eye and spiracle; mouth anteroventral; median part of upper lip bare; rest of lip having one row of papillae; a few other rows of small papillae at corners of mouth; tooth rows 2/3; first upper row entire, second upper row interrupted medially, shorter than first; lower rows shorter than upper rows, third shortest; beak moderately robust; spiracle nearer eye than anus; anal tube short, aperture not extending to border of ventral fin; caudal musculature slender, extending to tip of pointed tail; dorsal fin extending onto body (table 4). table 4.--sizes of tadpoles of _hyla foliamorta_ in relation to developmental stages. (means in parentheses below observed ranges; measurements in mm.) ======================================================= stage | n | body length | tail length | total length --------+---+-------------+-------------+------------- 25 | 2 | 5.0-5.2 | 8.0-8.5 | 13.0-13.7 | | (5.1) | (8.3) | (13.4) | | | | 26 | 3 | 7.0-7.5 | 12.0-12.4 | 17.0-19.5 | | (7.2) | (12.1) | (18.6) | | | | 28 | 2 | 6.5-7.0 | 18.0 | 25.0 | | (6.8) | | | | | | 35 | 1 | 9.5 | 25.0 | 34.5 in life, yellow above, white below; caudal fin greenish yellow with black or gray reticulations; dark line from snout to eye; dark spot behind eye; tail unpigmented except for fine dark reticulations. in preservative body creamy white, transparent below with dark pigment above in some specimens. _remarks._--_hyla foliamorta_ can be confused only with _hyla boulengeri_. the differences between adults of these species were discussed in _remarks_ on _h. boulengeri_. the tadpoles of _foliamorta_ have labial papillae on the lower lip and a stripe between the eye and the tip of the snout. by comparison the tadpoles of _boulengeri_ have a bare lower lip and no stripe between the eye and the tip of the snout. _distribution._--_hyla foliamorta_ inhabits the subhumid pacific lowlands (elevations of less than 100 meters) of central panamá and caribbean lowlands of northern colombia (fig. 4). _specimens examined._--panamá: _panamá_: 3 km wsw chepo, ku 77164-9, 101573-5, 104243-4 (tadpoles); 6 km wsw chepo, ku 77170, 101576-8; 1.5 km sw naranjal, ku 77171, 77687 (skeleton); 2 km n tocumen, ku 101579-83, 104349 (skeleton); 8 km ne tocumen, ku 101584-92. no specific locality: tnhc 24401. _hyla rubra_ laurenti _hyla rubra_ laurenti, synopsis reptilium emendatum, p. 35, 1768. daudin, hist. nat. rainettes grenouilles crapauds, ii:26, 1802. daudin, hist. nat. particuliere reptiles, 8:53, 1803. günther, catalogue batrachia salientia brit. mus., p. 110, 1859. boulenger, catalogue batrachia salientia s. ecaudata, p. 403, february 1, 1882. dunn, occas. papers, boston soc. nat. hist., 5:413, october 10, 1931. _hyla elaeochroa_ (part): dunn and emlen, proc. acad. nat. sci. philadelphia, 84:25, march 22, 1932. _diagnosis._--size medium; skull longer than wide; frontoparietal fontanelle absent in adults; snout subovoid; choanae rounded; dorsal stripes present; black vermiculations on posterior surfaces of thighs. _description._--head flat, longer than wide; snout long, subovoid, slightly protruding beyond lower lip; loreal oblique, concave; canthus rounded, indistinct; diameter of eye about equal to interorbital space; tympanum large, about three fifths diameter of eye, smaller than internarial distance; supratympanic fold indistinct; arms short; fingers free of webs; subarticular tubercles distinct; median palmar tubercle large, bifid; inner palmar tubercle on base of first finger flat, elongate; disc of third finger about one half diameter of tympanum; legs moderately long; tarsal fold absent; inner metatarsal tubercle distinct, oval; toes about half webbed; web on fourth toe extending to disc; discs of toes about size of those on fingers; skin smooth above with small granules on head and in scapular region in some specimens; skin on flanks, throat, belly, and lower surfaces of thighs granular; tongue oval, longer than wide, not free behind; choanae small, oval; vocal slits long, lateral to tongue. in preservative, dorsum pale brown with darker dorsolateral stripes; narrow dark brown line from nostril to eye; groin, anterior surface of thighs, and posteroventral surfaces of shanks creamy tan with dark brown vermiculations; white spots present on thighs in some specimens; throat flecked with brown; belly creamy white or gray. _remarks._--the taxonomic history of _hyla rubra_ laurenti is confused. seba (1734:70) illustrated and diagnosed a frog for which he used the name _ranula, americana, rubra_. linnaeus (1758:213) considered seba's frog to be a variety of _hyla arborea_. laurenti (1768:35) apparently examined the same individual that seba called _ranula, americana, rubra_. for this specimen, laurenti used the binominal _hyla rubra_ and provided a brief diagnosis. the type locality was given as america. daudin (1802:26) redescribed the same specimen(s?) treated by seba and laurenti and provided a fairly good description and figures. daudin restricted the type locality to surinam and indicated that marin de baize was the probable collector. daudin (1802:26 and 1803:53) neglected to consider laurenti's work, but he applied the same name used by laurenti. most authors have credited _hyla rubra_ to daudin, but rivero (1961:120) noted that _hyla rubra_ laurenti, 1768, has priority over _hyla rubra_ daudin, 1802. since both laurenti and daudin worked on seba's material, it is reasonable to assume that daudin redescribed the same frog that was named by laurenti; this was not an uncommon practice in the early nineteenth century. thus i conclude that _hyla rubra_ daudin, 1802, is a junior synonym of _hyla rubra_ laurenti, 1768. dunn (1931a:413) first reported _hyla rubra_ from central america; he recorded the species from the canal zone and san pablo, panamá. i have examined the material of _hyla rubra_ from panamá deposited in various museums. most of the specimens are faded, discolored, and do not have distinct brown vermiculations on the thighs. the specimens seem to be more like _hyla rubra_ than any of the other species in the _rubra_ group. the presence of oval choanae and a tympanum larger than the largest finger disc separate these specimens from _hyla elaeochroa_, a species with which _rubra_ has been confused. _hyla elaeochroa_ does not occur in the canal zone or eastern panamá. all museum specimens from nicaragua, costa rica, and western panamá that have been called _hyla rubra_, plus those mentioned by dunn and emlen (1932:25) and dunn (1933:61) are _hyla elaeochroa_. the taxonomic status of the many south american populations referred to _hyla rubra_ and of other populations now recognized as different species is not clear at the present time. considerable variation in external characters and in cranial features has been observed in south american _rubra_. a review of the taxonomy of these populations is beyond the scope of this paper. possibly the central american specimens herein referred to _rubra_ will ultimately be found to be specifically distinct from those in surinam. since i have no osteological material from central america, i have been unable to describe the cranium in this account. furthermore, i have no data on the ecology and life history of _rubra_ in central america. _distribution._--_hyla rubra_ inhabits lowland tropical forests from central-eastern panamá to northern south america and thence through lowlands east of the andes to northern argentina (fig. 6). _specimens examined._--panamá: _canal zone_: gatun, ummz 52720 (2); madden dam, fmnh 67820; no specific locality, ummz 56517 (3), usnm 37863. _colón_: cerro bruja, mcz 13248. _darién_: el real, usnm 140569-70, 140573. _panamá_: juan díaz, mcz 17973; las sabanas, mcz 17581; río trinidad, ummz 64003; san pablo, mcz 1398-9. _hyla elaeochroa_ cope _hyla elaeochroa_ cope, jour. acad. nat. sci. philadelphia, 8:105, 1876 [holotype.--usnm 30689, sipurio, limón province, costa rica; william m. gabb collector]. günther, biologia centrali-americana, reptilia and batrachia, p. 265, june 1901. taylor, univ. kansas sci. bull., 35:859, july 1, 1952. duellman, univ. kansas publ., mus. nat. hist., 17:270, june 17, 1966. _hyla quinquevittata_ cope, proc. amer. philos. soc., 23:273, april 1887 [holotype.--usnm 14187, nicaragua; j. f. bransford collector]. günther, biologia centrali-americana, reptilia and batrachia, p. 268, june 1901. noble, bull. amer. mus. nat. hist., 38:340, june 1918. _hyla rubra_ (part): dunn and emlen, proc. acad. nat. sci. philadelphia, 84:25, march 22, 1932. _hyla dulcensis_ taylor, univ. kansas sci. bull., 39:37, november 18, 1958 [holotype.--ku 32168, golfito, puntarenas province, costa rica; edward h. taylor collector]. _diagnosis._--size medium (male to 38 mm., female to 40 mm.); skull wider than long; nasals truncate anteriorly; frontoparietal fontanelle moderate in size; snout slightly protruding; tympanum about size of largest discs on fingers; dorsum marked by longitudinal stripes; dark stripe between eye and nostril; in life tan to olive-green with or without dark mark between eyes; bones greenish blue. _description._--head flat, longer than wide; snout long, rounded, protruding beyond mouth; canthus indistinct; length of eye equal to interorbital distance; loreal region not pronounced; tympanum distinct and about two-fifths diameter of eye; interorbital triangle present or absent; arms short; trace of web between fingers, extending as fringe along sides of fingers; first finger very short with small disc; other discs about size of those on toes; discs on third finger and fourth toe as large as tympanum; outer palmar tubercle moderate in size, partly bifid; inner palmar tubercle large, elongate, flat; subarticular tubercles distinct; legs moderately long; tarsal fold absent; inner metatarsal tubercle flat; outer metatarsal tubercle smaller, indistinct; subarticular tubercles moderate in size; fringe on toes to tip of disc of second toe; rest of toes about two-thirds webbed; foot length about two fifths snout-vent length; tibia length about one half snout-vent length; skin above smooth or with minute pustules; belly finely granular; ventral surfaces of thighs and areas below anus granular; skin on ventral surfaces of limbs smooth; tongue relatively large, longer than wide, barely notched behind; vocal slits elongate, lateral to tongue; choanae medium in size. in life, dorsum yellowish brown, olive green, or grayish brown with dark brown spots on snout, dark brown stripe from nostril to eye, dark yellow-brown interorbital triangle, and dark supratympanic region; generally five interrupted longitudinal dark brown stripes on dorsum (one on each flank, pair of paravertebral and one vertebral); flanks pale yellow; groin yellowish brown; thighs marked with one or two transverse yellow-brown blotches; shanks with two or three yellow-brown blotches above; spaces between blotches on thighs, shanks, tarsi, and feet yellow; brown spots on tarsi and in some specimens on feet; arm pale yellow with pale brown spots; belly creamy white having slight blue-green tint; vocal sac and chin yellow; axillary region yellow, blue-green in some specimens (pl. 2a). in preservative, head and dorsum yellowish brown; dark brown stripe from nostril to eye; dark brown spots on snout; a dark brown interorbital triangle with apex directed backward; dark brown supratympanic region; dorsal stripes same as in living individuals; flanks pale yellow with brown spots in some specimens; groin creamy white; thighs and shanks having or lacking transverse dark brown blotches; spaces between blotches creamy white or yellow-brown; arms pale yellowish brown; belly and vocal sac creamy white. _variation._--geographic variation in size and some proportions, such as the ratio of tibia length to snout-vent length and the ratio of the diameter of the tympanum to that of the eye, have been observed in this species. the largest individuals are from the golfo dulce region (samples from piedras blancas and rincón de osa), puntarenas province, costa rica. the smallest individuals are from el recreo, zelaya province, nicaragua, and from the caribbean lowlands of costa rica. the diameter of the tympanum is proportionately larger (relative to the size of the eye) in males from tilarán, guanacaste province; the tympanum is nearly as large in males from piedras blancas, puntarenas province, and puerto viejo, heredia province, costa rica. the lowest ratios occur in individuals from almirante, bocas del toro, panamá, in specimens from the caribbean lowlands of costa rica (except puerto viejo), and in those from el recreo, zelaya province, nicaragua. in general, the tympanum is proportionately larger in females than in males; the tympanum is largest in females from the pacific lowlands of costa rica (table 5). color variation has been observed in individuals from the same population, as well as in individuals from different localities, between males and females, and from night to day. in life, most individuals from the pacific lowlands of costa rica are dark tan to greenish gray above with dark brown longitudinal stripes that are entire or broken, but some specimens (mostly males) are dusky brown and lack longitudinal stripes or an interorbital triangle; females usually have the dark interorbital triangle and the stripes on the dorsum. individuals from turrialba, cartago province, costa rica, are pale olive-tan with olive-brown markings. individuals from puerto viejo, heredia province, costa rica, are uniformly yellowish brown with or without dark longitudinal stripes. specimens from el recreo, zelaya province, nicaragua, are like those from puerto viejo. males from almirante, bocas del toro, panamá, are pale brown with dark brown longitudinal stripes and an indistinct interorbital triangle. females have a distinct interorbital triangle and dark brown blotches on the thighs and shanks. by night, the dorsum usually is pale yellow, and the belly is creamy white. by day, the dorsum is dark tan; the stripes and spots are darker, and the belly is yellowish white. taylor (1952) noticed that considerable variation in color pattern occurred from night to day in individuals from turrialba, cartago province, costa rica. at night some individuals lacked a dorsal pattern, but by day many of these individuals developed dorsal stripes. _cranial osteology._--the skull of _hyla elaeochroa_ is slightly wider than it is long, and flat. the premaxillary is small and bears 10 to 15 teeth (mean for 9 specimens, 12.3). the alary process of the premaxillary is small, vertical, and slightly concave posteriorly. ventrally, the premaxillary is partially united to the prevomer by ossification. the maxillary is slender and bears 70 to 82 teeth (mean for 9 specimens, 74.3). the pars facialis of the maxillary is laterally convex and is about twice as high as the pars dentalis. the nasal is large, robust, anteriorly truncate, but pointed posteriorly in dorsal view. the nasal comprises about 45 per cent of the total length of the skull. there is an anterior cartilaginous septum nasi separating the two nasals; the latter overlap the sphenethmoid posteriorly. each nasal bears a shallow concavity in the midlateral side and lacks a maxillary process. dorsally, the sphenethmoid is wider than long, roughly pentagonal in shape; the frontoparietal is elongate, smooth, and bears a small anterior supraorbital process. the sphenethmoid and frontoparietal form the anterior margin of the frontoparietal fontanelle; the fontanelle is narrow anteriorly and wider posteriorly (fig. 5b). the entire distal surface of the proötic is in contact with the posterior arm of the squamosal. a narrow cartilaginous crista parotica is visible dorsally in some specimens. the squamosal is broad posteriorly but its anterior arm is slender and not in contact with the maxillary. table 5.--geographic variation in size and proportions in males of _hyla elaeochroa_. (means in parentheses below observed ranges.) ========================================================================== | |snout-vent| tibia | | | | length | length/ |tympanum/|foot length/ locality | n | (mm.) | snout-vent | eye | snout-vent ---------------------+----+----------+------------+---------+------------nicaragua: el recreo | 9 | 28.0-30.3| 0.51-0.57 |0.47-0.59| 0.39-0.54 | | (29.3) | (0.55) | (0.51) | (0.41) | | | | | costa rica: tilarán | 21 | 28.8-33.6| 0.47-0.57 |0.48-0.65| 0.40-0.46 | | (30.6) | (0.52) | (0.59) | (0.41) | | | | | costa rica: puerto | 22 | 26.3-32.4| 0.49-0.54 |0.48-0.65| 0.38-0.45 viejo | | (29.7) | (0.52) | (0.57) | (0.42) | | | | | costa rica: turrialba| 95 | 28.1-35.0| 0.47-0.56 |0.47-0.68| 0.37-0.46 | | (30.6) | (0.51) | (0.56) | (0.41) | | | | | costa rica: bataán, | 26 | 26.3-32.7| 0.47-0.54 |0.45-0.66| 0.36-0.44 limón, and suretka | | (30.0) | (0.51) | (0.50) | (0.41) | | | | | costa rica: piedras | 21 | 33.3-37.7| 0.50-0.54 |0.48-0.64| 0.40-0.46 blancas | | (35.2) | (0.51) | (0.57) | (0.43) | | | | | costa rica: rincón de| 24 | 31.4-35.9| 0.50-0.56 |0.45-0.61| 0.40-0.46 osa | | (34.1) | (0.53) | (0.54) | (0.43) | | | | | panamá: bocas del | 6 | 31.0-33.5| 0.49-0.54 |0.47-0.50| 0.41-0.43 toro | | (32.1) | (0.51) | (0.48) | (0.42) [illustration: plate 1 living _hyla_: (a) _h. boulengeri_ (ku 86322) and (b) _h. foliamorta_ (ku 101576), × 2.] [illustration: plate 2 living _hyla:_ (a) _h. elaeochroa_ (ku 91688), (b) _h. staufferi staufferi_ (ku 57791), and (c) _h. staufferi altae_ (ku 101688), × 2.] [illustration: plate 3 audiospectrograms and sections of mating calls of (a) _hyla boulengeri_ (ku tape no. 511) and (b) _h. foliamorta_ (ku tape no. 511) and (b) _h. foliamorta_ (ku tape no. 288).] [illustration: plate 4 audiospectrograms and sections of mating calls of (a) _hyla elaeochroa_ (ku tape no. 97), (b) _h. s. staufferi_ (ku tape no. 93), and (c) _h. staufferi altae_ (ku tape no. 502).] the prevomer is short, and broadest anteriorly. the prevomer is joined to the premaxillary by ossification. the posterior margin of the prevomer bears a narrow cartilaginous articulation with the sphenethmoid. the anterolateral and posterolateral processes of the prevomer form an incomplete bony margin to the small choanae; each prevomer bears four to seven teeth. the palatine is small, curved anteriorly and edentate. the anterior part of the parasphenoid is robust and ends in a point. the pterygoid is slender and weakly developed. _natural history._--_hyla elaeochroa_ inhabits humid lowland tropical forests in lower central america and breeds in temporary ponds. clasping pairs, gravid females, and calling males have been found mostly in june, july, and august. william e. duellman informed me that he also found males calling in mid-february, late april, and may. duellman (1967) reported detailed observations of the social organization in the mating call of _hyla elaeochroa_. the choruses in this species are initially organized, but when many individuals call, the chorus loses organization. i observed this species breeding in a temporary pond at puerto viejo, heredia province, cost rica, in late june. calling males and clasping pairs were extremely abundant within a few hours after a heavy rain. males were mostly found calling from low emergent herbs in the pond and less commonly from bushes and trees to heights of six meters above the water. calling males were also observed at ricón de osa, puntarenas province, costa rica, in late july. these breeding individuals were found in a shallow pond at the edge of a wet forest. calling stations were less than two meters in height. john d. lynch informed me that after a heavy rain in early august, he found several hundred individuals congregated in a small grassy pond less than a foot deep, at rincón de osa. males were calling from sites on grass stems a few centimeters above the water. the mating call of _hyla elaeochroa_ consists of short notes, repeated at intervals of about 0.40 second. each note has a duration of 0.12 to 0.24 second. the fundamental frequency varies from 48 to 65 cycles per second, and the notes have 40-50 pulses per second; the dominant frequency is at about 2,900 cycles per second (table 2, pl. 4a). the eggs are deposited in a mass in the water near floating vegetation. william e. duellman informed me that he observed hatchlings oriented vertically with the tip of the mouth at the surface of the water. they gradually sank to bottom, but swam back to surface again. no additional information is available concerning early development. tadpoles have been found in shallow grassy ponds in clearings and in temporary woodland ponds. _tadpoles._--three hundred and thirty-one tadpoles in various stages of development are available. thirty-five tadpoles in stage 35 have a mean body length of 8.1 mm. (8.0-9.0 mm.), tail length of 17.7 mm. (15.0-19.5 mm.), and total length of 25.9 mm. (23.0-27.5 mm.). the largest tadpole examined is in stage 40 and has a total length of 34.5 mm. (table 6). a typical tadpole, stage 35 of development (ku 104134, from puerto viejo, heredia province, costa rica), has a body length of 9.1 mm., tail length of 17.7 mm., and a total length of 26.8 mm. other characters are as follows: body depressed anteriorly; body length greater than depth of tail; internarial space as broad as interorbital distance; nostril equidistant between eye and tip of snout; eyes moderately large; mouth anteroventral and triangular; median fourth of upper lip bare; rest of lip bordered by one row of papillae; clumps of small papillae at corners of mouth; tooth rows 2/3; upper rows equal in length; second row interrupted medially; lower rows shorter than upper rows, diminishing in length; beak rather weak with small serrations; spiracle short and nearer eyes than anus; anal opening not reaching edge of ventral fin; caudal musculature attenuated distally (figs. 2b and 3b). table 6.--sizes of tadpoles of _hyla elaeochroa_ in relation to developmental stages. (means in parentheses below observed ranges; measurements in mm.) ------+---+-------------+-------------+------------- stage | n | body length | tail length | total length ------+---+-------------+-------------+------------- 24 | 2 | 4.0-4.0 | 8.5-9.0 | 12.5-13.0 | | (4.0) | (8.8) | (12.8) | | | | 25 |64 | 5.0-6.5 | 8.5-15.0 | 13.5-21.5 | | (5.7) | (11.8) | (17.6) | | | | 27 |30 | 7.0-7.5 | 13.0-16.0 | 20.0-23.0 | | (7.1) | (14.2) | (21.3) | | | | 30 |15 | 7.0-8.0 | 13.0-16.5 | 20.0-24.0 | | (7.3) | (15.0) | (22.4) | | | | 32 |30 | 7.5-8.5 | 15.0-17.0 | 22.5-25.0 | | (7.8) | (16.1) | (23.8) | | | | 35 |35 | 8.0-9.0 | 15.0-19.5 | 23.0-27.5 | | (8.1) | (17.7) | (25.9) | | | | 37 |22 | 8.5-9.5 | 16.0-22.0 | 25.0-31.0 | | (9.0) | (18.8) | (27.8) | | | | 39 |14 | 9.5-10.5 | 19.0-24.9 | 28.5-33.5 | | (9.9) | (21.1) | (31.0) | | | | 40 |27 | 7.0-11.5 | 15.0-23.0 | 23.0-34.5 | | (9.1) | (22.0) | (31.2) | | | | 43 |10 | 8.0-12.0 | 11.0-17.0 | 20.0-26.0 | | (10.2) | (13.5) | (23.7) | | | | 45 |16 | 10.0-12.0 | 1.0-7.0 | 12.0-17.0 | | (11.2) | (3.4) | (14.6) | | | | 46 |45 | 11.0-13.0 | | | | (11.8) | | in life, dorsum yellowish tan with gray-brown mottling; belly and ventrolateral surfaces silvery-gold or white; black stripe from tip of snout to eye; two black blotches below eye, another blotch extending from eye to base of caudal musculature; caudal musculature and fins gray-brown. in preservative, yellowish tan and silvery-gold colors lost; black reticulations present on tail. _remarks._--cope (1876:105) described _hyla elaeochroa_ from sipurio, limón province, costa rica. he based his description on a small specimen, 26.0 mm. in snout-vent length, having a dorsum uniformly colored and lacking an interorbital triangle and blotches on the thighs. cope (1887) described pigmented specimens from nicaragua as _hyla quinquevittata_, which he diagnosed as having dark brown bars on the hind limbs and five dark brown longitudinal stripes on the dorsum, the median one of which was expanded anteriorly so as to form a large triangular spot between the eyes. he thought this species was related to _hyla eximia_ baird and noted that "the hinder legs are much larger; the muzzle is more acuminate and the color bands are much wider" than in _eximia_. cope did not compare _quinquevittata_ with _elaeochroa_, which he had described ten years before. günther (1901:268), noble (1918:340), and nieden (1923:251) regarded both _elaeochroa_ and _quinquevittata_ as valid species. dunn and emlen (1932:25) regarded both as synonyms of _hyla rubra_, but they made no qualifying statements. taylor (1952:859) placed _quinquevittata_ as a synonym of _elaeochroa_ and indicated that _rubra_ was another species. taylor (1958:37) described _hyla dulcensis_ from the humid tropical forests of golfo dulce, puntarenas province, costa rica. he thought this species was "related to _h. elaeochroa_ but differs in its somewhat larger size, smaller finger and toe discs, the obsolete canthus rostralis; the loreal region concave and the choanae larger." duellman (1966a:270) compared adults, tadpoles, and mating calls of _dulcensis_ and _elaeochroa_ and concluded that a single species was involved. _hyla elaeochroa_ can be easily confused with the closely related _hyla staufferi_. although the durations of the calls are similar, the call of _elaeochroa_ has only about one third the number of pulses per second, a much lower fundamental frequency, and a lower dominant frequency than that of _staufferi_. _hyla elaeochroa_ is larger and has a less pointed snout than does _staufferi_. although the skulls of the two species are similar, that of _elaeochroa_ differs in having broad palatines and comparatively larger nasals that are truncate anteriorly. in _staufferi_ the nasal is rounded anteriorly and the palatine is absent. _distribution._--_hyla elaeochroa_ occurs on the caribbean lowlands from western panamá through costa rica to eastern nicaragua, and on the pacific lowlands of southeastern costa rica and extreme western panamá. most localities where it has been collected are below 800 meters, but the species has been found at two localities above 1000 meters (el silencio and pacuare, cartago province) on the caribbean slopes of the cordillera de talamanca, costa rica (fig. 6). _specimens examined._--nicaragua: _zelaya_: el recreo, ummz 79721 (9). costa rica: _alajuela_: laguna monte alegre, ku 64499. _cartago_: 2 km e chitaría, ku 107058; el silencio, 14.4 km ne turrialba, ku 107059-60; 4.6 km ene pacuare, ku 64451-75, 64628-37; 4 km s pavones, ku 64500; turrialba (instituto interamericano de ciéncias agrícolas), ku 30305-26, 24616-57, 30337-54, 31776-91, 31803, 31807-15, 64413-50, 68283-87 (skeletons), 68390-1 (young), 35042 (eggs), 25207-8 (skeletons), 25221 (skeleton), 41073-83 (skeletons). _guanacaste_: 2 km e tilarán, ku 86356-77, 87667-8 (young). _heredia_: puerto viejo, ku 36696, 46466, 64501-17, 68288-91, 68387, 68388-9 (young), 91803 (young), 91688-9, 104134 (tadpoles), 104135 (young), 104354-6 (skeletons); 1.5 km n puerto viejo, ku 64518-23, 68386 (tadpoles); 1 km s puerto viejo, ku 84985-6 (skeletons), 87669 (young), 87772-3 (skeletons). _limón_: bataán, ku 30327-36; la lola, ku 64478-98, 68281-2 (skeletons); los diamantes, ku 31800-02, 64476-7; peralta, ku 31816-21; puerto limón, ku 31792-99; suretka, ku 36467-79, 36697, 41084. _puntarenas_: 5 km nw buenos aires, ku 107057; 10 km e esparta, ku 87666 (tadpoles); golfito, ku 32166-8; 8 km e palmer norte ku 93939; 10.7 km se palmar sur, ku 93938 (skeleton), 93940-51, 93952 (eggs), 93953-6 (tadpoles); piedras blancas, ku 103646-59; 4.5 km w rincón de osa, ku 102208-41, 104298 (tadpoles). [illustration: fig. 6. map showing locality records for _hyla elaeochroa_ (circles) and _h. rubra_ (dots).] panamá: _bocas del toro_: almirante, ku 80079; isla bastimentos, ku 96008-11; río cricamola, 3.7 km from coast, ku 96012. _chiriquí_: río gariché, 8.3 km ese paso de canoas, ku 101571-2. _hyla staufferi_ cope _hyla staufferi_ cope, proc. acad. nat. sci. philadelphia, 17:165, october 1, 1865 [holotype.--usnm 15317, orizaba, veracruz, méxico; francis sumichrast collector]. _diagnosis._--small frogs (male to 29 mm., female to 31.6 mm.); skull longer than wide; palatine absent; large cartilaginous crista parotica present; snout flat, elongate and protruding; dark interorbital bar and dorsal stripes usually present. _description._--head flat, especially in females, longer than wide; snout long, protruding beyond mouth; loreal region concave; canthus ill-defined; length of eye greater than internarial distance or width of eyelid; length of eye less than interorbital space; tympanum distinct; interorbital spot irregular; supratympanic fold faint; arms short; fingers free of webs; discs on third and fourth fingers equal to diameter of tympanum; inner metatarsal tubercle on base of first finger distinct; first finger shorter than second; palmar tubercle distinct (fig. 1c); legs short (usually less than 50 per cent of snout-vent length); tarsal fold absent; metatarsal tubercles small, outer tubercle smaller than inner; subarticular tubercles small, simple, distinct; toes less than half webbed (fig. 1d); skin smooth above with a few small pustules on head, scapular region, flanks, and supratympanic region; arms and legs smooth; skin of belly coarsely granular; posteroventral surfaces of thighs finely granular; tongue small, rounded, longer than wide, slightly free and notched posteriorly; vocal slits small, lateral to tongue; choanae moderate in size. _variation._--the largest males of _hyla staufferi_ are from jalapa, guatemala, and from san salvador, el salvador. in these samples the average snout-vent length is 27 mm. in panamanian specimens the average snout-vent length is 23.6 mm. slight variation in the ratio of tibia length to snout-vent length exists throughout the range; more variation exists in the ratio of the diameter of the tympanum to that of the eye; the tympanum is proportionately larger in northern populations (table 7). the primary differences between panamanian and more northern populations are in size, color pattern on the dorsum and shanks, amount of webbing between the toes, and duration of notes in the mating call (table 2, pl. 4). the color in panamanian _staufferi_ is gray or gray-brown with a pair of distinct, complete, dark brown dorsolateral stripes, a pair of entire paravertebral stripes, and in some specimens a vertebral stripe. about five per cent of the individuals have interrupted stripes on the dorsum, whereas in the more northern populations complete paravertebral stripes are present in less ten per cent of the specimens; when complete stripes are present, they are irregular. the dorsal ground color in non-panamanian specimens is brown, olive-brown, or dark brown. transverse bars are present on the shanks in _hyla staufferi_ from costa rica northward to méxico, whereas in panamá all the individuals have a longitudinal stripe on the shank (table 7, pl. 2). the interorbital spot or bar is more noticeable in northern populations than in specimens from panamá. frogs from costa rica and northward have the toes about three fourths webbed, whereas in panamá the toes are about two fifths webbed. the mating calls of the northern and panamanian populations are similar, but the notes have a longer duration in the northern populations and a higher dominant frequency in panamanian populations. _hyla staufferi_ is the most variable member of the _hyla rubra_ group in central america. the panamanian populations are geographically separated from the costa rican and more northern populations by an area of tropical rainforest in the golfo dulce region in southeastern costa rica and adjacent panamá. _hyla staufferi_ does not occur on the caribbean versant of costa rica and panamá. the golfo dulce region and the caribbean versant are humid and inhabited by _hyla elaeochroa_. _hyla staufferi_ is an inhabitant of subhumid and xeric areas. on the basis of the discontinuous variation in several characters which correlate with the disjunct distribution of the two populations, two subspecies of _hyla staufferi_ are recognized. the accounts that follow apply equally to each. _cranial osteology._--the skull of _hyla staufferi_ is flat and longer than wide. the premaxillary is small and bears 9 to 13 teeth (mean for 5 specimens, 11.3). the alary process of the premaxillary is small, concave posteriorly and vertical. ventrally, the premaxillary is united to the prevomers by partially ossified cartilage. the maxillary is slender and usually bears 49 to 70 teeth (mean for 5 specimens, 60.7). the pars facialis of the maxillary is convex and less than twice the height of the pars dentalis. the nasal is large, rounded anteriorly, and pointed posteriorly in dorsal view. the nasal comprises about 40 per cent of the total length of the skull. anteromedially the two nasals converge; posteriorly they overlap the sphenethmoid. the nasals lack a concavity in the midlateral surface. dorsally, the sphenethmoid is wider than long, roughly pentagonal in shape; the frontoparietal is elongate, narrow, and smooth, with a small supraorbital process anteriorly. the frontoparietal fontanelle is narrow anteriorly and wide posteriorly. table 7.--geographic variation in size and color in males of _hyla staufferi_. (means in parentheses below observed ranges.) ================================================================= | | |complete dorsal| | |snout-vent | stripes |barred shanks locality | n |length (mm.)| (per cent) | (per cent) ---------------+-----+------------+---------------+------------- veracruz | 47 | 23.0-27.3 | 0.0 | 100 | | (25.4) | | | | | | campeche | 20 | 24.6-27.5 | 0.0 | 100 | | (25.5) | | | | | | oaxaca | 75 | 24.0-28.7 | 9.3 | 100 | | (26.4) | | | | | | chiapas | 20 | 23.2-27.8 | 10.0 | 100 | | (25.5) | | | | | | guatemala | 22 | 25.0-29.0 | 10.9 | 100 | | (26.9) | | | | | | el salvador | 21 | 24.7-28.6 | 0.0 | 100 | | (27.0) | | | | | | honduras | 34 | 20.6-27.0 | 3.3 | 100 | | (24.9) | | | | | | nicaragua | 67 | 21.5-26.8 | 3.0 | 92.7 | | (24.9) | | | | | | costa rica | 54 | 20.7-26.6 | 5.5 | 98.1 | | (24.2) | | | | | | total | 360 | 20.7-29.0 | 5.4 | 98.3 non-panamanian | | (25.9) | | | | | | panamá | 72 | 21.7-26.0 | 94.5 | 0.0 | | (23.6) | | only a narrow connection exists between the posterior, pointed arm of the squamosal and the lateral edge of the proötic. the crista parotica is visible dorsally along the lateral edge of the bony proötic. the squamosal is narrow anteriorly and posteriorly. the prevomers are short and separated anteriorly by partly ossified cartilage of the overlying solum nasi. the prevomer is joined to the premaxillary by cartilage. the posterior margin of the prevomer articulates directly with the sphenethmoid. the anterolateral and posterolateral processes of the prevomers form the incomplete bony internal margin of the choanae. each prevomer bears three to six teeth. the palatine is absent. the anterior part of the parasphenoid is narrow and ends in a point. the pterygoid is slender and weakly developed. _natural history._--throughout its range _hyla staufferi_ occurs in subhumid forests and savannas; consequently, the breeding activities are limited by the seasonal occurrence of rainfall, which accumulates in temporary ponds where this species breeds. clasping pairs and gravid females have been found mostly from june to august throughout its range. this species was observed calling at finca taboga, guanacaste province, costa rica, in mid-july. the males were calling from temporary grassy and weedy ponds in which _hyla microcephala_ also was calling, but the two species had different calling sites. _hyla staufferi_ called at stations at heights of five to 80 cm. near the edge of the pond, whereas _hyla microcephala_ called from emergent vegetation in the middle of the pond. charles w. myers informed me that at penonomé, coclé, panamá, he found _staufferi_ calling from grass in puddles where _microcephala_ was absent, and at el caño, coclé, panamá, _staufferi_ was calling from higher sites ("several inches to a few feet above water") than _microcephala_. stuart (1948:34) reported breeding individuals from la libertad, guatemala, after rainfall in late may, and schmidt and stuart (1941:239) reported _staufferi_ breeding in july in the salamá basin, alta verapaz, guatemala. stuart (1935:38) and duellman (1960:63 and 1963:226) agreed that this species breeds early in the rainy season. however, rand (1957:519) stated that in el salvador "these frogs did not begin to call until almost a month and a half after the beginning of the rains." blair (1960:133) reported that males call in june and july in chiapas, oaxaca, veracruz, and tamaulipas, méxico. the mating call of this species is a series of closely spaced notes having a fundamental frequency of about 100 cycles per second. each note has a duration of 0.13 to 0.23 second, repeated at intervals that are longer than the duration of the call. the notes are moderately low-pitched and have a dominant frequency of more than 3,000 cycles per second and about 120 pulses per second (table 2). _tadpoles._--measurements of the 33 tadpoles that are available are given in table 8. the largest tadpole examined is in stage 38 and has a total length of 29.5 mm. a typical tadpole in stage 38 of development (ku 104162, 5 km ese córdoba, veracruz, méxico) has a body length of 10 mm., tail length of 19.5 mm., and a total length of 29.5 mm. other characters are as follows: body as deep as wide, depressed anteriorly; body as long as depth of tail; interorbital space greater than distance between eye and snout but equal to internarial space; nostril equidistant between eye and tip of snout; distance between spiracle and eye less than distance between eye and snout; eyes large, situated dorsolaterally; mouth anteroventral, approximately triangular in outline; one row of papillae covering lower lip and all except median fourth of upper lip; scattered papillae at corners of mouth; tooth rows 2/3; first upper row entire, second row interrupted medially, shorter than first; lower rows shorter than upper rows; beak weak; spiracle short and nearer eyes than anus; anal opening not reaching edge of ventral fin; dorsal fin barely extending onto body; caudal musculature pointed distally. table 8.--sizes of tadpoles of _hyla s. staufferi_ in relation to developmental stages. (means in parentheses below observed ranges; measurements in mm.) ====================================================== stage | n | body length| tail length | total length --------+---+------------+-------------+------------- 25 | 3 | 6.0-7.0 | 12.0-13.0 | 18.0-20.0 | | (6.7) | (12.5) | (19.2) | | | | 26 | 2 | 7.0-7.5 | 14.0-15.0 | 21.5-22.0 | | (7.3) | (14.5) | (21.8) | | | | 27 | 9 | 7.0-8.0 | 13.0-17.0 | 21.0-25.0 | | (7.6) | (14.5) | (22.0) | | | | 32 | 1 | 8.5 | 15.5 | 24.0 | | | | 36 | 2 | 8.0-10.0 | 16.5-17.0 | 25.0-26.5 | | (9.0) | (16.8) | (25.8) | | | | 38 | 6 | 9.0-10.0 | 19.0-20.5 | 28.0-29.5 | | (9.6) | (19.5) | (29.1) | | | | 41 | 1 | 10.0 | 14.0 | 24.0 | | | | 42 | 6 | 11.0-14.0 | 10.0-13.0 | 20.0-29.0 | | (11.8) | (11.9) | (24.8) | | | | 45 | 1 | 12.5 | 0.5 | 13.0 | | | | 46 | 1 | 13.0 | - | -in life, body pale olive-tan, belly silvery white with pinkish-orange reticulations in some specimens; tail creamy white with silvery flecks and black or brown reticulations. in preservative, tan and pinkish-orange coloration lost; body transparent, reticulations on tail present. _remarks._--_hyla staufferi_ was described by cope (1865:195) on the basis of specimens from orizaba, veracruz, méxico. he described the color pattern as "color above dark olive, with a short black bar over each scapula, and one from eye to eye, with a trace along the coccyx." cope (1887:14) placed _staufferi_ as a subspecies of _hyla eximia_, but he did not justify his action. günther (1901:262) also considered _staufferi_ to be conspecific with _eximia_ without making any qualifying statement. dunn and emlen (1932:24) named _hyla culex_ from tela, honduras, on the basis of a male (mcz 16098) having a snout-vent length of 25.1 mm., and a female (usnm 20267) from patuca, honduras. they diagnosed the species as having "discs larger than tympanum ... black interorbital triangle, traces of black dorsal marking; three black bars on anterior and posterior face of thighs, two black bars on tibia, on tarsus and on forearm." the holotype now is faded but has some of the pattern described. dunn and emlen did not compare _culex_ with _staufferi_ but did compare it with _boulengeri_ and _rubra_. dunn (1933:61) named _hyla altae_ from summit, canal zone. his description was based on a male (mcz 17972) having a snout-vent length of 25.1 mm., the color pattern was described as "gray with four darker dorsal stripes ... a faint trace of mid-dorsal striping...." dunn defined the _hyla rubra_ group and recognized _boulengeri_, _altae_, _culex_, and _rubra_ as members. _hyla elaeochroa_ and _staufferi_ were omitted from his key to the group in central america. kellogg (1932:174) compared _staufferi_ with _eximia_ and concluded that the two were probably distinct species. stuart (1935:38) considered _altae_ to be a synonym of _culex_. gaige (1936:293) considered _altae_ and _culex_ to be conspecific but regarded _staufferi_ as a different species. she also suggested that _staufferi_ was not related to _eximia_ but belonged to the _rubra_ group. taylor (1952:865) and duellman (1966a:274) considered _altae_ and _culex_ to be synonyms of _staufferi_. the only other worker besides cope and günther to consider _hyla staufferi_ as a member of the _eximia_ group was blair (1960:129), who suggested the relationship on the basis of similarities in the structure of the calls of _eximia_ and _staufferi_. taylor (1938:421) and smith and taylor (1948:78) excluded _staufferi_ from the _eximia_ group on the basis of morphological characteristics. i consider _culex_ to be inseparable from _staufferi_, whereas _altae_ is recognizable as a panamanian subspecies of _staufferi_. _hyla staufferi staufferi_ cope, new combination _hyla staufferi_ cope, proc. acad. nat. sci. philadelphia, 17:195, october 1865 [holotype.--usnm 15317, orizaba, veracruz, méxico; francis sumichrast collector], brocchi, mission scientifique au mexique et dans l'amerique centrale, 1881, p. 36. boulenger, catalogue, of the bratrachia salientia s. ecaudata, p. 400, february 1, 1882. kellogg, bull. u.s. natl. mus., 160:173, march 31, 1932. smith and taylor, bull. u.s. natl. mus., 194:88, 1948. taylor, univ. kansas sci. bull., 35:862, july 1, 1952. rand, fieldiana zool. chicago nat. hist. mus., 34:518, april 18, 1957. duellman, univ. kansas publ, mus. nat. hist., 17:274, june 17, 1966. _hyla eximia staufferi_ cope, bull. u.s. natl. mus., 32:14, january 16, 1887. _hyla eximia_ (part): günther, biologia centrali-americana, reptilia and batrachia, p. 261, june 1901. nieden, das tierreich, anura i, p. 245, june 1923. _hyla culex_ dunn and emlen, proc. acad. nat. sci. philadelphia, 84:24, march 22, 1932 [holotype.--mcz 16098, tela honduras; raymond a. stadelman collector]. stuart, misc. publ., univ. michigan mus. zool., 29:38, october 1935. gaige, carnegie inst. washington publ., 457:293, 1936. _diagnosis._--small frogs (male to 29 mm., female to 31.6 mm.); dorsolateral stripes irregular; paravertebral stripes usually broken; two or three transverse bars on shanks; thighs spotted or not; arms usually barred; interorbital bar usually present; toes about three fourths webbed; color brown, tan, or olive-green. _variation._--three hundred and sixty males chosen at random from throughout the range have snout-vent lengths of 20.7 to 29 mm. (25.9 mm.). the smallest individuals are from costa rica and nicaragua (means 24.2 and 24.4 mm., respectively). the largest individuals are from guatemala and el salvador (mean of each 27.0 mm.). the ratio of the diameter of the tympanum to that of the eye is more than 60 per cent in most samples, but in those from costa rica and british honduras it is smaller. the color pattern is highly variable. some specimens are dark brown or pale brown in color. incomplete dorsal stripes are present in 94.6 per cent of the specimens, and transverse bars are present on the shanks in 98.3 per cent of the specimens. the interorbital spot varies from transverse to longitudinal in position, and an irregular white line extends from the upper jaw to the arm in some specimens (table 7). _distribution._--_hyla staufferi staufferi_ inhabits savanna and subhumid and xeric forests in the lowlands and moderate elevations from southern tamaulipas southward to nicaragua on the caribbean versant and from guerrero, méxico to northwestern costa rica on the pacific lowlands (fig. 7). duellman (1963:226) commented that a specimen from chinajá, guatemala, possibly was transported there in the cargo from toocog, because with this one exception the species is unknown in tropical rainforest in guatemala. [illustration: fig. 7. map showing locality records for _hyla staufferi staufferi_ (circles) and _h. staufferi altae_ (dots).] _specimens examined._--méxico: _campeche_: 5 km s champotón ku 71296-7; 7 km w escárcega, ku 71298-308; 13 km w, 1 km n escárcega, ku 71309-10, 75090-4. _chiapas_: 32 km s arriaga, ku 57789-92; 4 km n ixtapa, ku 5776-81; 3.6 km sw las cruces, ku 37740; 17 km s las cruces, ku 57793-4; 24 km s las cruces, ku 104160 (tadpoles); 11 km s tapachula, ku 57782-8, 60000 (young). _guerrero_: el limoncito, near la venta, ku 31392-401; mexcala, near balsa river, ku 31391; organos, s el trienta, ku 31390. _oaxaca_: 26 km n matías romero, ku 33878-82; 2.5 km s pochutla, ku 59924-7 (skeletons); 5 km s pochutla, ku 57795-801; 3.2 km e tapanatepec, ku 37877-902; 17.6 km wnw tapanatepec, ku 65033-4; temascal, usc 8243 (8); 3.2 km s tolocita, ku 39657-8; 0.5 km tuxtepec, ku 87073-81, 87610 (tadpoles); 17 km s tuxtepec, ku 65035-7; 1 km w zanatepec, ku 104161 (tadpoles). _quintana roo_: isla cozumel, 3.5 km n san miguel, ku 71710-11 (young). _san luis potosí_: valles, ku 31490. _tabasco_: teapa, ummz 118887 (3), 119203 (13); 9.6 km n teapa, ummz 119202; 24 km n teapa, ummz 119961 (5); 29 km n teapa, ummz 119960; 3.5 km s villahermosa, ummz 119201 (2); 17.6 km s villahermosa, ummz 119200 (8). _tamaulipas_: 1 km e chamal, ummz 110706; gómez farías, ummz 110701 (3); 5 km se gómez farías, ummz 110705; 8 km ne gómez farías, ummz 11282 (2), 11283 (3); kilometer 615 between río limón and llera, ummz 80455 (2); 5 km w san geraldo, ummz 110702 (4), 110703 (3); 8 km w san geraldo, near río frío, ummz 110704 (5). _veracruz_: 3 km sw boca del río, ku 10494-8; 5 km sw boca del río, ku 23701; 5 km ese córdoba, ku 104162 (tadpoles); cuautlapán, ku 57098-102, 26787; hacienda tamiahua, cabo rojo, ku 62871; 2 km ene mata oscura, ku 105627; 5 km se paso del toro, ku 40144; portrero viejo, ku 23911-2, 26786, 27413, 57094-7. guatemala: _alta verapaz_: chinajá, ku 57769; finca la cubilquitz, ummz 90871, 90872 (5), 91379 (2). _baja verapaz_: 1 km s san jerónimo, ummz 84077 (7), 84078 (14). _chiquimula_: 1.6 km se chiquimula, ummz 98114 (2); esquipulas, ummz 106784 (4), 106785 (14). _el petén_: no specific locality, usnm 25143, 24825-6; la libertad, fmnh 27096-7, ku 57770, ummz 75339 (15), 75340 (15), ummz 94341-2. _esquintla_: 20 km n san josé, amnh 74369-76. _guatamala_: 16 km ne guatamala, ku 43539. izábal: puerto barrios, tcwc 16671-73, 16646-56; 2.5 km ne río blanco, ku 57774-5. _jalapa_: jalapa, ummz 106788 (44). _jutiapa_: finca la trinidad, ummz 107730 (12), 107731 (16); jutiapa, ummz 106786 (2). _zacapa_: 14 km ene mayuelas, ku 57773; 7 km ene río hondo, ku 57771-2, 59999 (young). british honduras: belize: belize, fmnh 4406. _el cayo_: san agustín, ummz 80741 (8). _stann creek_: 10 km s stann creek on hummingbird highway, ummz 125720-1. el salvador: _cuscatlán_: 7 km wnw cojutepeque, tnhc 32004-10. _la libertad_: 16 km nw santa tecla, ku 43540-1. _la union_: 2.5 km santa rosa, tcwc 16669-70. _morazán_: dividendero, usnm 73288-92. _san salvador_: san salvador, fmnh 65101-06, ku 61932-44, 61989-92, 62152 (eggs), usnm 117588, 118391 (3), 118394; 1.6 km nw san salvador, ku 43162-3. honduras: _atlantidad_: ceiba, usnm 117592. _choluteca_: choluteca, ku 85361-6; 2 km e choluteca, ummz 118395 (7); 3.2 km ne choluteca, ku 100500-01; 6.2 km e choluteca, ku 65046-56; 10 km e choluteca, ku 65045: 5 km s choluteca, usc 2700 (4). _colón_: isla guanaja (islas de la bahía), tcwc 21551, tnhc 32011. _cortés_: agua azul, tcwc 19178-9; east side lago yojoa, ku 65038-44. _el paraiso_: valle de jamastrán, amnh 54800-04. _francisco morazán_: escuela agrícola panamericana, amnh 54963-73; 14.5 km nw comayaguela, ku 100499; el zamorano, ku 103224; 29 km n tegucigalpa, tnhc 32003, 32012. nicaragua: _chinandega_: finca san isidro, 10 km s chinandega, ku 85311-33. _managua_: 13 km e managua, ku 85339; 2 km s tipitapa, ku 85334-8. _rivas_: 9.5 km se rivas, ku 85355-6; 18 km se rivas, ku 85354; 7.7 km ne san juan del sur, ku 85346-53; 16.5 km ne san juan del sur, ku 85340-5; 5 km se san pablo, ku 43151-61. _zelaya_: isla grande del maiz, ku 85357-60. costa rica: _alajuela_: _los chiles_, usc 7215 (2), 7217. _guanacaste_: 4 km w bagaces, usc 7019 (5); finca taboga, ku 102265-5; 12 km s la cruz, usc 8091; las cañas, ku 41113 (skeleton); 27 km n las cañas, usc 8171 (5); guardia, río tempisque, usc 8214; 10 km n guardia, ku 102266-7; 1.6 km n guayabo de bagaces, usc 7023 (3); liberia, ku 36510-22; 4 km w liberia, ku 36449-64, usc 102 (10), 103 (9), 104 (7), 105; 6 km n liberia. usc 8096; 8 km nnw liberia, ku 65032; 14.5 km n liberia, usc 8079, 8138 (2); 14.5 km s liberia, usc 8238 (5); 6 km n nicoya, usc 8229 (11); 4 km s nicoya, usc 8230, 8231; peñas blancas, ku 102263; 8.6 km ese playa del coco, usc 8137 (14); 21 km e playa del coco, usc 8138 (2); santa cruz, usc 8232 (2); 3 km e santa rosa, tcwc 16663-68; tenorio, ku 32159; tilarán, ku 36509. _puntarenas_: 10 km wnw esparta, ku 65022-9, 68614 (skeleton); 4.5 km wnw esparta, ku 65030; 12 km wnw esparta, ku 65031; 6 km e esparta, ku 86477; hotel maribella, ku 32157-8; 3 km w puntarenas, tcwc 16657-62. _hyla staufferi altae_ dunn, new combination _hyla altae_ dunn, occas. papers boston soc. nat. hist., 8:61, june 7, 1933 [holotype.--mcz 17972, summit, canal zone, panamá; emmett r. dunn collector]. _hyla culex_: stuart, misc. publ. univ. michigan mus. zool., 29:38, october 1, 1935. gaige, carnegie inst. washington publ., 457:293, 1936. _hyla staufferi_: taylor, univ. kansas sci. bull., 35:862, july 1, 1952. duellman, univ. kansas publ., mus. nat. hist., 17:274, june 17, 1966. _diagnosis._--small frogs (male to 26 mm., female to 27 mm.); dorsolateral and paravertebral stripes complete; longitudinal dark gray stripe on shank; thighs unmarked; interorbital bar usually absent; toes about three fifths webbed; gray to brownish gray above. _variation._--_hyla staufferi altae_ is less variable in size, proportions, and color pattern than is _h. s. staufferi_. the size varies from 21.7 to 26 mm. (23.6) in 72 males. the ratio of tibia to snout-vent length is 0.42 to 0.50 (0.45), slightly less than in the northern subspecies. in color pattern 94.5 per cent of the individuals have complete dorsal stripes, and all have a longitudinal stripe on the shank (table 7). _distribution._--this subspecies is restricted to subhumid forests and savannas on the pacific lowlands of panamá. _hyla s. altae_ is presently known to occur from chepo in east-central panamá through the azuero peninsula to concepción, chiriquí, in western panamá (fig. 7). _specimens examined._--panamá: _canal zone_: no specific locality, tnhc 24406; 2.8 km sw fort kobbe, ku 101679. _chiriquí_: 14.4 km e concepción, amnh 69799-801; 6.6 km n david, tnhc 32013-4; 2 km s david, amnh 68802. _coclé_: 1 km ne el caño, ku 101662-75; el valle de antón, amnh 59601-5, ku 77333-47; 7 km ssw penonomé, ku 101654-61. _los santos_: tonosí, ku 101246 (tadpoles), 101697-701. _panamá_: 2 km wsw chepo, ku 101680-8; 6 km wsw chepo, ku 77324-27; el cangrejo (panamá), ku 101676-8; nueva gorgona, amnh 69991, 69798; 1.5 km w pacora, ku 77328-32; 2 km n tocumen, ku 101689-95; 8 km ne tocumen, ku 101696. evolutionary history my assumptions regarding the evolutionary history of the _hyla rubra_ group in central america were derived partly from interpretations of the evolutionary history of other animal groups (simpson, 1943, 1965; dunn, 1931b; stuart, 1950; duellman, 1958, 1960, 1963, 1965; and duellman and trueb, 1966). the origin and early evolution of the group probably occurred prior to the mid-pliocene in the lowlands of south america, because the greatest diversity of the group is in brazil. differentiation into two or more subgroups took place in south america prior to the late pliocene. at the end of the pliocene, shortly after the closure of the colombian portal, many south american animals migrated into central america (simpson, 1943, maldonado-koerdell, 1964, and savage, 1966). it is likely that the _hyla rubra_ group entered central america at that time; apparently two stocks (_rubra-elaeochroa-staufferi_ stock and _boulengeri-foliamorta_ stock) migrated into central america. _hyla elaeochroa_ is closely related to _rubra_ and probably differentiated from _rubra_ through spatial isolation. thus, we have _elaeochroa_ in central america and _rubra_ in south america; most likely only in relatively recent times has _rubra_ migrated into eastern panamá from northern south america. the differentiation and dispersal of _elaeochroa_ and _staufferi_ took place in central america after the pliocene. probably the events of the pleistocene resulted in the isolation of populations. one of these (_hyla staufferi_ stock) was restricted in the subhumid pacific lowlands, whereas the _hyla elaeochroa_ stock occupied the tropical wet forests of the caribbean lowlands. _hyla elaeochroa_ apparently more closely resembled the parental stock by being restricted to the tropical rain forests, whereas _staufferi_ adapted to subhumid environments and thereby was able to disperse throughout most of the subhumid regions of central america. after geographical separation took place the initial genetic divergence between the two populations was maintained by means of ecological and ethological isolating mechanisms. under these circumstances it can be supposed that the different ecological preferences of _elaeochroa_ and _staufferi_ depend on the climatic changes that took place during the pleistocene. on this basis it may be proposed that when the original prototype broke up into the two incipient species, the _staufferi_ stock became physiologically and behaviorally adapted to subhumid conditions and dispersed into dry areas of the lowlands of middle america. the tropical evergreen forests on the caribbean side of lower central america and the uplift of the talamanca range in the pliocene were barriers to the dispersal of _staufferi_. consequently, this frog dispersed along the pacific lowlands. at the present time _staufferi_ occupies the length of the pacific lowlands in central america, except in the rainforest of the golfo duce region, which apparently is a relict stand and now separates the ranges of two subspecies of _hyla staufferi_. this species crossed the central nicaraguan lowlands and reached the caribbean lowlands of nicaragua and nuclear central america. the species migrated through the subhumid corridor in northern honduras and eastern guatemala (comayagua valley in honduras and the motagua valley of guatemala) to the isthmus of tehuantepec. duellman (1960) hypothesized "that during the times of glacial advances (pleistocene) the lowlands of the isthmus probably were more extensive and had more semiarid tropical environments than at the present" and that when semiarid environments were continuous from the pacific slope across the isthmus to the gulf lowlands _staufferi_ and other amphibians migrated northward to southeastern tamaulipas, méxico. _hyla elaeochroa_ dispersed along caribbean lowland routes. this species not only occurs in the wet forests of the golfo dulce region but also in guanacaste. it is possible that _elaeochroa_ entered guanacaste and moved to the golfo dulce region when the intervening area was less xeric than now (duellman, 1966b). _hyla elaeochroa_ extended its range to eastern nicaragua, but even though northeastern nicaragua has over 2,000 mm. of precipitation annually (vivo escoto, 1964), this species has not spread into honduras and guatemala. _hyla boulengeri_ is widespread in amazonian and northern south america, whereas _foliamorta_ occurs only in eastern panamá and in north-central colombia. the ancestral _boulengeri-foliamorta_ stock probably invaded central america in the late pliocene and dispersed through humid forested environments to nicaragua. apparently a peripheral population established itself in the dry pacific lowlands of panamá. this population differentiated from _boulengeri_ of the humid caribbean lowlands and evolved into _foliamorta_, which subsequently expanded its range into colombia. literature cited blair, w. f. 1960. mating call as evidence of relations in the _hyla eximia_ group. southwestern nat., 5(3):129-135. november 1. boulenger, g. a. 1882. catalogue of the batrachia salientia s. ecaudata, 2nd. ed., pp. 1-503, pls. 1-30. february. brocchi, p. 1881-1883. etude des batraciens de l'amerique centrale. mission scientifique au mexique et dans l'amerique centrale, liv. 1, pp. 1-122, pls. 1-21. cochran, d. m. 1955. frogs of southeastern brazil. bull. u.s. natl. mus., 206:1-423. cope, e. d. 1865. third contribution to the herpetology of tropical america. proc. acad. nat. sci. philadelphia, 17:195. 1876. on the batrachia and reptilia of costa rica. jour. acad. nat. sci. philadelphia, new series, 8:93-154, pls. 23-28. 1887. thirteenth contribution to the herpetology of tropical america. proc. amer. philos. society, 23:273. april. daudin, f. m. 1802. histoire naturelle des rainettes, des grenouilles et des crapauds. paris, pp. 1-71, pls. 1-33. 1803. histoire naturelle générale et particulière des reptiles. paris, 8:1-439. duellman, w. e. 1956. the frogs of the hylid genus _phrynohyas fitzinger_, 1843. misc. publ. mus. zool., univ. mich., 96:1-47, pls. 1-6. february 21. 1958. a monographic study of the colubrid snakes genus _leptodeira_. bull. amer. mus. nat. hist., 114:1-152, pls. 1-31. february 24. 1960. a distributional study of the amphibians of the isthmus of tehuantepec, méxico. univ. kansas publ., mus. nat. hist, 13:19-72, pls. 1-8. august 16. 1963. amphibians and reptiles of the rainforests of southern el petén, guatemala. univ. kansas publ., mus. nat. hist., 15:205-249, pls. 7-10. october 4. 1965. a biogeographic account of the herpetofauna of michoacán, méxico. univ. kansas publ., mus. nat. hist., 15:627-709, pls. 29-36. december 30. 1966a. taxonomic notes on some mexican and central american hylid frogs. univ. kansas publ., mus. nat. hist., 17:263-279. june 17. 1966b. the central american herpetofauna: an ecological perspective. copeia, 4:700-719. december 23. 1967. social organization in the mating calls of some neotropical anurans. amer. midl. nat., 77(1):156-163. january. duellman, w. e., and l. trueb 1966. neotropical hylid frogs, genus smilisca. univ. kansas publ., mus. nat. hist., 17:281-375, pls. 1-12. july 14. dunn, e. r. 1931a. the amphibians of barro colorado island. occas. papers boston soc. nat. hist., 5:403-421. october 10. 1931b. the herpetological fauna of the americas. copeia, 3:106-119. october 30. 1933. a new hyla from panamá canal zone. occas. papers boston soc. nat. hist., 8:61-64. june 7. dunn, e. r., and j. emlen 1932. reptiles and amphibians from honduras. proc. acad. nat. sci. philadelphia, 84:24-25. march 22. fouquette, m. j. 1958. a new tree frog, genus _hyla_, from the canal zone. herpetologica, 14:125-128. april 25. gaige, h. t. 1936. some reptiles and amphibians from yucatan and campeche, méxico. carnegie inst. washington publ., 457:289-304. gosner, k. l. 1960. a simplified table for staging anuran embryos and larvae with notes on identification. herpetologica, 16:183-190. june 17. günther, a. c. l. 1859. catalogue of the batrachia salientia in the british museum. pp. 1-160. 1885-1902. biologia centrali americana. reptilia and batrachia. london, xx + 326 pp., pls. 1-76. kellogg, r. 1932. mexican tailless amphibians in the united states national museum, bull. u.s. natl. mus., 160:1-224. laurenti, j. n. 1768. specimen medicum exhibens synopsin reptilium emendatum cum experimentis circa venema et antidota reptilium austriacorum. vienna, pp. 1-216. linnaeus, c. 1758. systema naturae. holmiae, ed. 10 reformata, i:1-532. maldonado-koerdell, m. 1964. geohistory and paleography of middle america. in r. wauchope and r. c. west (eds.). handbook of middle american indians, vol. i, univ. texas press, austin, 570 pp. nieden, f. 1923. amphibia:anura i. das tierreich. berlin, 584 pp. noble, g. k. 1918. the amphibians collected by the american museum expedition to nicaragua in 1916. bull. amer. mus. nat. hist., 38:311-347. rand, a. s. 1957. notes on amphibians and reptiles from el salvador. fieldiana, zool., chicago nat. hist. mus., 43:505-534. april 18. rivero, j. a. 1961. salientia of venezuela. bull. mus. comp. zool., 126:1-207. november. savage, j. m. 1966. the origins and history of the central american herpetofauna. copeia, 4:719-756. december 23. schmidt, k. p., and l. c. stuart 1941. the herpetological fauna of the salamá basin, baja verapaz, guatemala. field mus. nat. hist., zool. ser., 24:233-247. seba, a. 1734. locupletissimi rerum naturalium thesauri accurata descriptio, et iconibus artificissimis expressio, per universam physis historiam. amsterdam, i xxxiv + 178 pp., pls. 1-111. simpson, g. g. 1943. turtles and the origin of the fauna of latin america. amer. jour. sci., 24:413-429. 1965. the geography of evolution. chilton books publishers. philadelphia, pp. 1-349. july. smith, h. m. and e. h. taylor 1948. an annotated checklist and key to the amphibians of méxico. bull. u.s. natl. mus., 194:1-118. starrett, p. 1960. description of tadpoles of middle american frogs. misc. publ. mus. zool., univ. michigan, 110:1-37, pl. 1. stuart, l. c. 1935. a contribution to a knowledge of the herpetology of a portion of the savanna region of central petén, guatemala. misc. publ. mus. zool., univ. michigan, 29:1-56, pls. 1-4. october 1. 1948. the amphibians and reptiles of alta verapaz, guatemala. misc. publ. mus. zool., univ. michigan, 69:1-109. june 12. 1950. a geographic study of the herpetofauna of alta verapaz, guatemala. contr. lab. vert. biol., univ. michigan, 45:1-77, pls. 1-9. may. taylor, e. h. 1938. frogs of the _hyla eximia_ group, with description of two new species. univ. kansas sci. bull., 25:421-445. june 1. 1952. the frogs and toads of costa rica. univ. kansas sci. bull., 35:577-942. july 1. 1958. additions to the known herpetological fauna of costa rica, with comments on other species. no. iii. univ. kansas sci. bull., 34:3-40. november 18. vivo escoto, j. a. 1964. weather and climate of méxico and central america. _in_ r. wauchope and r. c. west (eds.), handbook of middle american indians. vol. 1, univ. texas press, austin, 570 pp. _transmitted february 7, 1969._ university of kansas publications museum of natural history volume 17, no. 7, pp. 281-375, pls. 1-12, 17 figs. july 14, 1966 neotropical hylid frogs, genus smilisca by william e. duellman and linda trueb university of kansas lawrence 1966 university of kansas publications, museum of natural history editors: e. raymond hall, chairman, henry s. fitch, frank b. cross volume 17, no. 7, pp. 281-375, pls. 1-12, 17 figs. published july 14, 1966 university of kansas lawrence, kansas printed by robert r. (bob) sanders, state printer topeka, kansas 1966 [illustration: look for the union label.] 31-3430 neotropical hylid frogs, genus smilisca by william e. duellman and linda trueb contents page introduction 285 acknowledgments 286 materials and methods 287 genus smilisca cope, 1865 287 key to adults 288 key to tadpoles 289 accounts of species 289 _smilisca baudini_ (duméril and bibron) 289 _smilisca cyanosticta_ (smith) 303 _smilisca phaeota_ (cope) 308 _smilisca puma_ (cope) 314 _smilisca sila_ new species 318 _smilisca sordida_ (peters) 323 analysis of morphological characters 330 osteology 330 _descriptive osteology of smilisca baudini_ 331 _developmental cranial osteology of smilisca baudini_ 333 _comparative osteology_ 336 musculature 341 skin 342 _structure_ 342 _comparative biochemistry of proteins_ 343 external morphological characters 343 _size and proportions_ 343 _shape of snout_ 344 _hands and feet_ 344 _ontogenetic changes_ 344 coloration 344 _metachrosis_ 345 chromosomes 345 natural history 345 breeding 345 _time of breeding_ 345 _breeding sites_ 346 _breeding behavior_ 346 _breeding call_ 351 eggs 356 tadpoles 357 _general structure_ 357 _comparison of species_ 357 _growth and development_ 361 _behavior_ 365 phylogenetic relationships 366 interspecific relationships 366 evolutionary history 369 summary and conclusions 371 literature cited 372 introduction the family hylidae, as currently recognized, is composed of about 34 genera and more than 400 species. most genera (30) and about 350 species live in the american tropics. _hyla_ and 10 other genera inhabit central america; four of those 10 genera (_gastrotheca_, _hemiphractus_, _phrynohyas_, and _phyllomedusa_) are widely distributed in south america. the other six genera are either restricted to central america or have their greatest differentiation there. _plectrohyla_ and _ptychohyla_ inhabit streams in the highlands of southern mexico and northern central america; _diaglena_ and _triprion_ are casque-headed inhabitants of arid regions in méxico and northern central america. _anotheca_ is a tree-hole breeder in cloud forests in middle america. the genus _smilisca_ is the most widespread geographically and diverse ecologically of the central american genera. the definition of genera in the family hylidae is difficult owing to the vast array of species, most of which are poorly known as regards their osteology, colors in life, and modes of life history. the genera _diaglena_, _triprion_, _tetraprion_, _osteocephalus_, _trachycephalus_, _aparasphenodon_, _corythomantis_, _hemiphractus_, _pternohyla_, and _anotheca_ have been recognized as distinct from one another and from the genus _hyla_ on the basis of various modifications of dermal bones of the cranium. _phyllomedusa_ is recognized on the basis of a vertical pupil and opposable thumb; _plectrohyla_ is characterized by the presence of a bony prepollex and the absence of a quadratojugal. _gastrotheca_ is distinguished from other hylids by the presence of a pouch in the back of females. a pair of lateral vocal sacs behind the angles of the jaws and the well-developed dermal glands were used by duellman (1956) to distinguish _phrynohyas_ from _hyla_. he (1963a) cited the ventrolateral glands in breeding males as diagnostic of _ptychohyla_. some species groups within the vaguely defined genus _hyla_ have equally distinctive characters. the _hyla septentrionalis_ group is characterized by a casque-head, not much different from that in the genus _osteocephalus_ (trueb, ms). males in the _hyla maxima_ group have a protruding bony prepollex like that characteristically found in _plectrohyla_. ontogenetic development, osteology, breeding call, behavior, and ecology are important in the recognition of species. by utilizing the combination of many morphological and biological factors, the genus _smilisca_ can be defined reasonably well as a natural, phyletic assemblage of species. because the wealth of data pertaining to the morphology and biology of _smilisca_ is lacking for most other tree frogs in middle america it is not possible at present to compare _smilisca_ with related groups in more than a general way. _smilisca_ is an excellent example of an autochthonous middle american genus. as defined by stuart (1950) the autochthonous middle american fauna originated from "hanging relicts" left in central america by the ancestral fauna that moved into south america and differentiated there at a time when south america was isolated from north and middle america. the genus _smilisca_, as we define it, consists of six species, all of which occur in central america. one species ranges northward to southern texas, and one extends southward on the pacific lowlands of south america to ecuador. we consider the genus _smilisca_ to be composed of rather generalized hylids. consequently, an understanding of the systematics and zoogeography of the genus can be expected to be of aid in studying more specialized members of the family. acknowledgments examination of many of the specimens used in our study was possible only because of the cooperation of the curators of many systematic collections. for lending specimens or providing working space in their respective institutions we are grateful to doris m. cochran, alice g. c. grandison, jean guibe, robert f. inger, günther peters, gerald raun, william j. riemer, jay m. savage, hobart m. smith, wilmer w. tanner, charles f. walker, ernest e. williams, and richard g. zweifel. we are indebted to charles j. cole and charles w. myers for able assistance in the field. the cooperation of martin h. moynihan at barro colorado island, charles m. keenan of corozal, canal zone, and robert hunter of san josé, costa rica, is gratefully acknowledged. jay m. savage turned over to us many costa rican specimens and aided greatly in our work in costa rica. james a. peters helped us locate sites of collections in ecuador and coleman j. goin provided a list of localities for the genus in colombia. we especially thank charles j. cole for contributing the information on the chromosomes, and robert r. patterson for preparing osteological specimens. we thank m. j. fouquette, jr., who read the section on breeding calls and offered constructive criticism. permits for collecting were generously provided by ing. rodolfo hernandez corzo in méxico, sr. jorge a. ibarra in guatemala, and ing. milton lopez in costa rica. this report was made possible by support from the national science foundation (grants g-9827 and gb-1441) and the cooperation of the museum of natural history at the university of kansas. some of the field studies were carried out in panamá under the auspices of a grant from the national institutes of health (nih gm-12020) in cooperation with the gorgas memorial laboratory in panamá. materials and methods in our study we examined 4151 preserved frogs, 93 skeletal preparations, 88 lots of tadpoles and young, and six lots of eggs. we have collected specimens in the field of all of the species. observations on behavior and life history were begun by the senior author in méxico in 1956 and completed by us in central america in 1964 and 1965. osteological data were obtained from dried skeletons and cleaned and stained specimens of all species, plus serial sections of the skull of _smilisca baudini_. developmental stages to which tadpoles are assigned are in accordance with the table of development published by gosner (1960). breeding calls were recorded in the field on tape using magnemite and uher portable tape recorders. audiospectrographs were made by means of a vibralyzer (kay electric company). external morphological features were measured in the manner described by duellman (1956). in the accounts of the species we have attempted to give a complete synonymy. at the end of each species account the localities from which specimens were examined are listed alphabetically within each state, province, or department, which in turn are listed alphabetically within each country. the countries are arranged from north to south. abbreviations for museum specimens are listed below: amnh--american museum of natural history bmnh--british museum (natural history) byu--brigham young university cnhm--chicago natural history museum ku--university of kansas museum of natural history mcz--museum of comparative zoology mnhn--museu national d'histoire naturelle, paris uf--university of florida collections uimnh--university of illinois museum of natural history ummz--university of michigan museum of zoology usc--university of southern california usnm--united states national museum tnhc--texas natural history collection, university of texas zmb--zoologisches museum berlin =genus smilisca= cope, 1865 _smilisca_ cope, proc. acad. nat. sci. philadelphia, 17:194, oct., 1865 [type species _smilisca daulinia_ cope, 1865 = _hyla baudini_ duméril and bibron, 1841]. smith and taylor, bull. u. s. natl. mus., 194:75, june 17, 1948. starrett, copeia, 4:300, december 30, 1960. goin, ann. carnegie museum, 36:15, july 14, 1961. _definition._--medium to large tree frogs having: (1) broad, well ossified skull (consisting of a minimum amount of cartilage and/or secondarily ossified cartilage), (2) no dermal co-ossification, (3) quadratojugal and internasal septum present, (4) large ethmoid, (5) _m. depressor mandibulae_ consisting of two parts, one arising from dorsal fascia and other from posterior arm of squamosal, (6) divided _m. adductor mandibulae_, (7) paired subgular vocal sacs in males, (8) no dermal appendages, (9) pupil horizontally elliptical (10) small amounts of amines and other active substances in skin, (11) chromosome number of n = 12 and 2n = 24, (12) breeding call consisting of poorly modulated, explosive notes, and (13) 2/3 tooth-rows in tadpoles. _composition of genus._--as defined here the genus _smilisca_ contains six recognizable species. an alphabetical list of the specific and subspecific names that we consider to be applicable to species of _smilisca_ recognized herein is given below. names proposed valid names _hyla baudini_ duméril and bibron, 1841 = _s. baudini_ _hyla baudini dolomedes_ barbour, 1923 = _s. phaeota_ _hyla beltrani_ taylor, 1942 = _s. baudini_ _hyla gabbi_ cope, 1876 = _s. sordida_ _hyla labialis_ peters, 1863 = _s. phaeota_ _hyla manisorum_ taylor, 1954 = _s. baudini_ _hyla muricolor_ cope, 1862 = _s. baudini_ _hyla nigripes_ cope, 1876 = _s. sordida_ _hyla pansosana_ brocchi, 1877 = _s. baudini_ _hyla phaeota_ cope, 1862 = _s. phaeota_ _hyla phaeota cyanosticta_ smith, 1953 = _s. cyanosticta_ _hyla puma_ cope, 1885 = _s. puma_ _hyla salvini_ boulenger, 1882 = _s. sordida_ _hyla sordida_ peters, 1863 = _s. sordida_ _hyla vanvlietii_ baird, 1854 = _s. baudini_ _hyla vociferans_ baird, 1859 = _s. baudini_ _hyla wellmanorum_ taylor, 1952 = _s. puma_ _distribution of genus._--most of lowlands of méxico and central america, in some places to elevations of nearly 2000 meters, southward from southern sonora and río grande embayment of texas, including such continental islands as isla cozumel, méxico, and isla popa and isla cebaco, panamá, to northern south america, where known from caribbean coastal regions and valleys of río cauca and río magdalena in colombia, and pacific slopes of colombia and northern ecuador. key to adults 1. larger frogs ([m] to 76 mm., [f] to 90 mm.) having broad flat heads and a dark brown or black postorbital mark encompassing tympanum 2 smaller frogs ([m] to 45 mm., [f] to 64 mm.) having narrower heads and lacking a dark brown or black postorbital mark encompassing tympanum 4 2. lips barred; flanks cream-colored with bold brown or black mottling in groin; posterior surfaces of thighs brown with cream-colored flecks _s. baudini_, p. 289 lips not barred; narrow white labial stripe present; flanks not cream-colored with bold brown or black mottling in groin; posterior surfaces of thighs variable 3 3. flanks and anterior and posterior surfaces of thighs dark brown with large pale blue spots on flanks and small blue spots on thighs _s. cyanosticta_, p. 303 flanks cream-colored with fine black venation; posterior surfaces of thighs pale brown with or without darker flecks or small cream-colored spots _s. phaeota_, p. 308 4. fingers having only vestige of web; diameter of tympanum two-thirds that of eye; dorsum pale yellowish tan with pair of broad dark brown stripes _s. puma_, p. 314 fingers about one-half webbed; diameter of tympanum about one-half that of eye; dorsum variously marked with spots or blotches 5 5. snout short, truncate; vocal sacs in breeding males dark gray or brown; blue spots on flanks and posterior surfaces of thighs _s. sila_, p. 318 snout long, sloping, rounded; vocal sacs in breeding males white; cream-colored or pale blue flecks on flanks and posterior surfaces of thighs _s. sordida_, p. 323 key to tadpoles 1. pond tadpoles; tail about half again as long as body; mouth anteroventral 2 stream tadpoles; tail about twice as long as body; mouth ventral 5 2. labial papillae in two rows 3 labial papillae in one row 4 3. first upper tooth row strongly arched medially; third lower tooth row much shorter than other rows; dorsal fin deepest at about two-thirds length of tail; tail cream-colored with dense gray reticulations _s. puma_, p. 314 first upper tooth row not arched medially; third lower tooth row nearly as long as others; dorsal fin deepest at about one-third length of tail; tail tan with brown flecks and blotches _s. baudini_, p. 289 4. dorsal fin extending onto body _s. phaeota_, p. 308 dorsal fin not extending onto body _s. cyanosticta_, p. 303 5. mouth completely bordered by two rows of papillae; inner margin of upper beak not forming continuous arch with lateral processes; red or reddish brown markings on tail _s. sordida_, p. 323 median part of upper lip bare; rest of mouth bordered by one row of papillae; inner margin of upper beak forming continuous arch with lateral processes; dark brown markings on tail _s. sila_, p. 318 accounts of species =smilisca baudini= (duméril and bibron) _hyla baudini_ duméril and bibron, erpétologie général, 8:564, 1841 [holotype.--mnhn 4798 from "mexico;" baudin collector]. günther, catalogue batrachia salientia in british museum, p. 105, 1858. brocchi, mission scientifique au mexique ..., pt. 3, sec. 2, études sur les batrachiens, p. 29, 1881. boulenger, catalogue batrachia salientia in british museum, p. 371, feb. 1, 1882. werner, abhand. zool.-bot. gesell. wien., 46:8, sept. 30, 1896. günther, biologia centrali-americana: reptilia and batrachia, p. 270, sept. 1901. werner, abhand. konigl. akad. wiss. munchen, 22:351, 1903. cole and barbour, bull. mus. comp. zool., 50(5):154, nov. 1906. gadow, through southern méxico, p. 76, 1908. ruthven, zool. jahr. 32(4):310, 1912. decker, zoologica, 2:12, oct., 1915. stejneger and barbour, a checklist of north american amphibians and reptiles, p. 32, 1917. noble, bull. amer. mus. nat. hist., 38(10):341, june 20, 1918. nieden, das tierreich, amphibia, anura i, p. 243, june, 1923. gadow, jorullo, p. 54, 1930. dunn and emlen, proc. acad. nat. sci. philadelphia, 84:24, march 22, 1932. kellogg, bull. u. s. natl. mus., 160:160, march 31, 1932. martin, aquarien berlin, p. 92, 1933. stuart, occas. papers mus. zool., univ. michigan, 292:7, june 29, 1934; misc. publ. mus. zool. univ. michigan, 29:38, oct. 1, 1935. gaige, carnegie inst. washington, 457:293, feb. 5, 1936. gaige, hartweg, and stuart, occas. papers mus. zool. univ. michigan, 360:5, nov. 20, 1937. smith, occas. papers mus. zool. univ. michigan, 388:2, 12, oct. 31, 1938; ann. carnegie mus., 27:312, march 14, 1939. taylor, copeia, 2:98, july 12, 1939. hartweg and oliver, misc. publ. mus. zool. univ. michigan, 47:12, july 13, 1940. schmidt and stuart, zool. ser. field mus. nat. hist., 24(21):238, august 30, 1941. schmidt, zool. ser. field mus. nat. hist., 22(8):486, dec. 30, 1941. wright and wright, handbook of frogs and toads, ed. 2, p. 134, 1942. stuart, occas. papers mus. zool. univ. michigan, 471:15, may 17, 1943. bogert and oliver, bull. amer. mus. nat. hist., 83(6):343, march 30, 1945. taylor and smith, proc. u. s. natl. mus., 95(3185): 590, june 30, 1945. smith, ward's nat. sci. bull., 1, p. 3, sept., 1945. schmidt and shannon, fieldiana, zool. chicago nat. hist. mus., 31(9):67, feb. 20, 1947. stuart, misc. publ. mus. zool. univ. michigan, 69:26, june 12, 1948. wright and wright, handbook of frogs and toads, ed. 3, p. 298, 1949. stuart, contr. lab. vert. biol. univ. michigan, 45:22, may, 1950. mertens, senckenbergiana, 33:170, june 15, 1952; abhand. senckenb. naturf. gesell., 487:28, dec. 1, 1952. schmidt, a checklist of north american amphibians and reptiles, ed. 6, p. 69, 1953. stuart contr. lab. vert. biol. univ. michigan, 68:46, nov. 1954. zweifel and norris, amer. midl. nat., 54(1):232, july 1955. martin, amer. nat., 89:356, dec. 1955. duellman, copeia, 1:49, feb. 21, 1958. goin, herpetologica, 14:119, july 23, 1958. turner, herpetologica, 14:192, dec. 1, 1958. conant, a field guide to reptiles and amphibians, p. 284, 1958. duellman, univ. kansas publ., mus. nat. hist., 13(2):59, aug. 16, 1960; univ. kansas publ., mus. nat. hist., 15(1): 46, dec. 20, 1961. porter, herpetologica, 18:165, oct. 17, 1962. _hyla vanvlietii_ baird, proc. acad. nat. sci. philadelphia, 7:61, april 27, 1854 [holotype.--usnm 3256 from brownsville, cameron county, texas; s. van vliet collector]. baird, united states and mexican boundary survey, 2:29, 1859. smith and taylor, univ. kansas sci. bull., 33:361, march 20, 1950. cochran, bull. u. s. natl. mus., 220:60, 1961. _hyla vociferans_ baird, united states and mexican boundary survey, 2:35 1859 [_nomen nudum_]. diáz de león, indice de los batracios que se encuentran en la república mexicana, p. 20, june 1904. _hyla muricolor_ cope, proc. acad. nat. sci. philadelphia, 14(9):359, 1862 [holotype.--usnm 25097 from mirador, veracruz, méxico; charles sartorius collector]. smith and taylor, univ. kansas sci. bull., 33:349, march 20, 1950. cochran, bull. u. s. natl. mus., 220:56, 1961. _smilisca daulinia_ cope, proc. acad. nat. sci. philadelphia, 17:194, oct. 1865 [holotype.--"skeleton in private anatomical museum of hyrtl, professor of anatomy in the university of vienna"]. smith and taylor, univ. kansas sci. bull., 33:347, march 20, 1950. _smilisca daudinii_ [lapsus for _baudini_], cope, proc. acad. nat. sci. philadelphia, 23, pt. 2:205, 1871. _smilisca baudini_, cope, bull. u. s. nat. mus., 1:31, 1875; jour. acad. nat. sci. philadelphia, 8, pt. 2:107, 1876; proc. amer. philos. soc., 18:267, august 11, 1879. yarrow, bull. u. s. nat. mus., 24:176, july 1, 1882. cope, bull. u. s. nat. mus., 32:13, 1887; bull. u. s. nat. mus., 34:379, april 9, 1889. dickerson, the frog book, p. 151, july, 1906. smith and taylor, univ. kansas sci. bull., 33:442, march 20, 1950; taylor, u. kan. sc. bull., 34:802, feb. 15, 1952; univ. kansas sci. bull., 35:794, july 1, 1952. brattstrom, herpetologica, 8(3):59, nov. 1, 1952. taylor, u. kan. sci. bull., 35:1592, sept. 10, 1953. peters, occas. papers mus. zool. univ. michigan, 554:7, june 23, 1954. duellman, occas. papers mus. zool. univ. michigan, 560:8, oct. 22, 1954. chrapliwy and fugler, herpetologica, 11:122, july 15, 1955. smith and van gelder, herpetologica, 11:145, july 15, 1955. lewis and johnson, herpetologica, 11:178, nov. 30, 1955. martin, misc. publ. mus. zool. univ. michigan, 101:53, april 15, 1958. stuart, contr. lab. vert. biol. univ. michigan, 75:17, june, 1958. minton and smith, herpetologica, 17:74, july 11, 1961. nelson and hoyt, herpetologica, 17:216, oct. 9, 1961. holman, copeia, 2:256, july 20, 1962. stuart, misc. publ. mus. zool. univ. michigan, 122:41, april 2, 1963. maslin, herpetologica, 19:124, july 3, 1963. holman and birkenholz, herpetologica, 19:144, july 3, 1963. duellman, univ. kansas publ. mus. nat. hist., 15(5):228, oct. 4, 1963. zweifel, copeia, 1:206, march 26, 1964. duellman and klaas, copeia, 2:313, june 30, 1964. davis and dixon, herpetologica, 20:225, january 25, 1965. neill, bull. florida state mus., 9:89, april 9, 1965. _hyla pansosana_ brocchi, bull. soc. philom., ser. 7, 1:125, 1877 [holotype.--mnhn 6313 from panzós, alta verapaz, guatemala; m. bocourt collector]; mission scientifique au mexique ..., pt. 3, sec. 2, études sur les batrachiens, p. 34, 1881. _hyla baudini baudini_, stejneger and barbour, a checklist of north american amphibians and reptiles, ed. 3, p. 34, 1933. wright and wright, handbook of frogs and toads, p. 110, 1933. stejneger and barbour, a checklist of north american amphibians and reptiles, ed. 4, p. 39, 1939; a checklist of north american amphibians and reptiles, ed. 5, p. 49, 1943. smith and laufe, trans. kansas acad. sci., 48(3):328, dec. 19, 1945. peters, nat. hist. misc., 143:7, march 28, 1955. _hyla beltrani_ taylor, univ. kansas sci. bull. 28(14):306, nov. 15, 1942 [holotype.--uimnh 25046 (formerly eht-hms 29563) from tapachula, chiapas, méxico; a. magaña collector]. smith and taylor, bull. u. s. natl. mus. 194:87, june 17, 1948; univ. kansas sci. bull, 33:326, march 20, 1950. smith, illinois biol. mono., 32:23, may, 1964. _smilisca baudini baudini_, smith, jour. washington acad. sci., 37(11):408, nov. 15, 1947. smith and taylor, bull. u. s. natl. mus., 194:75, june 17, 1948; univ. kansas sci. bull., 33:347, march 20, 1950. brown, baylor univ. studies, p. 68, 1950. smith, smith, and werler, texas jour. sci., 4(2):254, june 30, 1952. smith and smith, anales inst. biol., 22(2):561, aug. 7, 1952. smith and darling, herpetologica, 8(3):82, nov. 1, 1952. davis and smith, herpetologica, 8(4):148, jan. 30, 1953. neill and allen, publ. res. div. ross allen's reptile inst., 2(1):26, nov. 10, 1959. maslin, univ. colorado studies, biol. series, 9:4, feb. 1963. holman, herpetologica, 20:48, april 17, 1964. _hyla manisorum_ taylor, univ. kansas sci. bull., 36:630, june 1, 1954 [holotype.--ku 34927 from batán, limón province, costa rica; edward h. taylor collector]. duellman and berg, univ. kansas publ. mus. nat. hist, 15(4):193, oct. 26, 1962. _diagnosis._--size large ([m] 76 mm., [f] 90 mm.); skull noticeably wider than long, having small frontoparietal fontanelle (roofed with bone in large individuals); postorbital processes long, pointed, curving along posterior border of orbit; squamosal large, contacting maxillary; tarsal fold strong, full length of tarsus; inner metatarsal tubercle large, high, elliptical; hind limbs relatively short, tibia length less than 55 per cent snout-vent length; lips strongly barred with brown and creamy tan; flanks pale cream with bold brown or black reticulations in groin; posterior surfaces of thighs brown with cream-colored flecks; dorsal surfaces of limbs marked with dark brown transverse bands. (foregoing combination of characters distinguishing _s. baudini_ from any other species in genus.) _description and variation._--considerable variation in size, and in certain proportions and structural characters was observed; variation in some characters seems to show geographic trends, whereas variation in other characters apparently is random. noticeable variation is evident in coloration, but this will be discussed later. in order to analyze geographic variation in size and proportions, ten adult males from each of 14 samples from various localities throughout the range of the species were measured. snout-vent length, length of the tibia in relation to snout-vent length, and relative size of the tympanum to the eye are the only measurements and proportions that vary noticeably (table 1). the largest specimens are from southern sinaloa; individuals from the atlantic lowlands of alta verapaz in guatemala, honduras, and costa rica are somewhat smaller, and most specimens from the pacific lowlands of central america are slightly smaller than those from the atlantic lowlands. the smallest males are from the atlantic lowlands of méxico, including tamaulipas, veracruz, the yucatán peninsula, and british honduras. table 1.--geographic variation in size and proportions in males of smilisca baudini. (means in parentheses below observed ranges; data based on 10 specimens from each locality.) ============================+============+==============+============ locality | snout-vent | tibia length/| tympanum/ | length | snout-vent | eye ----------------------------+------------+--------------+----------- southern sinaloa | 62.3-75.9 | 43.2-46.7 | 84.2-94.4 | (68.6) | (44.9) | (87.8) | | | ocotito, guerrero | 55.6-64.0 | 46.1-51.2 | 66.7-82.8 | (58.7) | (47.8) | (74.6) | | | pochutla, oaxaca | 56.1-65.1 | 44.7-49.4 | 73.0-84.2 | (60.2) | (47.5) | (77.4) | | | san salvador, el salvador | 57.0-68.0 | 42.1-46.1 | 74.6-83.3 | (62.1) | (44.9) | (77.6) | | | managua, nicaragua | 52.9-63.6 | 45.6-49.4 | 73.7-89.7 | (57.3) | (47.5) | (79.4) | | | esparta, costa rica | 57.6-66.0 | 44.6-49.3 | 65.5-83.6 | (61.3) | (47.3) | (75.2) | | | ciudad victoria, tamaulipas | 50.6-56.9 | 44.5-48.7 | 67.2-84.3 | (53.7) | (46.6) | (73.9) | | | córdoba, veracruz | 53.8-63.4 | 43.9-48.4 | 66.1-75.9 | (57.5) | (45.6) | (70.0) | | | isla del carmen, campeche | 47.3-56.6 | 44.7-48.9 | 61.5-72.6 | (50.9) | (47.6) | (65.7) | | | chichén-itzá, yucatán | 49.6-57.1 | 45.2-53.4 | 62.7-80.7 | (53.8) | (49.5) | (72.6) | | | british honduras | 49.0-59.6 | 47.5-50.7 | 67.9-76.8 | (54.9) | (49.1) | (72.2) | | | chinajá, guatemala | 56.8-67.6 | 47.0-51.0 | 70.0-82.8 | (63.2) | (49.5) | (73.6) | | | atlantidad, honduras | 52.5-65.1 | 49.8-53.6 | 56.1-76.5 | (57.6) | (51.5) | (67.0) | | | limón, costa rica | 57.7-71.3 | 50.4-52.3 | 63.9-73.0 | (62.4) | (51.2) | (68.5) ----------------------------+------------+--------------+-----------the ratio of the tibia to the snout-vent length varies from 42.1 to 53.6 in the 14 samples analyzed. the average ratio in samples from the pacific lowlands varies from 44.9 in sinaloa and el salvador to 47.8 in guerrero; on the gulf lowlands of méxico the average ratio varies from 45.6 in veracruz to 47.6 on isla del carmen, campeche. specimens from the yucatán peninsula and the caribbean lowlands have relatively longer legs; the variation in average ratios ranges from 49.1 in british honduras to 51.2 in costa rica and 51.5 in honduras. specimens from southern sinaloa are outstanding in the large size of the tympanum; the tympanum/eye ratio varies from 84.2 to 94.4 (average 87.8). in most other samples the variation in average ratios ranges from 72.2 to 79.3, but specimens from veracruz have an average ratio of 70.0; campeche, 65.7; honduras, 67.0; and limón, costa rica, 68.5. no noticeable geographic trends in size and proportions are evident. specimens from southern sinaloa are extreme in their large size, relatively short tibia, and large tympani, but in size and relative length of the tibia the sinaloan frogs are approached by specimens from such far-removed localities as san salvador, el salvador, and chinajá, guatemala. frogs from the caribbean lowlands of honduras and costa rica are relatively large and have relatively long tibiae and small tympani. the inner metatarsal tubercle is large and high and its shape varies. the tubercle is most pronounced in specimens from northwestern méxico, tamaulipas, and the pacific lowlands of central america. possibly the large tubercle is associated with drier habitats, where perhaps the frogs use the tubercles for digging. the ground color of _smilisca baudini_ is pale green to brown dorsally and white to creamy yellow ventrally. the dorsum is variously marked with dark brown or dark olive-green spots or blotches (pl. 6a). in most specimens a dark interorbital bar extends across the head to the lateral edges of the eyelid; usually this bar is connected medially to a large dorsal blotch. there is no tendency for the markings on the dorsum to form transverse bands or longitudinal bars. in specimens from the southern part of the range the dorsal dark markings are often fragmented into small spots, especially posteriorly. the limbs are marked by dark transverse bands, usually three on the forearm, three on the thigh, and three or four on the shank. transverse bands also are present on the tarsi and proximal segments of the fingers and toes. the webbing on the hands and feet is pale grayish brown. the loreal region and upper lip are pale green or tan; the lip usually is boldly marked with broad vertical dark brown bars, especially evident is the bar beneath the eye. a dark brown or black mark extends from the tympanum to a point above the insertion of the forearm; in some specimens this black mark is narrow or indistinct, but in most individuals it is quite evident. the flanks are pale gray to creamy white with brown or black mottling, which sometimes forms reticulations enclosing white spots. the anterior surfaces of the thighs usually are creamy white with brown mottling, whereas the posterior surfaces of the thighs usually are brown with small cream-colored flecks. a distinct creamy white anal stripe usually is present. usually, there are no white stripes on the outer edges of the tarsi and forearms. in breeding males the throat is gray. most variation in coloration does not seem to be correlated with geography. the lips are strongly barred in specimens from throughout the range of the species, except that in some specimens from southern nicaragua and costa rica the lips are pale and in some specimens the vertical bars are indistinct. six specimens from 7.3 kilometers southwest of matatán, sinaloa, are distinctively marked. the dorsum is uniformly grayish green with the only dorsal marks being on the tarsi; canthal and post-tympanic dark marks absent. a broad white labial stripe is present and interrupted by a single vertical dark mark below the eye. a white stripe is present on the outer edge of the foot. the flanks and posterior surfaces of the thighs are creamy white, boldly marked with black. two specimens from alta verapaz, guatemala (cnhm 21006 from cobán and ummz 90908 from finca canihor), are distinctive in having many narrow transverse bands on the limbs and fine reticulations on the flanks. two specimens from limón province, costa rica (ku 34927 from batán and 36789 from suretka), lack a dorsal pattern; instead these specimens are nearly uniform brown above and have only a few small dark brown spots on the back and lack transverse bands on the limbs. the post-tympanic dark marks and dark mottling on the flanks are absent. specimens lacking the usual dorsal markings are known from scattered localities on the caribbean lowlands from guatemala to costa rica. the coloration in life is highly variable; much of the apparent variation is due to metachrosis, for individuals of _smilisca baudini_ are capable of undergoing drastic and rapid change in coloration. when active at night the frogs usually are pale bright green with olive-green markings, olive-green with brown markings, or pale brown with dark brown markings. the dark markings on the back and dorsal surfaces of the limbs are narrowly outlined by black. the pale area below the eye and just posterior to the broad suborbital dark bar is creamy white, pale green, or ashy gray in life. the presence of this mark is an excellent character by which to identify juveniles of the species. the flanks are creamy yellow, or yellow with brown or black mottling. in most individuals the belly is white, but in specimens from southern el petén and northern alta verapaz, guatemala, the belly is yellow, especially posteriorly. the iris varies from golden bronze to dull bronze with black reticulations, somewhat darker ventrally. _natural history._--throughout most of its range _smilisca baudini_ occurs in sub-humid habitats; consequently the activity is controlled by the seasonal nature of the rainfall and usually extends from may or june through september. throughout méxico and central america the species is known to call and breed in june, july, and august. several records indicate that the breeding season in central america is more lengthy. gaige, hartweg, and stuart (1937:4) noted gravid females collected at el recreo, nicaragua, in august and september. schmidt (1941:486) reported calling males in february in british honduras. stuart (1958:17) stated that tadpoles were found in mid-february, juveniles in february and march and half-grown individuals from mid-march to mid-may at tikal, el petén, guatemala. stuart (1961:74) reported juveniles from tikal in july, and that individuals were active at night when there had been light rain in the dry season in february and march in el petén, guatemala. _smilisca baudini_ seeks daytime retreats in bromeliads, elephant-ear plants (_xanthosoma_), and beneath bark or in holes in trees. by far the most utilized retreat in the dry season in parts of the range is beneath the outer sheaths of banana plants. large numbers of these frogs were found in banana plants at cuautlapan, veracruz, in march, 1956, in march and december, 1959. large breeding congregations of this frog are often found at the time of the first heavy rains in the wet season. gadow (1908:76) estimated 45,000 frogs at one breeding site in veracruz. in the vicinity of tehuantepec, oaxaca, large numbers of individuals were found around rain pools and roadside ditches in july, 1956, and july, 1958; large concentrations were found near chinajá, guatemala, in june, 1960, and near esparta, costa rica in july, 1961. usually males call from the ground at the edge of the water or not infrequently sit in shallow water, but sometimes males call from bushes and low trees around the water. stuart (1935:38) recorded individuals calling and breeding throughout the day at la libertad, guatemala. _smilisca baudini_ usually is absent from breeding congregations of hylids; frequently _s. baudini_ breeds alone in small temporary pools separated from large ponds where numerous other species are breeding. in guerrero and oaxaca, méxico, _s. baudini_ breeds in the same ponds with _rhinophrynus dorsalis_, _bufo marmoreus_, _engystomops pustulosus_, and _diaglena reticulata_, and in the vicinity of esparta, costa rica, _s. baudini_ breeds in ponds with _bufo coccifer_, _hyla staufferi_, and _phrynohyas venulosa_. in nearly all instances the breeding sites of _s. baudini_ are shallow, temporary pools. the breeding call of _smilisca baudini_ consists of a series of short explosive notes. each note has a duration of 0.09 to 0.13 seconds; two to 15 notes make up a call group. individual call groups are spaced from about 15 seconds to several minutes apart. the notes are moderately high-pitched and resemble "wonk-wonk-wonk." little vibration is discernible in the notes, which have 140 to 195 pulses per second and a dominant frequency of 2400 to 2725 cycles per second (pl. 10a). the eggs are laid as a surface film on the water in temporary pools. the only membrane enclosing the individual eggs is the vitelline membrane. in ten eggs (ku 62154 from san salvador, el salvador) the average diameter of the embryos in first cleavage is 1.3 mm. and of the vitelline membranes, 1.5 mm. hatchling tadpoles have body lengths of 2.6 to 2.7 mm. and total lengths of 5.1 to 5.4 mm. the body and caudal musculature is brown; the fins are densely flecked with brown. the gills are long and filamentous. growth and development of tadpoles are summarized in table 9. a typical tadpole in stage 30 of development (ku 60018 from chinajá, alta verapaz, guatemala) has a body length of 8.7 mm., a tail length of 13.6 mm., and a total length of 22.3 mm.; body slightly wider than deep; snout rounded dorsally and laterally; eyes widely separated, directed dorsolaterally; nostril about midway between eye and tip of snout; mouth anteroventral; spiracle sinistral, located about midway on length of body and slightly below midline; anal tube dextral; caudal musculature slender, slightly curved upward distally; dorsal fin extending onto body, deepest at about one-third length of tail; depth of dorsal fin slightly more than that of ventral fin at mid-length of tail; dorsal part of body dark brown; pale crescent-shaped mark on posterior part of body; ventral surfaces transparent with scattered brown pigment ventrolaterally, especially below eye; caudal musculature pale tan with a dark brown longitudinal streak on middle of anterior one-third of tail; dorsum of anterior one-third of tail dark brown; brown flecks and blotches on rest of caudal musculature, on all of dorsal fin, and on posterior two-thirds of ventral fin; iris bronze in life (fig. 11). mouth small; median third of upper lip bare; rest of mouth bordered by two rows of conical papillae; lateral fold present; tooth rows 2/3; two upper rows about equal in length; second row broadly interrupted medially, three lower rows complete, first and second equal in length, slightly shorter than upper rows; third lower row shortest; first upper row sharply curved anteriorly in midline; upper beak moderately deep, forming a board arch with slender lateral processes; lower beak more slender, broadly v-shaped; both beaks bearing blunt serrations (fig. 15a). in tadpoles having fully developed mouthparts the tooth-row formula of 2/3 is invariable, but the coloration is highly variable. the color and pattern described above is about average. some tadpoles are much darker, such as those from 11 kilometers north of vista hermosa, oaxaca, (ku 87639-44), 3.5 kilometers east of yokdzonot, yucatán (ku 71720), and 4 kilometers west-southwest puerto juárez, quintana roo, méxico (ku 71721), whereas others, notably from 17 kilometers northeast of juchatengo, oaxaca, méxico (ku 87645), are much paler and lack the dark markings on the caudal musculature. the variation in intensity of pigmentation possibly can be correlated with environmental conditions, especially the amount of light. in general, tadpoles that were found in open, sunlit pools are pallid by comparison with those from shaded forest pools. these subjective comparisons were made with preserved specimens; detailed comparative data on living tadpoles are not available. the relative length and depth of the tail are variable; in some individuals the greatest depth of the tail is about at mid-length of the tail, whereas in most specimens the tail is deepest at about one-third its length. the length of the tail relative to the total length is usually 58 to 64 per cent in tadpoles in stages 29 and 30 of development. in some individuals the tail is about 70 per cent of the total length. on the basis of the material examined, these variations in proportions do not show geographical trends. probably the proportions are a reflection of crowding of the tadpoles in the pools where they are developing or possibly due to water currents or other environmental factors. stuart (1948:26) described and illustrated the tadpole of _smilisca baudini_ from finca chejel, alta verapaz, guatemala. the description and figures agree with ours, except that the first lower tooth row does not have a sharp angle medially in stuart's figure. he (1948:27) stated that color in tadpoles from different localities probably varies with soil color and turbidity of water. maslin (1963:125) described and illustrated tadpoles of _s. baudini_ from pisté, yucatán, méxico. these specimens are heavily pigmented like specimens that we have examined from the yucatán peninsula and from other places in the range of the species. maslin stated that the anal tube is median in the specimens that he examined; we have not studied maslin's specimens, but all tadpoles of _smilisca_ that we have examined have a dextral anal tube. newly metamorphosed young have snout-vent lengths of 12.0 to 15.5 mm. (average 13.4 in 23 specimens). the largest young are from la libertad, el petén, guatemala; these have snout-vent lengths of 14.0 to 15.5 mm. (average 14.5 in five specimens). young from 11 kilometers north of vista hermosa, oaxaca, méxico, are the smallest and have snout-vent lengths of 12.0 to 12.5 mm. (average 12.3 in three specimens). recently metamorphosed young usually are dull olive green above and white below; brown transverse bands are visible on the hind limbs. the labial markings characteristic of the adults are represented only by a creamy white suborbital spot, which is a good diagnostic mark for young of this species. in life the iris is pale gold. _remarks_: the considerable variation in color and the extensive geographic distribution of _smilisca baudini_ have resulted in the proposal of eight specific names for the frogs that we consider to represent one species. duméril and bibron (1841:564) proposed the name _hyla baudini_ for a specimen (mnhn 4798) from méxico. smith and taylor (1950:347) restricted the type locality to córdoba, veracruz, méxico, an area where the species occurs in abundance. baird (1854:61) named _hyla vanvlieti_ from brownsville, texas, and (1859:35) labelled the figures of _hyla vanvlieti_ [= _hyla baudini_] on plate 38 as _hyla vociferans_, a _nomen nudum_. cope (1862:359) named _hyla muricolor_ from mirador, veracruz, méxico, and (1865:194) used the name _smilisca daulinia_ for a skeleton that he employed as the basis for the cranial characters diagnostic of the genus _smilisca_, as defined by him. although we cannot be certain, cope apparently inadvertently used _daulinia_ for _baudini_, just as he used _daudinii_ for _baudini_ (1871:205). brocchi (1877:125) named _hyla pansosana_ from panzos, alta verapaz, guatemala. [illustration: plate 1 a b dorsal views of skulls of young _smilisca baudini_: (a) recently metamorphosed young (ku 60026), snout-vent length 12.6 mm. ×23; (b) young (ku 85438), snout-vent length 32.1 mm. ×9.] [illustration: plate 2 a b skull of adult female _smilisca baudini_ (ku 68184): (a) dorsal; (b) ventral. ×4.5.] [illustration: plate 3 a b c skull of adult female _smilisca baudini_ (ku 68184): (a) lateral; (b) dorsal view of left mandible; (c) posterior. ×4.5.] [illustration: plate 4 a b c d e f palmar views of right hands of _smilisca_: (a) _s. baudini_ (ku 87177); (b) _s. phaeota_ (ku 64276); (c) _s. cyanosticta_ (ku 87199); (d) _s. sordida_ (ku 91761); (e) _s. puma_ (ku 91716), and (f) _s. sila_ (ku 77408). ×3.] [illustration: plate 5 a b c d e f ventral aspect of right feet of _smilisca_: (a) _s. baudini_ (ku 87177); (b) _s. phaeota_ (ku 64276); (c) _s. cyanosticta_ (ku 87199); (d) _s. sordida_ (ku 91761); (e) _s. puma_ (ku 91716), and (f) _s. sila_ (ku 77408). ×3.] [illustration: plate 6 a b c living _smilisca_: (a) _s. baudini_ (ummz 115179) from 1.7 km. w xicotencatl, tamaulipas, méxico; (b) _s. cyanosticta_ (ummz 118163) from volcán san martín, veracruz, méxico; (c) _s. phaeota_ (ku 64282) from barranca del río sarapiquí, heredia prov., costa rica. all approx. nat. size.] [illustration: plate 7 a b c living _smilisca_: (a) _s. puma_ (ku 65307) from 5.9 km. w. puerto viejo, heredia prov., costa rica; (b) _s. sila_ (ku 77407) from finca palosanto, 6 km. wnw el volcán, chiriquí, panamá; (c) _s. sordida_ (ku 64257) from 20 km. wsw san isidro el general, san josé prov., costa rica. all approx. nat. size.] [illustration: plate 8 fig. 1. breeding site of _smilisca baudini_, 4 km. wnw of esparta, puntarenas prov., costa rica. fig. 2. breeding site of _smilisca phaeota_, puerto viejo, heredia prov., costa rica.] [illustration: plate 9 fig. 1. breeding site of _smilisca puma_, 7.5 km. w of puerto viejo, heredia prov., costa rica. fig. 2. breeding site of _smilisca sordida_, río la vieja, 30 km. e of palmar norte, puntarenas prov., costa rica.] [illustration: plate 10 audiospectrographs and sections of breeding calls of _smilisca_: (a) _s. baudini_ (ku tape no. 74); (b) _s. cyanosticta_ (ku tape no. 373); (c) _s. phaeota_ (ku tape no. 79).] [illustration: plate 11 audiospectrographs and sections of breeding calls of _smilisca_: (a) _s. puma_ (ku tape no. 382); (b) _s. sila_ (ku tape no. 385); (c) _s. sordida_ (ku tape no. 398).] [illustration: plate 12 lateral views of the heads of _smilisca_: (a) _s. baudini_ (ku 87177); (b) _s. sordida_ (ku 91765); (c) _s. phaeota_ (ku 64276); (d) _s. puma_ (ku 91716); (e) _s. cyanosticta_ (ku 87199); (f) _s. sila_ (ku 77408). ×3.2.] aside from the skeleton referred to as _smilisca daulinia_ by cope (1865:194), we have examined each of the types of the species synonymized with _s. baudini_. all unquestionably are representatives of _s. baudini_. taylor (1942:306) named _hyla beltrani_ from tapachula, chiapas. this specimen (uimnh 25046) is a small female (snout-vent length, 44 mm.) of _s. baudini_. taylor (1954:630) named _hyla manisorum_ from batán, limón, costa rica. the type (ku 34927) is a large female (snout-vent length, 75.3 mm.) _s. baudini_. in this specimen and a male from suretka, costa rica, the usual dorsal color pattern is absent, but the distinctive curved supraorbital processes, together with other structural features, show that the two specimens are _s. baudini_. _hyla baudini dolomedes_ barbour (1923:11), as shown by dunn (1931a:413), was based on a specimen of _smilisca phaeota_ from río esnápe, darién, panamá. [illustration: fig. 1. map showing locality records for _smilisca baudini_.] _distribution_.--_smilisca baudini_ inhabits lowlands and foothills usually covered by xerophytic vegetation or savannas, but in the southern part of its range _baudini_ inhabits tropical evergreen forest. the species ranges throughout the pacific and atlantic lowlands of méxico from southern sonora and the río grande embayment of texas southward to costa rica, where on the pacific lowlands the range terminates at the southern limits of the arid tropical forest in the vicinity of esparta; on the caribbean lowlands the distribution seems to be discontinuous southward to suretka (fig. 1). most localities where the species has been collected are at elevations of less than 1000 meters. three localities are notably higher; calling males were found at small temporary ponds in pine-oak forest at linda vista, 2 kilometers northwest of pueblo nuevo solistahuacán, chiapas, elevation 1675 meters, and 10 kilometers northwest of comitán, chiapas, at an elevation of 1925 meters. tadpoles and metamorphosing young were obtained from a pond in arid scrub forest, 17 kilometers northeast of juchatengo, oaxaca, elevation 1600 meters. stuart (1954:46) recorded the species at elevations up to 1400 meters in the south-eastern highlands of guatemala. _specimens examined._--3006, as follows: united states: texas: cameron county, brownsville, cnhm 5412-3, 6869, ummz 54036, usnm 3256. mexico: =campeche=: balchacaj, cnhm 102285, 102288, 102291, 102311, uimnh 30709-22, 30726; champotón, ummz 73172 (2), 73176, 73180; 16 km. e champotón, ummz 73181; 5 km. s champotón, ku 71369-75; 9 km. s champotón, ku 71367-8; 10.5 km. s champotón, ku 71365-6, 71722 (tadpoles), 71723 (yg.); 24 km. s champotón, ummz 73177 (2); chuina, ku 75101-3; ciudad del carmen, uimnh 30703-8; dzibalchén, ku 75413-31; encarnación, cnhm 102282, 102289, 102294-5, 102300, 102306-8, 102312, 102314, 102316-7, 102319, 102322, uimnh 30727-40, 30836-7; 1 km. w escárcega, ku 71391-6; 6 km. w escárcega, ku 71397-403; 7.5 km. w escárcega, ku 71376-89; 14 km. w escárcega, ku 71390; 13 km. w, 1 km. n escárcega, ku 71404; 3 km. n hopelchén, ku 75410-11; 2 km. ne hopelchén, ku 75412; matamoras, cnhm 36573; pitál, uimnh 30741; 1 km. sw puerto real, isla del carmen, ku 71345-64; san josé carpizo, ummz 99879; tres brazos, cnhm 102284, uimnh 30723-5; tuxpeña camp, ummz 73239. =chiapas=: acacoyagua, usnm 114487-92; 2 km. w acacoyagua, usnm 114493-4; 5 km. e arroyo minas, uimnh 9533-7; berriozabal, ummz 119186 (7); chiapa de corzo, ummz 119185 (2); cintalapa, uimnh 50077; colonia soconusco, usnm 114495-9; 5 km. w colonia soconusco, ummz 87885 (7); comitán, ummz 94438; 10 km. nw comitán, ku 57185; el suspiro, ummz 118819 (11); escuintla, ummz 88271 (7), 88278, 88327, 109233; 6 km. ne escuintla, ummz 87856 (26); 3 km. e finca juárez, uimnh 9538; finca prussia, ummz 95167; honduras, ummz 94434-7; la grada, ummz 87862; 21 km. s la trinitaria, uimnh 9540-1; 14.4 km. sw las cruces, ku 64239-44; palenque, uimnh 49286, usnm 114473-84; 2 km. nw pueblo nuevo solistahuacán, ku 57182-4, ummz 119948 (8), 121514; 1.3 km. n puerto madero, ku 57186-9; 4 km. n puerto madero, ku 57190-1; 8 km. n puerto madero, ummz 118379 (2); 12 km. n puerto madero, ku 57192; 17.6 km. n puerto madero, ummz 118378; rancho monserrata, uimnh 9531-2, ummz 102266-7; region soconusco, uimnh 33542-56; san bartola, uimnh 9519-30; san gerónimo, uimnh 30804; san juanito, usnm 114485-6; san ricardo, cnhm 102406; solosuchiapa, ku 75432-3; tapachula, cnhm 102208, 102219, 102239, 102405, uimnh 25046, 30802-3; tonolá, amnh 531, cnhm 102232, 102416, uimnh 30805-9, usnm 46760; tuina, ku 41593 (skeleton); tuxtla gutierrez, cnhm 102231, 102248; 6 km. e tuxtla gutierrez, uimnh 9539; 10 km. e tuxtla gutierrez, ummz 119949. =chihuahua=: 2.4 km. sw toquina, ku 47226-7; riito, ku 47228. =coahuila=: mountain near saltillo, uimnh 30833-4. =colima=: no specific locality, cnhm 1632; colima, amnh 510-11; hacienda albarradito, ummz 80029 (2); hacienda del colomo, amnh 6208; los mezcales, ummz 80028; manzanillo, amnh 6207, 6209; paso del río, cnhm 102207, 102229-30, uimnh 30819-21, ummz 110875 (3); periquillo, ummz 80025 (3), 80026 (14); 1.6 km. sw pueblo juárez, ummz 115564; queseria, cnhm 102204, 102216-7, 102224, uimnh 30816-8, ummz 80023 (7), 80024 (7); santiago, ummz 80027; 7.2 km. sw tecolapa, ummz 115184. =guerrero=: acahuizotla, uf 1338 (2), 1339-40, ummz 119182 (2), 119184; 3 km. s acahuizotla, ku 87183-7; acapulco, amnh 55276, ummz 121879 (4), usnm 47909; 3 km. n acapulco, ummz 110127; 8 km. nw acapulco, uf 11203 (7); 27 km. ne acapulco, uimnh 26597-610; agua del obispo, cnhm 102214, 102290, 102293, 102310, 102413, ku 60413, 87180-2, uimnh 30764-6; atoyca, ku 87175-8; buena vista, cnhm 102279, 102304, 102313, 102315, uimnh 30774; caculutla, ku 87179; 20 km. s chilpancingo, cnhm 102242, 102401, 102410-1, 102415; colonia buenas aires, ummz 119189; el limoncito, cnhm 102292, 102303, 102321, 102414; el treinte, cnhm 102212, 102221, 102237, 102240-1, uimnh 30783-5, usnm 114508-10; laguna coyuca, ummz 80960 (2); 3 km. n mazatlán, uimnh 30777-9; 9 km. s mazatlán, cnhm 102209, 102215, 102234, 102246, uimnh 30781-2; mexcala, cnhm 102399, 102403, 102409, 106539-40, uimnh 30775-6; ocotito, ku 60414-23; 5.4 km. n ocotito, ummz 119181 (4); 1.6 km. n organos, uimnh 30752-63; palo blanco, cnhm 102283, 102286, 102305, 102320, 102404, uimnh 30767-70; pie de la cuesta, amnh 55275, 59202-5; puerto marquéz, amnh 59200-1 (13); 5.6 km. s san andreas de la cruz, ku 87173-4; san vincente, ku 87172; zaculapán, ummz 119183. =hidalgo=: below tianguistengo, cnhm 102318. =jalisco=: atenqueque, ku 91435-6; 5 km. ne autlán, uimnh 30810; 5 km. e barro de navidad, ummz 110900; charco hondo, ummz 95247; puerto vallarta, uimnh 41346; between la huerta and tecomates, ku 91437; 3 km. se la resolana, ku 27619, 27620 (skeleton); 11 km. s, 1.6 km. e yahualica, ku 29039; zapotilitic, cnhm 102238. =michoacán=: aguililla, ummz 119179 (5); apatzingán, cnhm 38766-90, ku 69101 (skeleton); 7 km. e apatzingán, ummz 112843; 11 km. e apatzingán, ummz 112841 (3); 27 km. s apatzingán, ku 37621-3; 1.6 km. n arteaga, ummz 119180; charapendo, ummz 112840; coahuayana, ummz 104458; el sabino, cnhm 102205-6, 102210-1, 102220, 102228, 102233, uimnh 30822-3; la placita, ummz 104456; la playa, ummz 105163; 30 km. e nueva italia, ummz 120255 (2); 4 km. s nueva italia, ummz 112842; ostula, ummz 104457 (4); salitre de estopilas, ummz 104459; san josé de la montaña, ummz 104461 (2); 11 km. s tumbiscatio, ku 37626; 12 km. s tzitzio, ummz 119178. =morelos=: 3.5 km. w cuautlixco, ku 87188-90; 1 km. ne puente de ixtla, ku 60393-4; 20 km. s puente de ixtla, cnhm 102400, uimnh 30832; tequesquitengo, amnh 52036-9. =nayarit=: 3 km. s acaponeta, ummz 123030 (4); 56 km. s esquinapa (sinaloa), ku 73909; jesús maría, amnh 58239; san blas, ku 28087, 37624, 62360-2, usnm 51408; 8.6 km. e san blas, ummz 115185; tepic, uimnh 30812-5; 4 km. e tuxpan, ku 67786; 11 km. se tuxpan, uimnh 7329-31, 7335-59. =nuevo león=: galeana, cnhm 34389; salto de cola de caballo, cnhm 30628-31, 30632 (40), 30633-7, 34454-67. =oaxaca=: 11 km. s candelaria, uimnh 9515-8; cerro san pedro, 24 km. sw tehuantepec, ummz 82156; chachalapa, ku 38199; 8 km. s chiltepec, ku 87191; 12 km. s chivela, ummz 115182; coyul, usnm 114512; garza mora, uimnh 40967-8; juchatengo, ku 87193; 17 km. ne juchatengo, ku 87645 (tadpoles), 87646 (young); juchitán, usnm 70400; lagartero, uimnh 9514; matías romero, amnh 52143-5; 25 km. n matías romero, ku 33822-8; 7 km. s matías romero, uimnh 42703; mirador, amnh 6277, 13832-9, 13842-55; mira león, 1.6 km. n huatulco, uimnh 9503-4; mixtequillo, amnh 13924; pochutla, ku 57167-81, uimnh 9505-13; quiengola, amnh 51817, 52146; río del corte, uimnh 48677; río mono blanco, uimnh 36831; río sarabia, 5 km. n sarabia, ummz 115180 (4); 2.5 km. n salina cruz, ku 57165-6; san antonio, uimnh 37286; 5 km. nnw san gabriel mixtepec, ku 87192; san pedro del istmo, uimnh 37197; santo domingo, usnm 47120-2; 3.7 km. n sarabia, ummz 115181 (3); tapanatepec, ku 37793 (skeleton), 37794, uimnh 9542, ummz 115183; between tapanatepec and zanatepec, uimnh 42704-25; tecuane, ummz 82163 (3); tehuantepec, amnh 52625, 52639, 53470, ummz 82157-8, 82159 (9), 82160 (4), 82161 (8), 82162 (12), 112844-5, 118703, usnm 10016, 30171-4, 30188; 4.5 km. w tehuantepec, ku 59801-12 (skeletons), 69102-3 (skeletons); 10 km. s tehuantepec, ku 57163-4; temazcal, usc 8243 (3); 3 km. s tolocita, ku 39666-9; tolosa, amnh 53605; tuxtepec, ummz 122098 (2); 2 km. s valle nacional, ku 87194-5; 11 km. n vista hermosa, ku 87196, 87639-41 (tadpoles), 87642-3 (young), 87644 (tadpoles); yetla, ku 87197. =puebla=: 16 km. sw mecatepec (veracruz), uimnh 3657-8; san diego, amnh 57714, usnm 114511; vegas de suchil, amnh 57712; villa juárez, uf 11205. =quintana roo=: cóba, cnhm 26937; esmeralda, ummz 113551; 4 km. nne felipe carrillo puerto, ku 71417-8; pueblo nuevo x-can, ku 71405; 10 km. ene pueblo nuevo x-can, ku 71406; 4 km. wsw puerto juárez, ku 71407-11, 71721 (tadpoles); 12 km. w puerto juárez, ku 71412-6; san miguel, isla de cozumel, ummz 78542 (6), 78543 (10), 78544 (2); 3.5 km. n san miguel, isla de cozumel, ku 71419-22; 10 km. e san miguel, isla de cozumel, ummz 78541; telantunich, cnhm 26950. =san luis potosí=: ciudad valles, amnh 57776-81 (12), cnhm 37193, 102297, ku 23705; 21 km. n ciudad valles, ummz 118377; 6 km. e ciudad valles, uf 3524; 24 km. e ciudad valles, uf 7340 (2); 5 km. s ciudad valles, uimnh 30751; 16 km. s ciudad valles, amnh 52953; 30 km. s ciudad valles, cnhm 102394, 102402, 102412, uimnh 30749-50; 63 km. s ciudad valles, uimnh 19247-58; pujal, ummz 99872 (2); río axtla, near axtla, amnh 53211-5, 59516, ku 23706; tamazunchale, amnh 52675, cnhm 39621-2, 102226, 102281, uf 7615 (2), uimnh 26596, ummz 99506 (9), 118701 (2), usnm 114468; 17 km. n tamazunchale, uimnh 3659; 2.4 km. s tamazunchale, amnh 57743; 17 km. e tamuin, uf 11202 (2); xilitla, uimnh 19259-60. =sinaloa=: 8 km. n. carrizalejo, ku 78133; 4 km. ne concordia, ku 73914; 5 km. sw concordia, ku 75438-9; 6 km. e cosalá, ku 73910; costa rica, 16 km. s. culiacán, uimnh 34887-9; 51 km. sse culiacán, ku 37792; el dorado, ku 60392; 1.6 km. ne el fuerte, cnhm 71468; isla palmito del verde, middle, ku 73916-7; 21 km. nne los mochis, uimnh 40536-7; matatán, ku 73913; 7.3 km. sw matatán, ku 78464, 78466-70; mazatlán, amnh 12562, ummz 115197 (3); 57 km. n mazatlán, uimnh 38364; plomosas, usnm 47439-40; presidio, uimnh 30811, usnm 14082; rosario, ku 73911-2; 5 km. e rosario, uimnh 7360-76; 8 km. sse rosario, ku 37625; 5 km. sw san ignacio, ku 78465; 1.6 km. ene san lorenzo, ku 47917-24; teacapán, isla palmito del verde, ku 73915; 9.6 km. nnw teacapán, ku 91410; villa unión, ku 78471; 9 km. ne villa unión, ku 75434-7; 1 km. w villa unión, amnh 59284. =sonora=: guiracoba, amnh 51225-38 (25). =tabasco=: 4 km. ne comalcalco, amnh 60313; teapa, ummz 119943; 5 km. n teapa, ummz 119940, 119944, 122997 (2); 10 km. n teapa, ummz 119187, 119188 (2); 13 km. n teapa, ummz 119941 (2), 119945 (3), 120254 (2); 21 km. n teapa, ummz 119942, 119947; 29 km. n teapa, ummz 119946 (11); tenosique, usnm 114505-7. =tamaulipas=: acuña, ummz 99864; 5 km. s acuña, ummz 101180; 13 km. n antiguo morelos, uimnh 40532-5; 3 km. s antiguo morelos, uf 11204; 3 km. ne chamal, ummz 102867; 1.6 km. e chamal, ummz 110734; ciudad mante, ummz 80957, 80958 (3), 106400 (3); 16 km. n ciudad victoria, cnhm 102408; 34 km. ne ciudad victoria, ku 60395-411; 8.8 km. s ciudad victoria, uimnh 19261-3; 11 km. w ciudad victoria, uimnh 30924; 16 km. w ciudad victoria, uimnh 30825; 3 km. w el carizo, ummz 111279; gómez farías, ummz 110837-8; 8 km. ne gómez farías, ummz 102265, 102916 (4), 102917, 104110 (5), 105493, 110836 (2), 111274-7; 8 km. nw gómez farías, ummz 101178 (7), 101179 (3), 101362-3, 101364 (2), 108799 (2), 110129, 111278, 111280; 8 km. w gómez farías, ummz 102859 (2); 16 km. w gonzales, ku 37795-6; jiménez, ku 60412; la clementina, 6 km. w forlan, usnm 106244; limón, uimnh 30831; llera, usnm 140137-40; 3 km. e llera, uimnh 16858; 21 km. s llera, uimnh 30828-9; 23 km. s llera, uimnh 30830; 11 km. sw ocampo, ummz 118956; 22 km. w, 5 km. s piedra, ku 37568-71; rio sabinas, ummz 97976; 5 km. w san gerardo, ummz 110733 (2); santa barbara, ummz 111272-3; villagrán, cnhm 102280, 102287, 102299, 102309, uimnh 30826-7; 1.7 km. w xicotencatl, ummz 115179. =veracruz=: 1.6 km. nw acayucan, ummz 115189; 28.5 km. se alvarado, ummz 119933; 2.4 km. ssw amatitlán, ummz 115195; barranca metlac, uimnh 38365; boca del río, uimnh 26619-30, ummz 74954 (9); 16 km. s boca del río, uimnh 26631; between boca del río and anton lizardo, uimnh 42701; canadá, cnhm 102397; catemaco, ummz 118702 (4); ciudad alemán, ummz 119608 (3); córdoba, cnhm 38665-7, usnm 30410-3; 5.2 km. ese córdoba, ku 71423-35, 89924 (skeleton); 7 km. ese córdoba, ummz 115176 (4); cosamaloapan, ummz 115193 (2); coyame, uimnh 36853-6, 38366, ummz 111461 (3), 111462-3; 1 km. se coyame, ummz 121202 (3); cuatotolapam, ummz 41625-39; cuautlapan, cnhm 38664, 70591-600, 102218, 102398, ku 26300, 26302, 26309, 26312-3, 26315-6, 26321, 26336, 26339, 26347 (skeleton), 26354, 55614-21 (skeletons), uimnh 11236-67, 11269-71, 11273, 26611-8, 30792-5, ummz 85466 (6), 115173 (25), 115175 (7), usnm 114433-57; dos ríos, cnhm 39623; 5 km. ene el jobo, ku 23843, 23845, 23847; 6.2 km. e encero, uimnh 30835; escamilo, cnhm 102298, uimnh 30788; 1 km. n fortín, uf 11201; 4 km. sw huatusco, ummz 115177; 1 km. sw huatusco, ummz 123119; 10 km. se hueyapan, ummz 115190; 20 km. s jesús carranza, ku 23844, 23846, 27399; 38 km. se jesús carranza, ku 23417; laguna catemaco, ummz 119932 (62); 1.6 km. n la laja, uimnh 3651; la oaxaqueña, amnh 43930-1; 17 km. e martínez de la torre, uimnh 36630-2; 6.2 km. w martínez de la torre, uimnh 3652-4; minatitlán, amnh 52141-2; mirador, usnm 25097-8, 115178; 6 km. s monte blanco, uf 11200 (4); 21 km. e nanchital, ummz 123004; 2 km. s naranja, ummz 115188 (3); 1.6 km. ne novillero, ummz 115194 (2); 3 km. ne novillero, ummz 115196; 5.2 km. ne novillero, ummz 115192 (4); 6 km. ne novillero, ummz 115191; 5 km. n nuevo colonia, ummz 105066; orizaba, usnm 16563-6; 4 km. ne orizaba, ummz 120251 (2); panuco, ummz 118922; paraje nuevo, ummz 85465 (2), 85467 (35), 85468 (36); paso del macho, uimnh 49281; paso de talaya, jicaltepec, usnm 32365, 84420; pérez, cnhm 1686 (5); 20 km. n piedras negras, río blanco, ku 23708; plan del río, ku 26310, 26333-5, 26345, 26354, ummz 102069, 102070 (5); potrero, uimnh 49282-5, ummz 88799, 88805, 88806 (2), usnm 32391-5; potrero viejo, cnhm 102296, ku 26301, 26304-5, 26307-8, 26311, 26317-20, 26323-25, 26326-8 (skeletons), 26329-31, 26332 (skeleton), 26337-8, 26340-4, 26346, 26348, 26351, 26353, 27400-12, uimnh 30800, ummz 88800 (2), 88802 (15), 88803 (9), 88804, usnm 114458-67; 5 km. s potrero viejo, ku 26303, 26314, 26322; puente nacional, uimnh 21783-8; 3 km. n rinconada, ummz 122099 (5); río de las playas, usnm 118635-6; río seco, ummz 88801 (9); rodriguez clara, cnhm 102225; san andrés tuxtla, cnhm 102213, 102222, 102227, 102247, uimnh 30789-91; 10 km. nw san andrés tuxtla, ummz 119935; 13.4 km. nw san andrés tuxtla, ummz 119939 (2); 19.8 km. nw san andrés tuxtla, ummz 119938; 27.2 km. nw san andrés tuxtla, ummz 119936 (6); 48 km. nw san andrés tuxtla, ummz 119937; 4 km. w san andrés tuxtla, ummz 115187; 37.4 km. s san andrés tuxtla, ummz 119934 (12); 15 km. ese san juan de la punta, ku 23707; san lorenzo, usnm 123508-12; 3 km. sw san marcias ku 23841; 1.5 km. s santa rosa, uimnh 42702; 2 km. s santiago tuxtla, ummz 121201 (4); sauzel, ummz 121239; 14 km. e suchil, uimnh 46880; 15 km. s tampico (tamaulipas), ummz 103322 (4); 4 km. n tapalapan, ummz 115186 (2); tecolutla, uimnh 42677-700; 16 km. nw tehuatlán, uimnh 3660-3; 5 km. s tehuatlán, ku 23842; teocelo, ku 26306; tierra colorado, cnhm 102393, 102395-6, uimnh 30789-91; veracruz, amnh 6301-4, 59398-402, uimnh 30801, ummz 115174, 122060 (2); 24 km. w veracruz, cnhm 104570-2. =yucatán=: no specific locality, cnhm 548, 49067, usnm 32298; chichén-itzá, cnhm 20636, 26938-49, 36559-62, uimnh 30742-6, ummz 73173 (6), 73174 (14), 73175 (14), 73178-9, 76171, 83107 (2), 83108, 83109 (2), 83915 (30), usnm 72744; 9 km. e chichén-itzá, ku 71438-9; 12 km. e chichén-itzá, ku 71440; mérida, cnhm 40659-66, uimnh 30747-8, ummz 73182; 6 km. s mérida, ku 75194; 8.8 km. se ticul, ummz 114296; valladolid, cnhm 26934-6; xcalah-op, cnhm 53906-14; 3.5 km. e yokdzonot, ku 71441-3, 71720 (tadpoles). british honduras: belize, cnhm 4153, 4384-5, 4387, ummz 75310, usnm 26065; bokowina, cnhm 49064-5; cocquercote, ummz 75331 (2); cohune ridge, ummz 80738 (15); double falls, cnhm 49066; el cayo, ummz 75311; 6 km. s el cayo, mcz 37856; gallon jug, mcz 37848-55; manatee, cnhm 4264-7; mountain pine ridge, mcz 37857-8; san augustin, ummz 80739; san pedro, columbia, mcz 37860-2; valentin, ummz 80735 (4), 80736 (2), 80737 (2); 5 km. s waha loaf creek, mcz 37859. guatemala: =alta verapaz=: 5.1 km. ne campur, ku 68464 (tadpoles), 67465 (young); 28.3 km. ne campur, ku 64203-22, 68183-4 (skeletons); chamá, mcz 15792-3, ummz 90895 (7), 90896 (5), 90897 (29), 90898 (12), 90899; chinajá, ku 55939-41, 57193-8, 60018-20 (tadpoles), 60021 (eggs), 60022 (tadpoles); cobán, cnhm 21006; cubilquitz, ummz 90902 (10); finca canihor, ummz 90908; finca chicoyou, ku 57246-8, 60026 (young), 64202, 68466-7 (tadpoles); finca los alpes, ku 64197-201, 68463 (tadpoles); finca los pinales, ummz 90903 (2); finca tinajas, byu 16031; finca volcán, ummz 90905 (4), 90906-7; panzos, mnhn 6313, ummz 90904; samac, ummz 90900; samanzana, ummz 90901 (6). =baja verapaz=: chejel, ummz 90909 (7), 90910 (3); san gerónimo, ummz 84076 (16). =chiquimula=: 1.6 km. se chiquimula, ummz 98112; esquipulas, ummz 106793 (28). =el petén=: 20 km. nnw chinajá (alta verapaz), ku 57199-240; flores, ummz 117985; la libertad, ku 60024 (young), ummz 75313-20, 75323 (2), 75324 (7), 75325 (13), 75326 (2), 75327 (11), 75328 (12), 75329 (2); 3 km. se la libertad, ku 57243-4; 13 km. s la libertad, mcz 21458 (2); pacomon, usnm 71334; piedras negras, usnm 114469-71; poptún, ummz 120475; poza de la jicotea, usnm 114672; ramate-yaxha trail, ummz 75321; río de la pasión between sayaxché and subín, ku 57151; río san román, 16 km. nnw chinajá (alta verapaz), ku 55942-6; sacluc, usnm 25131; sayaxché, ku 57144-5; tikal, ummz 117983 (7), 117984 (5), 117993 (5), 120474 (5); toocog, ku 57241-2, 60023 (young), 60025 (young); uaxactún, ummz 70401-3; yaxha, ummz 75322; 19 km. e yaxha, ummz 75330 (4). =el quiché=: finca tesoro, ummz 89166 (3), 90549 (tadpoles). =escuintla=: río guacalate, masagua, usnm 125239; tiquisate, ummz 98262 (7). =guatemala=: 16 km. ne guatemala, ku 43545-53. =huehuetenango=: finca san rafael, 16 km. se barillas, cnhm 40912-6; 45 km. wnw huehuetenango, ku 64223-4; jacaltenango, ummz 120080 (6), 120081 (14), 120082 (13). =izabál=: 2 km. sw puerto matías de gálvez, ku 60027-8 (tadpoles); quiriguá, cnhm 20587, ummz 70060. =jalapa=: jalapa, ummz 98109, 106792 (11). =jutiapa=: finca la trinidad, ummz 107728 (10); jutiapa, ummz 106789; 1.6 km. se mongoy, ku 43069; santa catarina mita, ummz 106790. =progreso=: finca los leones, ummz 106791. =quetzaltenango=: coatepeque, amnh 62204. =retalhueleu=: casa blanca, ummz 107725 (18); champerico, ummz 107726 (3). =san marcos=: talisman bridge, usnm 128056-7. =santa rosa=: finca la guardiana, ummz 107727 (6); finca la gloria, ummz 107724 (6); 1.6 km. wsw el molino, ku 43065-8. el salvador: =la libertad=: 16 km. nw santa tecla, ku 43542-4. =morazán=: divisadero, usnm 73284. =san salvador=: san salvador, cnhm 65087-99, ku 61955-88, 62138-9 (skeletons), 62154 (eggs), 62155-60 (tadpoles), 68462 (tadpoles), ummz 117586 (3), 118380 (3), usnm 140278. honduras: state unknown: guaimas, ummz 58391. =atlantidad=: isla de roatán, cnhm 34551-4; la ceiba, usnm 64985, 117589-91; lancetilla, mcz 16207-11; tela, mcz 15774-5, 28080, ummz 58418, usnm 82173-4. =choluteca=: 1.5 km. nw choluteca, ku 64228-32; 10 km. nw choluteca, ku 64233; 10 km. e choluteca, ku 64226-7; 12 km. e choluteca, ku 64225; 5 km. s choluteca, usc 2700 (2). =colón=: bambú, uf 320; belfate, amnh 45692-5; patuca, usnm 20261. =comayagua=: la misión, 3.5 leagues n siguatepeque, mcz 26424-5. =copán=: copán, ummz 83026 (2). =cortés=: cofradía, amnh 45345-6; hacienda santa ana, cnhm 4724-31; lago de yojoa, mcz 26410-1; río lindo, amnh 54972. =el paraiso=: el volcán, mcz 26436. =francisco morazán=: tegucigalapa, byu 18301-4, 18837-41, mcz 26395-7, usnm 60499. =gracias a dios=: río segovia, mcz 24543. =santa barbara=: santa barbara, usnm 128062-5. nicaragua: =chinandega=: 4 km. n, 2 km. w chichigalpa, ku 85385; chinandega, mcz 2632; río tama, usnm 40022; san antonio, ku 84944-9 (skeletons), 85386-403. =chontales=: 1 km. ne acoyapa, ku 64234. =estelí=: finca daraili, 5 km. n, 15 km. e condega, ku 85404-8; finca venecia, 7 km. n, 16 km. e condega, ku 85409. =león=: 1.6 km. ene poneloya, ku 43084-5. =managua=: managua, usnm 79989-90; 8 km. nw managua, ku 43094-110; 20 km. ne managua, ku 85412; 21 km. ne managua, ku 85413-4; 5 km. sw managua, ku 43086-93; 2 km. n sabana grande, ku 85411; 3 km. n sabana grande, ku 43070-8; 20 km. s, 0.5 km. w tipitapa, ku 85410. =matagalpa=: guasqualie, ummz 116493; matagalpa, ummz 116492; 19 km. n matagalpa, ummz 116494. =río san juan=: greytown, usnm 19585-6, 19767-8. =rivas=: javillo, ummz 123001; moyogalpa, isla ometepe, ku 85428-37, 87706 (tadpoles); peñas blancas, ku 85417; río javillo, 3 km. n, 4 km. w sapoá, ku 85418-20, 85438 (skeleton); 13.1 km. se rivas, ku 85415; 14.8 km. se rivas, ku 85421-3; 11 km. s, 3 km. e rivas, ku 85416; 16 km. s rivas, mcz 29009-10; 7.7 km. ne san juan del sur, ku 85426-7; 16.5 km. ne san juan del sur, ku 85424-5, 87705 (young); 5 km. se san pablo, ku 43079-83. =zelaya=: cooley, amnm 7063-8, 8019-20, 8022, 8034-5; cukra, amnh 8016-7; musahuas, río huaspuc, amnh 58428-31; 11 km. nw rama, río siquia, ummz 79708, 79709 (5), 79710 (2); río escondido, usnm 19766, 20701; río siquia at río mico, ummz 79707 (10); sioux plantation, amnh 7058-61, 8023-33. costa rica: =alajuela=: los chiles, amnh 54639; orotina, mcz 7960-1; san carlos, usnm 29991. =guanacaste=: la cruz, usc 8232 (3); 4.3 km. ne la cruz, ummz 123002; 18.4 km. s la cruz, usc 8136; 23.5 km. s la cruz, usc 8094 (4); 3 km. w la cruz, usc 8233 (4); 2 km. ne las cañas, ku 64235-7; las huecas, ummz 71212-3; liberia, ku 36787, usc 8161; 11.5 km. n liberia, usc 8149; 13 km. n liberia, usc 8139; 22.4 km. n liberia, usc 8126; 8 km. nnw liberia, ku 64238; 8.6 km. ese playa del coco, usc 8137; 21.8 km. ese playa del coco, usc 8138; río piedra, 1.6 km. w bagaces, usc 7027; río bebedero, 5 km. s bebedero, ku 64158; 5 km. ne tilarán, ku 36782-6. =heredia=: 13 km. sw puerto viejo, ku 64142-6. =limón=: batán, ku 34927; guacimo, usc 621; pandora, usc 505 (3); suretka, ku 36788-9; tortugero, uf 7697, 10540-2. =puntarenas=: barranca, cnhm 35254-6; 15 km. wnw barranca, ku 64155-7, ummz 118381; 18 km. wnw barranca, ummz 118382 (4); 4 km. wnw esparta, ku 64159-96, 68178-82 (skeletons); 19 km. nw esparta, ku 64147-54. =smilisca cyanosticta= (smith), new combination _hyla phaeota_, taylor, univ. kansas sci. bull., 28(5):80, may 15, 1942. taylor and smith, proc. u. s. natl. mus., 95(3185):589, june 30, 1945. _hyla phaeota cyanosticta_ smith, herpetologica, 8:150, jan. 30, 1953 [holotype.--usnm 111147 from piedras negras, el petén, guatemala; hobart m. smith collector]. shannon and werler, trans. kansas acad. sci., 58:386, sept. 24, 1955. poglayen and smith, herpetologica, 14:11, april 25, 1958. cochran, bull. u. s. natl. mus., 220:57, 1961. smith, illinois biol. mono., 32:25, may, 1964. _smilisca phaeota cyanosticta_, stuart, misc. publ. mus. zool. univ. michigan, 122:42, april 2, 1963. duellman, univ. kansas publ. mus. nat. hist., 15(5):229, oct. 4, 1963. _diagnosis._--size moderately large ([m] 56.0 mm., [f] 70.0 mm.); skull as long as wide; frontoparietal fontanelle large; narrow supraorbital flanges having irregular margins anteriorly; large squamosal not in contact with maxillary; tarsal fold moderately wide, full length of tarsus; inner metatarsal tubercle moderately large, low, flat, elliptical; hind limbs relatively long; tibia usually more than 52 per cent of snout-vent length; labial stripe silvery-white; lips lacking vertical bars; loreal region pale green; pale bronze-colored stripe from nostril along edge of eyelid to point above tympanum narrow, bordered below by narrow dark brown stripe from nostril to eye, and broad dark brown postorbital mark encompassing tympanum and terminating above insertion of arm; flanks, dark brown with large pale blue spots; anterior and posterior surfaces of thighs dark brown with small pale blue spots on thighs. (foregoing combination of characters distinguishing _s. cyanosticta_ from any other species in genus.) _description and variation._--the largest males are from piedras negras, el petén, guatemala, and they average 52.5 mm. in snout-vent length whereas males from los tuxtlas, veracruz, average 50.6 mm. and those from northern oaxaca 50.3 mm. the smallest breeding male has a snout-vent length of 44.6 mm. the average ratio of tibia length to snout-vent length is 54.8 per cent in males from piedras negras, and 56.4 and 56.3 per cent in males from los tuxtlas and oaxaca, respectively. the only other character showing noticeable geographic variation is the size of the tympanum. the average ratio of the diameter of the tympanum to the diameter of the eye is 76.3 per cent in males from piedras negras, 71.8 from oaxaca, and 66.9 from los tuxtlas. the dorsal ground color of _smilisca cyanosticta_ is pale green to tan and the venter is creamy white. the dorsum is variously marked with dark olive-green or dark brown spots or blotches (pl. 6b). an interorbital dark bar usually is present. the most extensive dark area is a v-shaped mark in the occipital region with the anterior branches not reaching the eyelids; this mark is continuous, by means of a narrow mid-dorsal mark, with an inverted v-shaped mark in the sacral region. in many specimens this dorsal marking is interrupted, resulting in irregular spots. in some specimens the dorsum is nearly uniform pale green or tan with a few small, dark spots. the hind limbs are marked by dark transverse bands, usually three or four each on the thigh and shank, and two or three on the tarsus. the webbing on the feet is brown. the loreal region is pale green, bordered above by a narrow, dark brown canthal stripe extending from the nostril to the orbit, which is bordered above by a narrow, bronze-colored stripe extending from the nostril along the edge of the eyelid to a point above the tympanum. the upper lip is white. a broad dark brown mark extends posteriorly from the orbit and encompasses the tympanum to a point above the insertion of the forelimb. the flanks are dark brown with many pale blue, rounded spots, giving the impression of a pale blue ground color with dark brown mottling enclosing spots. the anterior and posterior surfaces of the thighs are dark brown with many small pale blue spots. the inner surfaces of the shank and tarsus are colored like the posterior surfaces of the thighs. pale blue spots are usually present on the proximal segments of the second and third toes. a distinct white stripe is present on the outer edge of the tarsus and fifth toe; on the tarsus the white stripe is bordered below by dark brown. a white stripe also is present on the outer edge of the forearm and fourth finger. the anal region is dark brown, bordered above by a narrow transverse white stripe. the throat in breeding males is dark, grayish brown with white flecks. no geographic variation in the dorsal coloration is evident. specimens from the eastern part of the range (piedras negras and chinajá, guatemala) have bold, dark reticulations on the flanks enclosing large pale blue or pale green spots, which fade to tan in preservative. specimens from oaxaca and veracruz characteristically have finer dark reticulations on the flanks enclosing smaller blue spots; in many of these specimens the ventrolateral spots are smallest and are white. all living adults are easily recognized by the presence of pale, usually blue, spots on the flanks and thighs. individuals under cover by day have a tan dorsum with dark brown markings. a hiding individual at chinajá, alta verapaz, guatemala (ku 55936), had a pale tan dorsum when found; later the dorsal color changed to chocolate brown. a pale green patch was present below the eye; the spots on the posterior surfaces of the thighs were pale blue, and those on the flanks were yellowish green. a calling male obtained 10 kilometers north-northwest of chinajá (ku 55934) was reddish brown when found at night; later the dorsal color changed to pale tan. a green patch below the eye was persistent. usually these frogs are green at night. the coloration of an adult male (ku 87201) from 11 kilometers north of vista hermosa, oaxaca, méxico, was typical: "when calling dorsum pale green; later changed to dull olive-green. flanks dark brown with pale blue spots in axilla and groin and bluish white flecks on mid-flank. anterior and posterior surfaces of thighs, inner surfaces of shanks, and median dorsal surfaces of tarsi dark brown with blue spots. canthal and postorbital stripes dark chocolate brown; labial stripe creamy white. forearm, tarsal, and anal stripes pale cream-color. throat dark brown with yellow flecks; belly and ventral surfaces of limbs creamy buff; webs pinkish tan; iris deep bronze, brown below pupil." (duellman, field notes, june 24, 1964.) some individuals have both green and brown coloration in life. an individual obtained at night on the south slope of volcán san martín, veracruz, méxico, had a pale tan dorsum changing peripherally to pale green. the dorsal markings were dark brown and dark olive-green. in contrast to the color-changes noted above, the labial region below the eye is always pale green, and pale spots are always present on the flanks and thighs in adults. the iris is invariably golden or bronze above and darker, usually brown, below. minute black flecks are present on the iris, and in some individuals these flecks are so numerous that the eye appears gray. recently metamorphosed young have pale tan flanks, and the posterior surfaces of the thighs are orange-yellow; pale spots are absent. a juvenile (ku 55935) from chinajá, alta verapaz, guatemala, having a snout-vent length of 35.0 mm. was pale yellowish tan above with olive-green markings; the flanks were dark brown with pale blue spots, and the anterior and posterior surfaces of the thighs were uniform bright tomato red. a juvenile (ummz 121298), 18.6 mm. in snout-vent length, from the southeast slope of volcán san martín, veracruz, méxico, had pale tan flanks lacking blue spots, but had red thighs. apparently the ontogenetic changes in coloration proceed as follows: (1) flanks pale tan and thighs orange-yellow, both lacking spots, (2) flanks pale tan and thighs red, both lacking spots, (3) flanks dark brown with blue spots and thighs red, lacking spots, and (4) flanks and thighs dark brown, both having pale blue spots. _natural history._--_smilisca cyanosticta_ inhabits humid tropical forest and cloud forest from the lowlands to elevations of about 1200 meters in los tuxtlas and to about 900 meters in northern oaxaca. in these moist environments the frogs apparently are active throughout the year. active individuals have been obtained in january, july, and august in los tuxtlas, veracruz, in june and july in northern oaxaca, in february and march at chinajá, guatemala, and taylor and smith reported (1945:589) activity in may at piedras negras, guatemala. calling males were observed as follows; in a rain barrel 11 kilometers north of vista hermosa, oaxaca, méxico, on june 23-28, 1964; in a quiet pool in a stream 8 kilometers south of yetla, oaxaca, méxico, in july, 1963 (dale l. hoyt, personal communication); in and near springs flowing into a stream at dos amates, veracruz, méxico, on august 4, 1959 (douglas robinson, personal communication); and in a water-filled depression in a log 10 kilometers west-northwest of chinajá, guatemala, on march 13, 1960. taylor and smith (1945:589) reported that individuals were found at night on the ground at the edge of temporary pools at piedras negras, guatemala, on may 28-29, 1939. a clasping pair was found on a rock at the edge of a small stream on the south slope of volcán san martín, veracruz, méxico, on july 11, 1959 (douglas robinson, personal communication). only one individual has been observed in a tree at night. in the daytime, individuals were found in elephant ear plants (_xanthosoma_) at chinajá, guatemala. the breeding call consists of one or two moderately short notes that are lower pitched than those of _s. baudini_, but higher pitched than those of _s. phaeota_. each note has a duration of 0.25 to 0.45 seconds and is repeated at intervals of one-half minute to several minutes. each note is a vibrant "waunk," having 110 to 180 pulses per second and dominant frequency of 1600 to 2100 cycles per second (pl. 10b). apparently the eggs are deposited as loose clumps in the water. in eggs in the yolk plug stage of development, the diameter of the embryo is about 2.3 mm.; that of the outer envelope is 4.0 mm. hatchling tadpoles have total lengths of 5.8 to 6.5 mm. and body lengths of 2.8 to 3.1 mm. the external gills are moderately long, slender, and filamentous; the yolk sac is still moderately large. the body and anterior part of the caudal musculature are dark brown; posteriorly the caudal musculature is pale brown. the caudal fins are creamy tan. the oral discs are large and ovoid. the growth of the tadpole is summarized in table 10. a typical tadpole in stage 30 of development (ku 87652 from 11 km. n vista hermosa, oaxaca, méxico) can be described as follows: body length 9.5 mm.; tail length 15.5 mm.; total length 25.0 mm.; body slightly wider than deep; snout rounded laterally, broadly ovoid dorsally; eyes widely separated, directed dorsolaterally; nostril about midway between eye and tip of snout; mouth anteroventral; spiracle sinistral, slightly posterior to midpoint of body and slightly below midline; anal tube dextral; caudal musculature slender, barely curved upward distally; dorsal fin not extending onto body, depth of dorsal fin slightly more than that of ventral fin on mid-length of tail; dorsal part of body dark brown; ventral surfaces transparent, lacking pigment; posterior edge of body pale cream-color; caudal musculature creamy white with interconnected brown spots; caudal fins transparent with small brown blotches on dorsal fin and posterior half of ventral fin; iris coppery bronze in life (fig. 12). mouth small, median part of upper lip bare; rest of mouth bordered by single row of bluntly rounded papillae; lateral fold present; tooth rows 2/3; all tooth-rows approximately equal in length; second upper row broadly interrupted medially; other rows complete; upper beak moderately deep, forming broad arch with slender lateral processes; lower beak slender, broadly v-shaped; both beaks finely serrate (fig. 15c). all tadpoles having fully developed mouth parts have 2/3 tooth rows. little variation is noticeable in coloration. in many specimens the posterior edge of the body is dark brown instead of pale cream-color. mottling is rather dense on the caudal fins in all specimens; in some individuals pigment is concentrated along the anterior one-third of the lateral groove. in life the body is dark brown with greenish gold flecks ventrally; the caudal musculature is gray. in each of two recently metamorphosed young the snout-vent length is 14.0 mm. coloration of young in life (ku 87653 from 11 km. n vista hermosa, oaxaca, méxico): "dorsum pale tan with dark brown markings. thighs orange-yellow; labial stripe white; iris bronze" (duellman, field notes, july 10, 1964.) _remarks._--smith (1953:150) named _cyanosticta_ as a subspecies of _hyla phaeota_. the differences in cranial characters and certain external characters between _phaeota_ and _cyanosticta_ indicate that they are distinct species. furthermore, a gap of about 350 kilometers exists between the known geographic ranges of the two kinds. _distribution._--_smilisca cyanosticta_ inhabits wet forests on the atlantic slope of southern méxico and northern central america from northern oaxaca and southern veracruz through northern chiapas in méxico and into el petén and northern alta verapaz in guatemala (fig. 2). apparently the range is discontinuous, for in southern méxico the species is found in cloud forest at elevations of 830 to 900 meters on the northern slopes of the sierra de juárez. in the sierra de los tuxtlas in southern veracruz the species is found in wet forest at elevations of 300 to 1200 meters, but is absent in the intervening lowlands characterized by drier forest. in the west forests of northern alta verapaz and el petén, guatemala, the species is found at low elevations. _specimens examined._--78, as follows: mexico: =chiapas=: monte libano, mcz 28271-9; 8 km. n san fernando, 24 km. ne tuxtla gutierrez, uimnh 41588. =oaxaca=: 11 km. n vista hermosa, ku 84918-20 (skeletons), 87198-212, 87647 (eggs), 87648-52 (tadpoles), 87653 (young), uimnh 57199-201; 8 km. s yetla, ku 87213, ummz 124838 (8). =veracruz=: coyame, ummz 111459-60; between coyame and tebanco, ummz 121198; dos amates, ummz 121297; between laguna de catemaco and volcán san martín, ummz 121200; volcán san martín, uimnh 35403-4, 35408-12, ummz 118163; se slope volcán san martín, ummz 121199, 121295 (2), 121296, 121298. guatemala: =alta verapaz=: chinajá, ku 55935-7, 55938 (skeleton). =el= =petén=: 10 km. nnw chinajá (alta verapaz), ku 55934; piedras negras, cnhm 99006-7, 99227, uimnh 28853, usnm 111139-41, 111143-7; 8 km. s piedras negras, cnhm 99008; semicoch, usnm 35907. [illustration: fig. 2. map showing locality records for _smilisca cyanosticta_ (triangles) and _smilisca phaeota_ (circles).] =smilisca phaeota= (cope) _hyla phaeota_ cope, proc. acad. nat. sci. philadelphia, 14 (9):358, 1862 [holotype.--usnm 4347 from turbo, colombia; j. cassin collector]. boulenger, catalogue batrachia salientia in british museum, p. 402, feb. 1, 1882. werner, sitzungsb. akad. wiss. münchen, 27:215, 1897. günther, biologia centrali-americana: reptilia and batrachia, p. 269, sept. 1901. nieden, das tierreich, amphibia, anura i, p. 261, june 1923. dunn, occas. papers boston soc. nat. hist., 5:413, oct. 10, 1931. gaige, hartweg, and stuart, occas. papers mus. zool. univ. michigan, 357:4, oct. 26, 1937. cooper, copeia, 2:122, june 30, 1944. breder, bull. amer. mus. nat. hist., 86(6):416, aug. 26, 1946. smith and taylor, bull. u. s. natl. mus., 194:88, june 17, 1948; univ. kansas sci. bull, 33:364, march 20, 1950. taylor, univ. kansas sci. bull., 35(1):837, july 1, 1952. brattstrom and howell, herpetologica, 10:117, aug. 1, 1954. goin, herpetologica, 14:120, july 23, 1958. cochran, bull. u. s. natl. mus., 220:57, 1961. _hyla labialis_ peters, monats. konigl. akad. wissen. berlin, p. 463, 1863 [holotype.--zmb 4913 from "region of bogotá," colombia]; monats. konigl. akad. wissen. berlin, p. 618, oct. 16, 1873. boulenger, catalogue batrachia and salientia in british museum, p. 397, feb. 1, 1882. _hyla baudini dolomedes_ barbour, occas. papers mus. zool. univ. michigan, 129:11, jan. 25, 1923 [holotype.--mcz 8539 from río esnápe, sambú valley, darién, panamá; barbour and brooks collectors]. barbour and loveridge, bull. mus. comp. zool. harvard, 69:278, june, 1929. _hyla phaeota phaeota_, smith, herpetologica, 8:152, jan. 30, 1953. minton and smith, herpetologica, 16:103, june 17, 1960. _smilisca phaeota_, starrett, copeia, 4:303, dec. 30, 1960. _diagnosis._--size large ([m] 65 mm., [f] 78 mm.); skull as long as wide, lacking frontoparietal fontanelle; large supraorbital flanges having straight edges and extending posterolaterally; large squamosal not in contact with maxillary; tarsal fold moderately wide, full length of tarsus; inner metatarsal tubercle moderately large, low, flat, elliptical; hind limbs relatively long, tibia averaging more than 53 per cent of snout-vent length; labial stripe silvery white; lips lacking vertical bars; loreal region pale green; dark brown or black tympanic mark dispersing into brown venated pattern on flanks; posterior surfaces of thighs pale brown, with or without darker flecks or small cream-colored flecks. (foregoing combination of characters distinguishing _s. phaeota_ from any other species in genus.) table 2.--geographic variation in size and proportions in males of smilisca phaeota. (means in parentheses below observed ranges; data based on ten specimens from each locality.) ================================================================== |snout-vent |head width/|interorbital locality |length |snout-vent |distance/ | |length |head width -----------------------------+-----------+-----------+----------- bonanza, nicaragua | 40.8-47.7 | 34.1-38.0 | 31.0-36.1 | (43.7) | (36.3) | (35.4) -----------------------------+-----------+-----------+----------- heredia prov., costa rica | 46.3-59.0 | 32.5-36.0 | 30.5-39.6 | (51.7) | (35.0) | (34.7) -----------------------------+-----------+-----------+----------- puntarenas prov., costa rica | 53.6-64.9 | 32.6-36.1 | 31.0-38.0 | (61.4) | (34.5) | (34.4) -----------------------------+-----------+-----------+----------- canal zone, panamá | 52.4-65.5 | 33.5-37.6 | 31.3-37.2 | (56.5) | (35.6) | (34.7) -----------------------------+-----------+-----------+----------- río quesada, colombia | 52.6-61.0 | 33.1-37.1 | 30.1-33.9 | (56.0) | (35.0) | (32.1) -----------------------------------------------------------------_description and variation._--for the purposes of analyzing geographic variation in size and proportions, measurements were taken on ten adult males from each of five samples throughout the range of the species. aside from the data summarized in table 2, the average ratio of tibia length to snout-vent length is noticeably less in colombian specimens (53.4 per cent, as compared with 54.8 to 57.8 per cent in the other samples) and the ratio of head length to snout-vent length is noticeably less in costa rican specimens (33.5 per cent as compared with 34.9 to 35.1 per cent in the other samples). also, specimens from heredia province, costa rica, have a relatively smaller tympanum (62.7 to 80.4 [mean 68.4] per cent of the diameter of the eye, as compared with means of 74.0 to 77.9 per cent in the other samples). two populations are distinctive as regards the size of adult males. specimens from the northern caribbean lowlands of nicaragua (bonanza, the northernmost locality for the species) are remarkably small. males having snout-vent lengths of between 40 and 43 mm. were breeding; the largest male found had a snout-vent length of 47.7 mm. the other extreme in size is attained in specimens from the pacific lowlands of eastern costa rica and western panamá, where most breeding males have snout-vent lengths of more than 55 mm.; the largest male had a snout-vent length of 64.9 mm. the rather striking differences in size among certain samples and the minor differences in proportions among other samples show no geographic trends. instead, the variations apparently are random among the samples. the data presented here possibly are the results of inadequate sampling, but more likely reflect actual differences in the populations. the dorsal ground color of _smilisca phaeota_ is pale green to tan; the venter is creamy white. the dorsum is variously marked with dark olive-green or dark brown spots or blotches (pl. 6c). a dark interorbital bar is usually present. usually a large dark dorsal mark extends from the occiput to the sacral region, but in many individuals this blotch is replaced by two or three dark marks. the dorsal markings are irregular in shape and do not tend to form transverse bands or longitudinal bars. the hind limbs are marked by dark transverse bands, usually four or five on the thigh, five or six on the shank, and four on the tarsus. two or three narrow bands are usually present on the proximal part of the fourth toe. the webbing on the feet is brown. the loreal region is pale green, bordered above by a narrow dark brown canthal stripe extending from the nostril to the orbit. the upper lip is silvery white. a broad dark brown or black mark extends posteriorly from the orbit, encompassing the tympanum, to a point above the insertion of the forelimb. the flanks are pale green or pale tan and marked with a fine dark brown or black venation. the anterior surfaces of the thighs usually are pale brown or grayish tan, sometimes having small, indistinct darker flecks. the posterior surfaces of the thighs are similarly colored, but in most specimens small but distinct dark flecks are present; in some specimens small cream-colored spots are also present on the posterior surfaces of the thighs. a distinct, narrow creamy white anal stripe usually is present. a distinct white stripe is present on the outer edge of the tarsus and fifth toe; on the tarsus the white stripe is bordered below by dark brown. a white stripe also is present on the outer edge of the forearm and fourth finger. in breeding males the throat is dark gray. little geographic variation in color or pattern is evident. few, if any, specimens from the pacific lowlands of south america are green in life. (we have seen no living individuals from south america.) some living individuals from costa rica and all those seen alive from nicaragua have a tint of pale blue on the flanks. in some specimens the dorsal pattern is so faint as to be barely discernible, whereas in most specimens the pattern is bold. the coloration in the living frogs is highly variable due to extreme metachrosis. individuals of this species are capable of changing the dorsal coloration from green to brown in a short period of time. both green and brown individuals have been found active at night. usually those individuals found hiding by day are brown. one individual from finca la sumbadora, panamá (now ku 91914), was kept alive in the laboratory for nearly one month. this individual usually was pale green with tan dorsal markings at night and tan with pale green markings by day. on occasion the pale green dorsal markings were boldly outlined by bright dark green. in living individuals from throughout the range of the species the iris is a bronze color, darkest medially with fine black reticulations. _natural history._--_smilisca phaeota_ inhabits humid lowland tropical forest and seldom ascends the foothills to more than 1,000 meters. the rather equable climatic conditions, especially more or less evenly distributed rainfall throughout the year, permit this frog to be active most of the year. dunn (1931:413) reported males calling on barro colorado island, panamá, in february and in july, and breder (1946:416) noted calling individuals in the chucanaque drainage of darién, panamá in january, march, july, august and october and in costa rica in april through august inclusively. calling males were found at bonanza, nicaragua in march and in july. at all times of year the usual daytime retreats for these frogs are near water; the frogs have been found in elephant ear plants (_xanthosoma_) and in bromeliads; occasional individuals have been found sitting on shaded branches of bushes and trees. none has been observed on the ground or beneath ground-cover by day. the length of the breeding season cannot be determined definitely. the earliest date on which eggs have been found is may 23; gaige, hartweg, and stuart (1937:5) reported a gravid female taken at el recreo, nicaragua, in september, and we have a gravid female taken at almirante, panamá, in march. males usually call from secluded spots at the edge of water. all calling males that we observed were on the ground within a few centimeters of the water. the males usually are hidden beneath an overhanging leaf or some other cover; they definitely do not sit in the open like _smilisca baudini_. most calling males and clasping pairs have been found at the edges of small pools or shallow ditches, although occasional individuals are found at the edges of large ponds or streams. the breeding call consists of one or two moderately short, low-pitched notes (duration 0.33 to 0.42 seconds), repeated at intervals of about 20 seconds to several minutes. each note is a low, vibrant "wauk," having 100 to 130 pulses per second and a dominant frequency of 330 to 420 cycles per second (pl. 10c). the eggs are deposited in loose clumps amidst vegetation in the water. hatchling tadpoles have total lengths of 8.7 to 10.6 mm., and body lengths of 4.1 to 4.5 mm. the external gills are long and filamentous, and the yolk sac is large. the head and caudal musculature are dark brownish black, and the caudal fins are gray. the oral discs are large and roughly circular. the growth and development of the tadpoles are summarized in table 11 and figure 16. a typical tadpole in stage 30 of development (ku 68482 from the río chitaría, cartago province, costa rica) may be described as follows: body length 9.7 mm.; tail length 14.6 mm.; total length 24.3 mm.; body as wide as deep; snout rounded dorsally and laterally; eyes widely separated, directed dorsolaterally; nostril about midway between eye and tip of snout; mouth anteroventral; spiracle sinistral, about midway on length of body and slightly below midline; anal tube dextral; caudal musculature slender, curved upward distally; dorsal fin extending onto body; depth of dorsal fin slightly less than that of ventral fin at mid-length of tail; dorsal part of body pale brown; ventral surfaces transparent with scattered pigment; pale cream-colored, crescent-shaped mark on posterior edge of body; caudal musculature pale creamy tan with scattered pale brown spots; caudal fins transparent with scattered small brown blotches on dorsal and ventral fins; iris pale bronze in life (fig. 13); mouth small; median part of upper lip bare; rest of mouth bordered by one row of pointed papillae; lateral fold present; tooth-rows 2/3, first upper row longest; second upper row slightly shorter, broadly interrupted medially; three lower rows complete, equal in length, slightly shorter than second upper row; upper beak moderately deep, forming broad arch with slender lateral processes; lower beak slender, broadly v-shaped; both beaks serrate (fig. 15e). in tadpoles having fully developed mouthparts the tooth-row formula of 2/3 is invariable. the pale crescent-shaped mark on the posterior part of the body curves anterodorsally on the dorsal surface of the body. these marks in dorsal view give the appearance of a pair of short, pale stripes on the posterior part of the body. most specimens from costa rica have the pale coloration like that described above, but some individuals (notable ku 87683 from guápiles, costa rica, ku 87707 from finca tepeyac, nicaragua, and ku 87708 from bonanza, nicaragua) have much more pigment. in these specimens the same color pattern obtains as in the pallid individuals, but the pigmentation is dense. this is especially noticeable on the tail. recently metamorphosed young have snout-vent lengths of 12.7 to 16.7 mm. (average, 14.3 mm. in eleven specimens). coloration of young in life (ku 68484 from río chitaría, cartago province, costa rica): "dorsum pale tan; side of head and flanks darker brown, separated from tan dorsum by an indistinct cream stripe. limbs pale yellow; thighs flecked with brown; shank and tarsus yellowish tan with indistinct brown bars. soles of feet brown. belly white; throat dusty cream flecked with silvery white. upper lip silvery white. iris bright gold with black flecks. heels, tarsal and anal stripes white" (duellman, field notes, may 23, 1961). _remarks._--peters (1863:463) named _hyla labialis_ from the "region of bogotá, colombia", but in 1873 regarded his new species as identical with _hyla phaeota_ cope, 1862, from turbo, colombia. the name _hyla labialis_ has been used for frogs from the northern andes in colombia (see dunn, 1944:72, and stebbins and hendrickson, 1959:522, for discussion of nomenclature). rivero (1961:131) used the name _hyla vilsoniana_ cope, 1899, for the frogs from the northern andes previously referred to _hyla labialis_. a review of the nomenclature and taxonomy of these frogs, which superficially resemble _smilisca_ but are unrelated, is beyond the scope of the present study. _hyla baudini dolomedes_ barbour, 1923, is based on a small _smilisca phaeota_ (mcz 8539) having a snout-vent length of 45.5 mm. dunn (1931a:413) placed _dolomedes_ in the synonymy of _smilisca phaeota_. we have examined the holotype of _dolomedes_ and agree with dunn's assignment. smith (1953:150) described _hyla phaeota cyanosticta_ from guatemala. our studies on the external morphology, coloration, and especially the cranial osteology provide evidence that _cyanosticta_ is a species distinct from _phaeota_. _distribution._--_smilisca phaeota_ inhabits humid tropical forests from northeastern nicaragua southward on the caribbean lowlands to elevations of about 1000 meters and on the pacific lowlands of costa rica, exclusive of the arid regions of guanacaste, throughout the lowlands of panamá, exclusive of the savannas of the pacific lowland and the azuero peninsula, and southward on the pacific slopes of south america through colombia to west-central ecuador; also the valleys of the río cauca and río magdalena in colombia (fig. 2). _specimens examined._--528, as follows: nicaragua: =matagalpa=: finca tepeyac, 10 km. n, 9 km. e matagalpa, ku 85439, 87707 (tadpoles); matagalpa, mcz 3546-7, ummz 92367; 19 km. n matagalpa, ummz 116495-6. zelaya: bonanza, ku 84854-62, 84950-2 (skeletons), 85440-50, 87708-9 (tadpoles); cukra, amnh 80618; río mico, 16 km. e recreo, ummz 79711 (6), 79712 (4); junction río mico and río siguia, ummz 79713 (10); río siguia, 11 km. nw rama, ummz 79714 (14), 79715 (11), 79716 (21), 79717, 79718 (3). costa rica: =alajuela=: cinchona, ku 32255, 64286-8; 5 km. s ciudad quesada, usc 8077; laguna monte alegre, ku 64289-90; las playuelas, 11 km. s los chiles, usc 7216; san carlos, usnm 29961. =cartago=: moravia de turrialba, ku 32212-47, 37133-5, 41093 (skeleton), 64280-1, usc 7243 (3); peralta, ku 32271-2; río chitaría, 3 km. nne pavones, ku 64273-9, 68477 (eggs), 68478-83 (tadpoles), 68484 (young); río reventazón, mcz 29196-203, ummz 117677 (9); turrialba, ku 25720-2, 32209-11, 32266-8, 32273-4, 37136-67, 41090-2 (skeletons), 64270-2, mcz 29221, 29222 (tadpoles), 29269-70, usnm 29934. =guanacaste=: tilarán, ku 36805-7; 8 km. ne tilarán, ku 36803-4. =heredia=: barranca del río sarapiquí below isla bonita, ku 64282-3; cariblanco, ku 32256-60, 41094 (skeleton), 64284, mcz 7967; isla bonita, ku 32250-4; 4.2 km. w puerto viejo, ku 64285, 68485; 7.5 km. w puerto viejo, ku 68486; 1 km. s puerto viejo, ku 86518. =limón=: bambú, usc 7182 (4); batán, ummz 118582; coén, mcz 9825; la lola, ku 32262-4, uf 4029, ummz 117678 (3); los diamantes, cnhm 101295-8, ku 25723-4, 32265, 64267-9; pandora, ummz 122650 (2), usc 7188 (3), 7190; puerto limón, ku 32261; río larí at río dipari, 21 km. sw amubre, usc 7177; río toro amarillo, 7 km. w guápiles, ku 86519, 87683 (tadpoles); suretka, ku 36808-10, 37168. =puntarenas=: agua buena, ku 36790; 1.6 km. e buenos aires, ummz 117578; 3 km. nw buenos aires, ku 64304; 4 km. n, 15 km. w dominical, ku 68491-2 (tadpoles); esparta, mcz 8029-30, 8032; golfito, ku 32270; 6 km. e golfito, ku 84999-500 (skeletons); gromaco, ummz 123677 (4); palmar, ku 32269; 4 km. ese palmar sur, ku 64305-6; 5.6 km. se palmar sur, ku 68489 (tadpoles); 7.0 km. se palmar sur, ku 68490 (young); 8.5 km. se piedras blancas, ku 64292-303; quebrada boruca, 22 km. e palmar norte, ku 64291; rincón, "camp seattle," peninsula de osa, ummz 123676 (3), usc 7254; río ferruviosa, 7 km. s rincón, usc 7235; 1.6 km. wnw villa neily, ku 68493 (young), 68494 (tadpoles). =san josé=: san isidro el general, ku 32249, ummz 75025; 10 km. n san isidro el general, mcz 29099-103; 13 km. wsw san isidro el general, ku 86517; 15 km. wsw san isidro el general, ku 68487 (tadpoles), 68488 (young), 68495 (young); 20 km. wsw san isidro el general, ku 32248. panama: no province: cano saddle, usnm 69588; punta de pena, usnm 38733; quipo, amnh 18925-6. =bocas del toro=: almirante, ku 80080, 91835-6; 1.6 km. w almirante, ku 91837; 3 km. w almirante, ku 91824 (skeleton), 91838-43, 91906-7; 11 km. nw almirante, cnhm 67853-61; 13 km. w almirante, ku 91825-7 (skeletons), 91844-9; fish creek, ku 92329; isla popa, ku 91850-1. =canal zone=: barro colorado island, cnhm 6007, 13316, 13325, 13331, 13360-2, 13377-8, mcz 24191-5, uf 7523, ummz 63547-60, 64457, 69497 (3); 3.7 km. w cocoli, ku 67916; fort sherman, mcz 10139; gatun, mcz 35644; junction roads c25b and c16, tnhc 23839; madden forest preserve, tnhc 23837-8. =coclé=: el valle, ku 77521-4, 77649 (tadpole), tnhc 23369. =comarca del barú=: progreso, ummz 61085-9. colón: achiote, ku 77516-20, 77648 (young); río candelaria, cnhm 67851-2. =darién=: río esnápe, sambú valley, mcz 8539; río sucubti, chalichiman's creek, amnh 40512; camp creek, amnh 40758-9; camp creek, camp townsend, amnh 40988. =panamá=: nw slope cerro prominente, ku 80459; finca la sumbadora, ku 91914 (skeleton). =chiriquí=: 2 km. w concepción, amnh 68910. columbia: =antioquia=: puerto berrio, cnhm 30805 (goin); turbo, usnm 39899. =caldas=: pueblorrica, santa cecilia, cnhm 54768-71 (goin). =choco=: no specific locality, amnh 3984-6; andagoya, bmnh 1915.10.21. 69-70, cnhm 81857 (goin); golfo de urabá, cnhm 63881 (goin); peña lisa, condoto, bmnh 1913.11.12. 118-125, 1913.11.12. 137-146 (goin); pizarro, cnhm 4451-3, 4455-61 (goin); río san juan, playa del oro, cnhm 54772 (goin); río quesada, amnh 13615-77; 37 km. up río puné, amnh 13688; 48 km. up río puné, amnh 13689. =narino=: tumaco, río rosario, cjg 2310-13 (goin). =valle=: buenaventura, bmnh 1895.11.16.82 (goin); raposa, wat 166, 346-47, 388 (goin); río calima above córdoba, cjg 2249-57 (goin). ecuador: no province: bulun, amnh 10620. =esmeraldas=: cachabé, amnh 10625-8; río capayas, cnhm 35712; río sapaya, ummz 58910 (5); salidero, amnh 10623-4; san javier, amnh 10618. =guayas=: hacienda balao chico, ummz 123904. =imbabura=: pambelar, amnh 10629, 10631. =pichincha=: hacienda espinosa, 9 km. w santo domingo de los colorados, ku 40220. =smilisca puma= (cope), new combination _hyla puma_ cope, proc. amer. philos. soc., 22:183, 1885 [holotype.--usnm 13735 from nicaragua; j. f. moser collector]. günther, biologia centrali-americana: reptilia and batrachia, p. 270, sept., 1901. nieden, das tierreich, amphibia, anura i, p. 251, june, 1923. cochran, bull. u. s. natl. mus., 220:58, 1961. _hyla wellmanorum_ taylor, univ. kansas sci. bull. 25(1):843, july 1, 1952 [holotype.--ku 30302 from batán, limón, costa rica, edward h. taylor collector]; univ. kansas sci. bull., 36(1):626, june 1, 1954. duellman and berg, univ. kansas publ. mus. nat. hist., 15:194, oct. 26, 1962. _smilisca wellmanorum_, starrett, copeia, 4:303, dec. 30, 1960. _diagnosis._--size small ([m] 38.0 mm., [f] 46.0 mm.), differing from other species in the genus by the following combination of characters: skull about as long as broad; frontoparietal fontanelle keyhole-shaped; supraorbital flanges absent; squamosal small, not in contact with maxillary; bony portion of ethmoid terminating at anterior edge of orbit; tarsal fold weak, two-thirds length of tarsus; inner metatarsal tubercle small, low, flat, elliptical; snout rounded in dorsal profile; lips thin and flaring; fingers having only vestige of web; toes one-half webbed; diameter of tympanum about two-thirds that of eye; narrow labial stripe white; pair of dark brown (sometimes interconnected) stripes on tan dorsum; no blue spots on flanks or thighs; vocal sac in breeding males pale brown. (foregoing combination of characters distinguishing _s. puma_ from other species in genus.) _description and variation._--ten breeding males from the vicinity of puerto viejo, heredia province, costa rica, have snout-vent lengths of 32.5 to 37.9 mm. (34.8 mm.). in these specimens, the length of the tibia to the snout-vent length is 0.48 to 0.53 (0.51), and the tympanum/eye ratio is 0.52 to 0.72 (0.65). seven females have snout-vent lengths of 40.8 to 45.8 mm. (43.9 mm.). no individual has more than a vestige of a web between the second and third and fourth fingers. none has a web between the first and second fingers. breeding males lack nuptial excrescences on the thumbs. the vocal sac is moderately large and bilobed. in preserved specimens the dorsal ground color varies from yellowish tan to grayish brown. all specimens have dark brown dorsal markings in the form of a pair of dorsal stripes, variously modified (pl. 7a). in some specimens, such as ku 91716, the stripes are discrete and extend from the postorbital region nearly to the vent. in most specimens the stripes are connected by a transverse mark in the scapular region and in many others also by a transverse mark in the sacral region. in some specimens the stripes are fragmented posteriorly; fragmentation is extreme in ku 30300, in which the dorsal pattern consists of two series of dark longitudinal dashes. the other extreme is a nearly complete fusion of the stripes, as in ku 91714. a dark brown interorbital bar usually extends onto the eyelids, but in some specimens this is reduced to a short v-shaped mark or small spot between the eyes. there is no dark post-tympanic mark, but dark brown pigment forms a venated pattern from the axilla to the mid-flank; the inguinal region is white, finely mottled with dark brown. the dorsal surfaces of the hind limbs are colored like the body and have two or three dark brown transverse marks on the thighs, three to five marks on the shanks, and one or two marks or irregularly arranged dark flecks on the tarsi. the anterior and posterior surfaces of the thighs are pale tan to brown. the webbing of the feet is tan to grayish brown. a narrow white labial stripe, white anal stripe, and narrow white stripes on the tarsi and outer edges of the forelimbs are invariably present. the ventral surfaces are creamy white. in life the dorsum is tan or pale brown with dark brown markings. some individuals have scattered metallic green flecks on the dorsum. the flanks are mottled dark brown and creamy white. the posterior surfaces of the thighs are dark brown. the vocal sacs are grayish brown, and the iris is a deep bronze color. _natural history._--_smilisca puma_ inhabits humid lowland tropical forests having more or less evenly distributed rainfall throughout the year. the equable climatic conditions seemingly permit these frogs to be active throughout most of the year. taylor (1952:846) found calling males at batán, costa rica, on july 20, 1951. we found the species breeding near puerto viejo, costa rica, on february 19, june 18, july 13, and july 31. specimens of calling males from costa rica in the collection at the university of southern california were obtained in february at la fortuna, on august 22 at los diamantes, on august 30 at jabillos, and on september 5 at la lola. gravid females were collected in june, july and august. males call from shallow water. all breeding congregations of this species that we have found were in a grassy marsh, 7.5 kilometers west of puerto viejo, costa rica. tadpoles were found in water-filled depressions in the marsh at night. when first observed, tadpoles were near the surface of the water; they responded to light by quickly taking refuge in the dense grass. no tadpoles were observed by day. the breeding call consists of a low squawk, usually followed by a series of one or more rattling secondary notes (duration of primary notes, 0.06-0.35 seconds; of secondary notes, 0.10 to 0.47 seconds), repeated at intervals of 5 to 55 seconds. the primary notes have 187 to 240 pulses per second and major frequencies of about 740 to 1870 cycles per second (pl. 11a). only six tadpoles are available for study. four of them in stage 34 of development have body lengths of 9.0 to 9.5 mm., tail lengths of 14.0 to 15.0 mm., and total lengths of 23.0 to 24.5 mm. one tadpole in stage 38 and one in stage 40 have total lengths of 31.0 mm. a typical tadpole in stage 34 of development (ku 91807 from 7.5 km. w puerto viejo, heredia province, costa rica) has a body length of 9.5 mm., tail length of 15.0 mm., and total length of 24.5 mm.; body about three-fourths as deep as wide; snout rounded dorsally and laterally; eyes widely separated, directed dorsolaterally; nostril about midway between eye and tip of snout; mouth anteroventral; spiracle sinistral, about two-thirds distance from snout to posterior end of body and slightly below midline; anal tube dextral; caudal musculature slender, barely curved upward distally; dorsal fin extending onto body; at mid-length of tail, depth of caudal musculature equal to that of dorsal fin and ventral fin; body grayish brown, palest ventrally; caudal musculature pale creamy yellow with bold gray reticulations; caudal fins transparent with gray reticulations anteriorly and black flecks posteriorly on both fins (fig. 14a). median part of upper lip bare; rest of mouth bordered by two rows of short blunt papillae; lateral fold present; tooth-rows 2/3; upper rows equal in length; second upper row broadly interrupted medially; three lower rows complete, first and second rows equal in length, slightly shorter than upper rows; third lower row noticeably shorter; upper beak shallow, forming broad, continuous arch with slender lateral processes; lower beak slender, broadly v-shaped, both beaks finely serrate (fig. 15b). all six tadpoles are colored alike, except that in the larger specimens scattered white flecks are present on the ventral surface of the body, and the dark reticulations continue farther posteriorly on the caudal fins than in the smaller tadpoles. in two specimens the third lower tooth-row is only about one-half the length of the other lower rows, and in one specimen the second lower tooth-row is shorter than the first. coloration of tadpoles in life: "body olive-brown with silvery green flecks laterally. caudal musculature olive-brown with greenish tan flecks. fins brown with greenish gold flecks. iris deep bronze." (duellman, field notes, february 19, 1965). one recently metamorphosed young (ku 91808) has a snout-vent length of 12.4 mm. in life this frog had a pale tan dorsum with dark brown markings, yellowish tan posterior surfaces of thighs, grayish brown throat, and bronze iris. _remarks._--the identity of cope's _hyla puma_ has not been known. the name has appeared in various compilations, but no workers have referred any of their specimens to that species. examination of the holotype (usnm 13735), an adult female, revealed the presence of the following combination of characters: snout-vent length 45.8 mm., snout blunt above and rounded laterally, nostrils close to tip of snout, lips thin and flaring, a vestige of a web on the hands, feet about one-half webbed, tarsal fold weak and extending about two-thirds length of tarsus, dorsal markings consisting of a faded dark interorbital bar and a pair of faded longitudinal brown marks connected by a transverse band in the scapular region. the type agrees well with specimens of _smilisca wellmanorum_ (taylor, 1952); the vestigial webbing on the hands and the dorsal coloration are especially significant. consequently, we consider _hyla wellmanorum_ taylor, 1952, to be a synonym of _hyla puma_ cope, 1885. cope gave only "nicaragua" as the locality for _hyla puma_. the specimen was part of a collection received at the united states national museum from lt. j. f. moser. among the species in the collection are _dentrobates pumilio_, _phyllomedusa helenae_, _corythophanes cristatus_, _pliocercus dimidatus_, _tretanorhinus nigroluteus_, and others characteristically found on the caribbean lowlands of central america. thus, it seems reasonable to assume that the type specimen of _hyla puma_ came from the caribbean lowlands. though no other nicaraguan specimens have been found by us, numerous specimens are known from the caribbean lowlands of costa rica. cochran (1961:58), in her catalogue of type specimens in the united states national museum, listed _hyla puma_ cope, 1885, as a synonym of _hyla molitor_ schmidt, 1857. she made no qualifying statements. schmidt (1858:246), in his descriptions of the species in the year following his publication of the names and latin diagnoses, stated: "dorsum uniformly gray, more intensive on back, fading away laterally and on extremities; in every-day-life this blue would be called _mueller's blau_. a delicately dotted black line runs on the canthus rostralis from the opening of the nose to the corner of the eye. in the armpits, on the flanks and the thighs two of our three specimens have black marblings." [free translation] certainly on the basis of coloration _hyla puma_ is distinctly different from _hyla molitor_. _distribution._--this species lives in the wet, forested regions of the caribbean lowlands of costa rica and presumably southern nicaragua (fig. 3). all specimens are from low elevations; the highest known elevation for the occurrence of this frog is 285 meters at laguna bonilla. [illustration: fig. 3. map showing locality records for _smilisca puma_ (triangles) and _smilisca sila_ (circles).] _specimens examined._--62, as follows: nicaragua: no specific locality, usnm 13735. costa rica: =alajuela=: jabillos, 5 km. n santa clara, usc 8058 (6); 5 km. w la fortuna, usc 8078 (2); río la fortuna at la fortuna, usc 7151 (3). =cartago=: laguna bonilla, tunnel camp near peralta, ku 32171. =heredia=: puerto viejo, ku 86521; 5.9 km. w puerto viejo, ku 64307; 7.5 km. w puerto viejo, ku 64308-10, 64311 (skeleton), 64312-15, 68635-6 (skeletons), 85001-2 (skeletons), 86520, 87770-1 (skeletons), 91709-16, 91791-2, 91807 (tadpoles), 91808 (young). =limon=: batán, ku 30300-2; la lola, ku 32169, usc 141, 201, 8067; los diamantes, ku 32170, ummz 118470 (6), usc 212; 2.4 km. e los diamantes, usc 8049 (5). =smilisca sila= new species _hyla gabbi_, noble, proc. biol. soc. washington, 37:66, feb. 21, 1924. dunn, occas. papers boston soc. nat. hist., 5:413, oct. 10, 1931. schmidt, smithsonian misc. coll., 89(1):6, march 16, 1933. _hyla sordida_, dunn, copeia, 3:166, nov. 19, 1937. cooper, copeia, 2:121, june 30, 1944. breder, bull. amer. mus. nat. hist., 86(8):417, aug. 26, 1946. _hyla phaeota_, breder, bull. amer. mus. nat. hist., 86(8): pl. 55, aug. 26, 1946. _holotype._--adult male, ku 91852 from a small stream at the north edge of the village of el volcán, chiriquí province, panamá, elevation 1280 meters; obtained on feb. 5, 1965, by william e. duellman. _paratypes._--ku 91853-74, collected with the holotype. _diagnosis._--size moderate ([m] 45.0 mm., [f] 62.2 mm.); skull wider than long, having large, ovoid frontoparietal fontanelle; supraorbital flanges absent; squamosal small, not contacting maxillary; bony section of ethmoid extending anteriorly between nasals; tarsal fold weak, full length of tarsus; inner metatarsal tubercle low, flat, elliptical; lips thick, rounded, not flaring; fingers one-third webbed; toes three-fourths webbed; diameter of tympanum about one-half that of eye; margin of upper lip faintly marked by interrupted creamy white stripe; dark spots on dorsum; pale flecks on flanks and posterior surfaces of thighs; vocal sacs in breeding males dark brown. (foregoing combination of characters distinguishing _s. sila_ from any other species in genus.) _description of holotype._--snout-vent length 36.6 mm.; tibia length 19.8 mm., 54.1 per cent of snout-vent length; foot length 15.5 mm., 42.3 per cent of snout-vent length; head length 12.7 mm., 34.7 per cent of snout-vent length; head width 13.3 mm., 36.8 per cent of snout-vent length; snout short, in lateral profile truncate, only slightly rounded above, in dorsal profile rounded; canthus rounded; loreal region noticeably concave; lips thick, rounded, not flaring; nostrils not protuberant, directed laterally; internarial distance 3.0 mm.; internarial area flat; top of head flat; interorbital distance 3.5 mm., 26.3 per cent of head width; diameter of eye 4.2 mm., thrice distance (1.4 mm.) from tympanum to eye, and half again distance (2.8 mm.) from orbit to nostril; pupil horizontally ovoid; width of eyelid 2.8 mm., 21.1 per cent of head width; dermal fold from posterior corner of orbit covering upper edge of tympanum to point above insertion of forelimb; diameter of tympanum 2.3 mm., 54.7 per cent of diameter of eye; no axillary membrane; arms moderately robust; weak fold on wrist; faintly scalloped fold along ventrolateral margin of forearm; fingers short, slender; fingers from shortest to longest, 1-2-4-3; vestige of web between first and second fingers; others about two-fifths webbed; discs moderate, diameter of that on third finger about one-third diameter of eye; triangular outer palmar tubercle; elliptical inner palmar tubercle on base of pollex; subarticular tubercles large, conical, none bifid; supernumerary tubercles few, large, conical; brown nuptial excrescence on prepollex; heels overlap by about one-fifth length of shank when hind limbs adpressed; tibiotarsal articulation extending to nostril; tarsal fold weak, extending nearly full length of tarsus; inner metatarsal tubercle elliptical, flat; outer metatarsal tubercle absent; toes moderately long; toes from shortest to longest, 1-2-3-5-4, third and fifth about equal in length; discs about same size as those on fingers; webbing extending to middle of penultimate phalanx on all toes, except only to distal end of antepenultimate phalanx of fourth toe; subarticular tubercles round; supernumerary tubercles large, round, present only on proximal digits. anal opening directed posteriorly at level of upper edge of thighs; no noticeable anal sheath; flat tubercles ventrolateral to anal opening large; skin of chest, belly, and posterior surfaces of thighs granular; other surfaces smooth; tongue broadly cordiform, shallowly notched posteriorly, and barely free behind; vomerine teeth 4-4, situated on ventral surfaces of separated rounded prominences between posterior margins of small, ovoid inner nares; vocal slits long, each situated along inner margin of ramus; color (in preservative) pinkish tan above with irregular olive-brown markings forming interconnected spots on back; four bars on dorsal surface of each thigh; five bars on shank, and three on tarsus; inguinal region white with black mottling; posterior surfaces of thighs yellowish tan proximally, dark brown distally; margins of lips grayish white with brown markings; ventral surfaces of hands and feet grayish brown; belly and posterior part of throat creamy white; anterior part of throat brown. _description and variation._--ten breeding males from finca la sumbadora, panamá, have snout-vent lengths of 40.0 to 44.8 mm. (42.3 mm.). in these specimens the tibia/snout-vent length ratio is 0.50 to 0.57 (0.54), and the tympanum/eye ratio is 0.48 to 0.58 (0.53). there is a geographic gradient in size; specimens from the western part of the range (southern costa rica) are smaller than those from the eastern part of the range (eastern panamá). five males from the pacific lowlands of southern costa rica have snout-vent lengths of 31.6 to 38.2 mm. (34.7 mm.); ten males from el volcán, chiriquí, panamá, 32.6 to 37.9 mm. (36.4 mm.), and eight males from barro colorado island, canal zone, 38.2 to 42.0 mm. (35.6 mm.). these are smaller than the males from finca la sumbadora, which is east of the canal zone. ten females from el volcán have snout-vent lengths of 44.2 to 55.6 mm. (49.2 mm.), as compared 56.1 to 62.2 mm. (58.2 mm.) in three females from finca la sumbadora. large females have scattered small tubercles on the head and back; tubercles occur in males from costa rica and in some males from western panamá. the truncate snout is characteristic of both sexes. the coloration of _smilisca sila_ consists of a gray, tan, or pale reddish brown dorsal ground color and a creamy white venter. the dorsum is marked by dark brown, olive-brown, or dark reddish brown spots or blotches (pl. 7b). usually the blotches are discrete, but in some individuals they are interconnected and form an irregular dark mark on the dorsum. there is no tendency for the blotches to form transverse bars as in _smilisca sordida_. in one specimen (ku 80467) the blotches are fused and form two wide irregular longitudinal stripes, as in _smilisca puma_. in some females the dorsal markings are reduced to a few small spots or are nearly absent (ku 92332), whereas in other females the dorsal markings are bold. in one female (ku 91894) the dorsal markings are narrowly bordered by pale blue, and numerous pale blue flecks are present on the pale brown dorsum. in many individuals of both sexes small white flecks are present on the dorsal surfaces. usually the flanks and posterior surfaces of the thighs have black mottling enclosing pale blue spots and flecks, respectively. the dorsal surfaces of the limbs are marked by dark brown transverse bars; usually three or four bars are present on each forearm, thigh, and shank. the coloration of the flanks and limbs varies geographically. specimens from southern costa rica and western panamá have distinct bars on the limbs; the posterior surfaces of the thighs have brown reticulations enclosing small blue flecks in specimens from costa rica and bolder, black reticulations enclosing large pale blue spots in specimens from western panamá. in specimens from costa rica the flanks are brown with pale blue flecks, whereas in those from chiriquí, panamá, the flanks are pale blue with dark brown mottling in the inguinal region. frogs from el valle and cerro la campana usually have distinct bars on the limbs; the posterior surfaces of the thighs are colored as in frogs from chiriquí, and the inguinal region is pale blue with coarse brown mottling. specimens from barro colorado island are marked like those from el valle and cerro la campana, except that on the posterior surfaces of the thighs fine black reticulations enclose many dark blue spots. in specimens from darién and from panamá province east of the canal zone (altos de pacora, cerro jefe, finca la sumbadora, and río pacora), the markings on the dorsal surfaces of the limbs are indistinct or absent in males, but distinct in some females. intense brown and black pigment forms fine reticulations delimiting bold blue spots on the flanks; this coloration extends to the axilla in many specimens. fine black reticulations enclose many dark blue spots on the posterior surfaces of the thighs. in females, the throat is creamy white; in some specimens scattered brown flecks are present on the chin and throat. in breeding males the anterior part of the throat is dark gray or dark brown. the coloration in life is as variable as it is in preservative. in life the holotype had a tan dorsum with dark olive-green irregular markings and small green flecks. the limbs were tan with dark brown transverse bars. the flanks were grayish tan anteriorly; the inguinal region and posterior surfaces of thighs were blue with black mottling. the belly was creamy white, and the throat was brown with creamy yellow flecks. the iris was a dull bronze color. among the paratypes, some individuals had green flecks, others did not. the inguinal region and posterior surfaces of the thighs were pale blue, pale yellowish green, or grayish tan with black mottling. the blue was most noticeable in females. colors of a male from finca la sumbadora, panamá, were described as follows: "dorsum olive-brown; irregular dark brown blotches, pale green flecks, and raised creamy yellow spots on dorsal surfaces; belly creamy white; throat grayish brown; undersides of limbs grayish tan; groin, anterior and posterior surface of thigh, inner surface of shank, anterior edge of tarsus, and proximal parts of third and fourth toes pale blue marbled with dark brown and black; webbing brown; iris pale bronze, finely reticulated with black." (duellman, field notes, january 28, 1964.) a female (now ku 91894) from altos de pacora, panamá, was described as follows: "an irregular dark brown, green-bordered figure on head and back; dark brown, green-bordered bands on limbs--all on a lighter brown and heavily green-spotted background. these markings are more vivid at night than during the day. lower sides, from midbody onto front of thighs and rear of thighs onto venter of shanks to heels and thence dorsally onto basal portions of toes heavily blue spotted on a light brown (front of thighs and venter of shanks) to blackish brown background. venter cream. iris gray-brown, finely veined with dark brown." (charles w. myers, field notes, december 14, 1964.) note that in the earlier discussion of coloration of preserved specimens, the green spots and borders have changed to pale blue after six months in alcohol. in living individuals from costa rica and panamá west of the canal zone, the blue coloration on the flanks and thighs is much less conspicuous than in specimens from eastern panamá. the color of the iris is variable, even in frogs from one locality. the coloration of the iris in 13 living frogs (now ku 92333-45) from valle hornito, chiriquí, panamá, was described as follows: "iris variable--from pale to dark brown; in a few the iris has a golden cast to the brown; in a few others the lower half of the iris is pale gray with the upper half being light brown." (charles w. myers, field notes, april 24, 1965). _natural history._--_smilisca sila_ inhabits the pacific slopes of lower central america where a pronounced dry season occurs. we have records of males calling in december through may and also in august (latter date from el volcán, chiriquí, panamá). the breeding season seems to be correlated with the time of the year when the water is clear and at a low level in the streams where these frogs breed. males call from the edges of small, shallow streams, from rocks in the streams, or less frequently from vegetation overhanging the streams. females are most frequently found on the banks of streams, and clasping pairs usually are in shallow pools in streams. one individual was found in a bromeliad about three meters above the ground in the daytime. the breeding call consists of a low squawk, usually followed by a series of one or more rattling secondary notes (duration of primary notes, 0.06 to 0.28 seconds; of secondary notes, 0.14 to 0.48 seconds), repeated at intervals of 4 to 20 seconds. the primary notes have 97 to 120 pulses per second and major frequencies of about 900 to 2220 cycles per second (pl. 11b). eggs were obtained artificially in the field; the average length of ten embryos in the neural groove stage is 2.4 mm., and the average diameter of the outer envelope is 4.9 mm. hatchlings have large, conical oral discs, heavy gills, and a large amount of yolk; their average total length is 6.3 mm. tadpoles have been found in pools in clear streams; some tadpoles have been observed to cling by their mouths to rocks in the stream; others were found on the bottom where they seek refuge among pebbles or under rocks and leaves. a complete developmental series of tadpoles is not available. eleven tadpoles in stage 25 of development have body lengths of 8.3 to 10.2 mm. (9.3 mm.), tail lengths of 17.3 to 21.0 mm. (18.8 mm.), and total lengths of 25.9 to 31.0 mm. (28.1 mm.). one tadpole in stage 41 and one in stage 42 have body lengths of 11.5 and 12.5 mm., tail lengths of 27.2 and 29.5 mm., and total lengths of 38.7 and 42.0 mm., respectively. the snout-vent lengths of two specimens in stage 43 and one in stage 45 are 12.7, 13.0, and 13.6 mm., respectively. a typical tadpole in stage 25 of development (ku 80620 from finca la sumbadora, panamá) has a body length of 9.5 mm., tail length of 19.0 mm., and a total length of 28.5 mm.; body only slightly wider than deep, nearly flat dorsally; snout broadly rounded in dorsal view, bluntly rounded in lateral view; eyes widely separated, directed dorsolaterally; nostril slightly closer to eye than to tip of snout; mouth ventral; spiracle sinistral, located about two-thirds distance from snout to posterior edge of body; anal tube dextral; caudal musculature moderately heavy, straight; dorsal fin not extending onto body; fins deepest at about two-fifths length of tail, where depth of caudal musculature about equal to depth of dorsal and depth of ventral fin; musculature extending nearly to tip of tail; body dark grayish brown above and pale grayish tan below with small dark brown spots dorsally and white flecks laterally; caudal musculature pale tan with dark brown flecks over entire surface and dark brown streaks on posterior one-half of ventral fin and on all of dorsal fin (fig. 14b). median one-third of upper lip bare; rest of mouth bordered by a single row of conical papillae; lateral fold present; tooth rows 2/3; upper rows cone-shaped, about equal in length, broadly /\-shaped; second upper row narrowly interrupted medially; lower rows complete, about equal in length, but slightly shorter than upper rows; upper beak moderately massive, its inner surface forming a continuous arch with short lateral processes; lower beak broadly \/-shaped; both beaks finely serrate (fig. 15d). tadpoles from el volcán, chiriquí (ku 91833), are more heavily pigmented than those from finca la sombadora; the spots on the tail are larger. in life these tadpoles had dark brownish black bodies with golden and green lichenous flecks; the tail was tan with dark brown markings, and the iris was a grayish bronze color. in life tadpoles from finca la sumbadora were olive-tan above and dark gray with pale bluish gray irridescent spots ventrally. the caudal musculature was creamy tan with brown flecks and streaks, and the iris was pale bronze. metamorphosing young have been found on vegetation at the edge of streams and have been raised in the laboratory. seven recently metamorphosed young have snout-vent lengths of 13.6 to 15.6 mm. (14.6 mm.). a living juvenile (ku 91913) raised in the laboratory from a tadpole obtained at finca la sumbadora had a brown dorsum with darker brown markings, a white spot below the eye, and a narrow white labial stripe. the belly was white; the flanks were brown with white spots, and the posterior surfaces of the thighs were yellowish tan. the iris was a golden bronze color with much black reticulation. _remarks._--this species has been confused with _smilisca sordida_; most authors have referred both species to _hyla (smilisca) gabbi_. examination of the types of _hyla sordida_, _gabbi_, _salvini_, and _nigripes_ revealed that all of the names were referable to a single species (_s. sordida_), and that the small, blunt-snouted species in panamá and southern costa rica probably was without a name. possibly _hyla molitor_ schmidt (1857) is based on the species that we have named _s. sila_, but several discrepancies in his description, plus the unknown provenance of the type, have led us to discount the applicability of that name to the species under consideration. _distribution._--_smilisca sila_ ranges along the pacific slopes and lowlands of southern costa rica and panamá at elevations from sea level to about 1300 meters; in northern south america the species occurs in the caribbean lowlands and in the valleys of the northward draining rivers of colombia (fig. 3). _specimens examined_, 234, as follows: costa rica: =puntarenas=: 6 km. e golfito, ku 91717; quebrada boruca, 22 km. e palmar norte, ku 64265-6; río zapote, 7 km. e palmar norte, usc 7100 (2). =san josé=: san isidro el general, ku 28200; 14 km. nw san isidro el general, usc 7098 (2); 15 km. wsw san isidro el general, usc 7097. panama: =canal zone=: barro colorado island, amnh 62320-3, cnhm 13324, 13326-8, 13330, 13338, 13359, 13423-5, ku 80460-6, 80619 (young), 80625 (skeleton), ummz 63542-6, usc 7051. =chiriquí=: boquete, amnh 69815, ummz 58441-5; el volcán, ku 77413, 91828-31 (skeletons), 91852-74, 91832 (eggs), 91833 (tadpoles); 6 km. s el volcán, cnhm 60442; 16 km. nnw el volcán, ku 91879-90; finca palosanto, 6 km. wnw el volcán, ku 77406-12, 77692 (skeleton), 91875-7, 92330-1; río colorado, 17 km. nnw el volcán, ku 91878, 92332; valle hornito, 19 km. ne gualaca, ku 92333-45. =coclé=: el valle, amnh 55440-5 (13), 59607-14, cnhm 48140, 60349-2, 60387-92, 60401-4, 60443, 67842-5, ku 91834 (young), 91902-4, tnhc 23751-2, usnm 140653. =colón=: río candelaria, amnh 53708-15, cnhm 67826-36. =darién=: camp creek, camp townsend, amnh 40756-7, 40936-9, 40992; río chico, amnh 39784, 40986-7; río pita, cnhm 67823-5; tacarcuna, usnm 141796-802; three falls creek, amnh 41684, 51788. =los santos=: cerro hoya, usnm 148213-4; lajamina, río puria, ku 67915. =panamá=: altos de pacora, ku 91894; cerro jefe, ku 91895-6; cerro la campana, cnhm 67846, ku 91897-900, usnm 139689; finca la sumbadora, ku 80467-81, 80620 (tadpoles), 91910 (eggs), 91911-2 (tadpoles), 91913 (young), 91908-9 (skeletons); río calobra, usnm 53722, río pacora, 9 km. nne pacora, ku 91901. =veraguas=: cerro carbunco, usnm 129066; cerro tute, cnhm 67837-41; isla cebaco, río platanal, ku 91891-3. colombia: =antioquia=: urabá, villa arteaga, cnhm 63893 (goin). =atlantico=: sabanalarga, río causa, amnh 14506. =smilisca sordida= (peters), new combination _hyla sordida_ peters, monatsb. konigl. akad. wissen. berlin., p. 460, 1863 [syntypes.--zmb 3141 (two specimens) from "veragua," panamá; j. von warszewicz collector]. brocchi, mission scientifique au mexique ..., pt. 3, sec. 2, études sur les batrachiens, p. 42, 1881. boulenger, catalogue batrachia salientia in british museum, p. 393, feb. 1, 1882. günther, biologia centrali-americana: reptilia and batrachia, p. 273, sept. 1901. nieden, das tierreich, amphibia, anura, i, p. 258, june, 1923. _hyla gabbi_ cope, jour. acad. nat. sci. philadelphia, new ser., 8, pt. 2:103, 1876 [syntypes.--usnm 30658-9 from near sipurio, limón, costa rica; william m. gabb collector]. brocchi, mission scientifique au mexique ..., pt. 3, sec. 2, études sur les batrachiens, p. 37, 1881. boulenger, catalogue batrachia salientia in british museum, p. 372, feb. 1, 1882. cope, bull. u. s. natl. mus., 32:32, 1887. günther, biologia centrali-americana: reptilia and batrachia, p. 274, sept. 1901. werner, abhand. konigl. akad. wissen. münchen., 22:351, 1903. nieden, das tierreich, amphibia, anura i, p. 252, june, 1923. taylor, univ. kansas sci. bull., 35(1):840, july 1, 1952. cochran, bull. u. s. natl. mus., 220:54, 1961. _hyla nigripes_ cope, jour. acad. nat. sci. philadelphia, new ser., 8, pt. 2:104, 1876 [syntypes.--usnm 30685-6, from pico blanco, costa rica; william m. gabb collector]. brocchi, mission scientifique au mexique ..., pt. 3, sec. 2, études sur les batrachiens, p. 38, 1881. boulenger, catalogue batrachia salientia in british museum, p. 394, feb. 1, 1882. cope, bull. u. s. natl. mus., 32:32, 1887. günther, biologia centrali-americana: reptilia and batrachia, p. 278, sept., 1901. nieden, das tierreich, amphibia, anura i, p. 253, june, 1923. james, copeia, 3:147, sept. 30, 1944. taylor, univ. kansas sci. bull, 35(1):853, july 1, 1952. cochran, bull. u. s. natl. mus., 220:56, 1961. _hyla salvini_ boulenger, catalogue batrachia salientia in british museum, p. 372, feb. 1, 1882 [syntypes.--bmnh 1947.2.24.13-14 from cartago, costa rica; osbert salvin collector]. günther, biologia centrali-americana: reptilia and batrachia, pl. 71, fig. b., sept., 1901. werner, abhand. zool.-bot. gesell. wien, 46:8, sept. 30, 1896. _smilisca gabbi_, starrett, copeia, 4:303, dec. 30, 1960. _diagnosis._--size moderate ([m] 45 mm., [f] 64 mm.); skull slightly wider than long, having large and elongate frontoparietal fontanelle; supraorbital flanges absent; squamosal small, not contacting maxillary; bony section of ethmoid terminating just anterior to anterior edge of orbit; tarsal fold weak, full length of tarsus; inner metatarsal tubercle long, low, flat, elliptical; lips thin and flaring; fingers one-half webbed; toes four-fifths webbed; diameter of tympanum about one-half that of eye; no white labial stripe; dorsal dark markings irregular, sometimes forming broad transverse bars; pale flecks on flanks and usually on posterior surfaces of thighs; vocal sacs in breeding males white. (foregoing combination of characters distinguishing _s. sordida_ from any other species in genus.) _description and variation._--ten breeding males from 15 to 20 kilometers west-southwest of san isidro el general, san josé, costa rica, have snout-vent lengths of 38.1 to 42.6 mm. (40.5 mm.). in these specimens, the tibia/snout-vent length ratio is 0.50 to 0.54 (0.52), and the tympanum/eye ratio is 0.45 to 0.57 (0.49). specimens from the pacific slopes of costa rica are larger than those from the meseta central and the caribbean lowlands. ten males from 6 kilometers east of golfito, puntarenas, have snout-vent lengths of 38.4 to 44.6 mm. (41.8 mm.), and five males from rincón, peninsula de osa, have snout-vent lengths of 38.8 to 41.6 mm. (40.3 mm.). snout-vent lengths of ten males from la fortuna, alajuela, are 31.9 to 36.0 mm. (34.4 mm.), of ten males from pandora, limón, 33.8 to 37.6 mm. (35.9 mm.), and of ten males from escazú and río jorco on the meseta central, 34.3 to 37.6 mm. (36.0 mm.). eight females from the río jorco on the meseta central have snout-vent lengths of 48.8 to 53.8 mm. (50.4 mm.), and six females from various localities on the pacific slopes of costa rica have snout-vent lengths of 56.5 to 64.0 mm. (59.8 mm.). the only noticeable differences in proportions between males and females is in the tympanum/eye ratio; for example, this ratio is 0.47 to 0.53 (0.49) and 0.54 to 0.68 (0.61) in ten males and eight females, respectively, from the meseta central. the shape of the snout and the associated cranial elements of _s. sordida_ vary geographically and ontogenetically. specimens from the caribbean lowlands have blunt snouts in lateral view; those from the pacific lowlands have longer, more slender snouts that are pointed in lateral view, and those from the meseta central are intermediate in snout shape between the two lowland populations (fig. 4). these differences in shape of the snout are dependent on the nature of the underlying cranial bones, principally the maxillaries and nasals. in specimens from the caribbean lowlands the nasals are long, wide, and narrowly separated from the ethmoid; the anterior edge is just posterior to the nostril. the maxillary flanges are nearly vertical. in specimens from the pacific lowlands the nasals are relatively shorter, narrower, and rather widely separated from the ethmoid; the anterior edges of the nasals do not extend so far forward as in specimens from the caribbean lowlands. the maxillary flanges slant medially. in these cranial characters, specimens from the meseta central are intermediate between the two lowland populations. superimposed on this geographic variation are ontogenetic changes, which are most noticeable in males. in smaller, and presumably younger, specimens the snouts are more pointed than in larger specimens; consequently some small males from the caribbean lowlands resemble larger males from the pacific lowlands, since the nasals and maxillaries of the former are not fully ossified. in addition, in small breeding males the ethmoid is only about one-half ossified, a large frontoparietal foramen is present, the anterior arm of the squamosal extends only about one-fourth the distance to the maxillary (two-thirds the distance in larger specimens), and the tegmen tympani are short, as compared with the long, thin elements in larger specimens. [illustration: fig. 4. variation in the shape of the snout in _smilisca sordida_; left column females, right column males; all from costa rica: (a) camp seattle, rincón de osa, puntarenas prov. (ummz 123684); (b) quebrada agua buena, 3 km. sw rincón de osa, puntarenas prov. (usc 7236); (c) río oro, 28.5 km. nw villa neily, puntarenas prov. (ku 91742); (d) río jorco, near desamparados, san josé prov. (ku 91765); (e-f) bambú, limón prov. (usc 7183). ×3.] the dorsal ground-color of _smilisca sordida_ is gray to pale tan or reddish brown; the venter is white. the dorsum is variously marked with dark gray, dark brown, reddish brown, or olive-green spots or blotches (pl. 7c). a dark interorbital bar usually is present. the dorsal markings on the body usually consist of a blotch, or two or more spots, on the occiput, in the scapular region, and in the sacral region. in many specimens, especially females, these markings are in the form of broad transverse bars. a female (usc 7164) from las cañas, guanacaste, costa rica, has a tan dorsum with many black flecks and round brown spots bordered by darker brown. one female (ku 91763) from the río jorco, san josé, costa rica, has a unicolor tan dorsum. some individuals have scattered, small white spots on the dorsum; these are most evident in a male (usc 7153) from la fortuna, alajuela. white labial stripes and anal stripes are absent in all specimens. the limbs are marked by dark brown transverse bars; these are indistinct in some specimens from the meseta central and caribbean lowlands, whereas they are distinct in all specimens from the pacific lowlands. specimens from the caribbean lowlands have two to six bars on each shank, whereas specimens from the pacific slopes have four to six bars on each shank, and specimens from the meseta central have as many as eight bars on each shank. a narrow, sometimes broken white line is present on the ventrolateral edge of the forearm. the webbing on the hand is tan or pale gray, and the ventral surfaces of the tarsi and the webbing on the feet are dark gray or brown. breeding males have dark brown nuptial excrescences on the prepollex. the flanks and posterior surfaces of the thighs usually are marked by bluish white and creamy tan flecks, respectively, but vary considerably. in specimens from the caribbean lowlands a small amount of flecking is present in the inguinal region, and on the posterior surfaces of the thighs flecks are few or absent. in specimens from the meseta central, numerous large flecks or small, round spots (pale bluish white in life) are on the posterior half of the flanks; small flecks are on the posterior surfaces of the thighs. specimens from the pacific slopes and lowlands of southern costa rica (puntarenas and san josé provinces) have bold mottling of black and bluish white on the flanks and many bluish white flecks on the posterior surfaces of the thighs. the flanks are reticulated from the axilla to the groin in two females (ummz 123684 and usc 7236) from rincón, peninsula de osa. in specimens from the pacific slopes of guanacaste in northwestern costa rica, flecks are present in the inguinal region; indistinct flecks are on the posterior surfaces of the thighs. the throat is immaculate in specimens from the caribbean lowlands in limón province; the throats are dusky laterally in most other specimens except some from the meseta central, in which the throats are heavily flecked with black. this variation occurs in males and females. the color and pattern in life are highly variable. a composite description of living individuals (now ku 91718-41) from 6 kilometers east of golfito, puntarenas, costa rica, illustrates the variability: "dorsum pale olive-green, fading to tan posteriorly, or tan all over with dark olive-green or dark brown spots on back and bars on limbs. flanks dark brown with cream, greenish gray, or bluish gray mottling. posterior surfaces of thighs dark brown with pale blue, pale green, or tan flecks. iris creamy silver. throats white with some brown flecks peripherally." (duellman, field notes, february 15, 1965.) a male from the río jorco, san josé, costa rica, was dull olive-tan above with olive-green marks; the flanks were brown with pale tan flecks, and the posterior surfaces of the thighs were pale brown with cream-colored flecks. six females from the same locality were reddish brown above with olive-brown or dark brown markings; one was uniform orange-tan, and another was dull olive-green with darker markings. the color of the iris in living frogs varies from creamy silver to grayish yellow or bronze with a variable amount of black reticulation. _natural history._--_smilisca sordida_ is not associated with any one type of vegetation; instead it lives in the vicinity of rocky streams having low gradients. breeding takes place primarily in the dry season, when the water in the streams is clear and at a low level. through most of the range of _s. sordida_ showers, or even short heavy rains, occur in the dry season. after such rains the breeding activity is maximal. breeding congregations have been found from december through april, but a few calling males and gravid females have been taken in june, july, and august. in the rainy season non-breeding individuals are found sitting on bushes near streams at night. taylor (1952:843) found specimens in bromeliads by day. males usually call from rocks or gravel bars in, or at the edge of, streams. some individuals perch in low bushes overhanging the streams, and some sit in shallows in the streams. clasping pairs have been found on the banks of streams and in shallow water in streams. the breeding call consists of one to six moderately short, rather high-pitched notes (duration 0.18 to 0.45 seconds) repeated at intervals of 12 seconds to several minutes. each note is a vibrant rattle having 78 to 135 pulses per second and major frequences of about 1200 to 2600 cycles per second (pl. 11c). the tadpoles live in shallow parts of the streams, where they cling to the surfaces of small rocks and hide beneath leaves and rocks. a complete developmental series of tadpoles is not available; measurements of those stages examined are summarized in table 12. a typical tadpole in stage 36 of development (ku 68475 from 15 km. wsw of san isidro el general, costa rica) has a body length of 11.7 mm., tail length of 22.8 mm., and a total length of 34.5 mm.; body about three-fourths as deep as wide; snout broadly rounded in dorsal view, sloping and rounded in lateral view; eyes widely separated, directed dorsolaterally; nostril slightly closer to eye than to tip of snout; mouth ventral; spiracle sinistral, about two-thirds distance from snout to posterior end of body and slightly below midline; anal tube dextral; caudal musculature heavy, straight; dorsal fin not extending onto body; fins deepest at about mid-length of tail; there depth of caudal musculature equal to depth of dorsal fin and half again as deep as ventral fin; musculature extending nearly to tip of tail; body reddish brown above and pale grayish brown with white flecks below; caudal musculature pale tan with brown flecks; a series of reddish brown dashes at base of caudal fin separated from others in series and from dashes on other side by creamy white; fins transparent with reddish brown flecks on posterior one-half of ventral fin and on all of dorsal fin (fig. 14c). mouth bordered by two rows of short, pointed papillae; lateral fold present; tooth-rows 2/3; upper rows equal in length; second upper row narrowly interrupted medially; three lower rows complete, nearly as long as upper rows, deeply indented medially; upper beak robust, inner surface not forming continuous arch with short lateral processes; lower beak deep, v-shaped; both beaks bearing short serrations (fig. 15f). little variation occurs in structure. in some specimens the second upper tooth-row is complete; no individuals were found to have the row broadly interrupted medially. the series of dark dashes on the dorsal edge of the caudal musculature is diagnostic of all stages studied. in life, tadpoles from 15 and 20 kilometers west-southwest of san isidro el general, costa rica, had a tan body, often with an olive-tan tinge; the caudal musculature was tan; the flecks and dashes were dull red or reddish brown. tadpoles from 6 kilometers east of golfito, costa rica, had bodies with olive-green flecks. the caudal musculature was brown with bluish green flecks; the fins were transparent with reddish brown flecks. the belly was a silvery golden color. tadpoles from bajos de jorco, costa rica, had brown bodies with bluish green flecks; the tail and fins had reddish brown flecks and dashes. the iris was a bronze color in specimens from all three localities, as well as in the young mentioned in the following paragraph. nine recently metamorphosed young were found on vegetation at the edges of streams in april. these specimens have snout-vent lengths of 13.1 to 15.7 mm. (14.9 mm.) and in life were pale greenish tan or olive-tan above and white below. the hands, feet, and thighs were pale yellowish tan. _remarks._--the foregoing synonymies indicate that confusion has existed in the application of various names, to this species, as well as in use of the names _sordida_ and _gabbi_ to include the species that we describe and name _smilisca sila_. correct allocation of the names involved was possible only after studying and comparing the type specimens, for the descriptions given by the various authors are not sufficiently explicit to determine the nature of many essential features. the presence of a rounded snout and a long white throat in males distinguishes _s. sordida_ from _s. sila_, which has a high truncate snout and short dark throat in males. the two syntypes of _hyla sordida_ peters, 1863, (zmb 3141) are males having snout-vent lengths of 36.9 and 37.0 mm. the two syntypes of _hyla gabbi_ cope, 1876 (usnm 30658-9), are females having snout-vent lengths of 52.8 and 53.7 mm., respectively. also included in the collections made by gabb is eastern costa rica are two males (usnm 30685-6), which cope (1876) named and described as _hyla nigripes_. these specimens are soft and faded, but are recognizable as the same as _hyla sordida_ peters; the syntypes of _hyla nigripes_ have snout-vent lengths of 37.6 and 37.7 mm. we have examined one of the syntypes of _hyla salvini_ boulenger, 1882 (bmnh 1947.2.24.13), a female having a snout-vent length of 54.6 mm. we are convinced that all of these type specimens are representatives of one species, the earliest name for which is _hyla sordida_ peters, 1863. the type localities for three of the named species are in costa rica--_h. gabbi_ from sipurio on the caribbean lowlands, _h. nigripes_ from the caribbean slopes of pico blanco, and _h. salvini_ from cartago on the meseta central. the type locality of _h. sordida_ was given as "veraguas" by peters (1863). at that time veraguas was often considered to be most of western panamá. though we have not seen panamanian specimens other than the types of _s. sordida_ and one specimen from the pacific lowlands of western panamá, the species probably occurs on the caribbean slopes of western panamá. the species has been taken on the caribbean lowlands of costa rica within a few kilometers of panamá; collecting on the caribbean slopes in the provinces of bocas del toro and veraguas should reveal the presence of _smilisca sordida_ there. _distribution._--_smilisca sordida_ is found along the pacific slopes and lowlands from guanacaste, costa rica, southeastward to extreme western panamá, to elevations of about 1200 meters on the meseta central in costa rica, and on the caribbean slopes and lowlands of costa rica and probably adjacent panamá (fig. 5). one specimen purportedly comes from "río grande, nicaragua." [illustration: fig. 5. map showing locality records for _smilisca sordida_.] _specimens examined._--412, as follows: nicaragua: "río grande" (? depto. zelaya), mcz 2634. costa rica: =alajuela=: between atena and salto de san mateo, usc 6185; 8 km. n ciudad quesada, usc 7155 (4); la fortuna, usc 7153 (20); 3 km. e la fortuna, usc 7150; san carlos, usnm 29969; sarchi, ku 32990-9, 36792-3. =cartago=: cartago, bmnh 1947.2.24.13; headwaters of río pacuare, usc 119; instituto interamericano de ciéncias agricolas, turrialba, ku 37012, usc 420, 437; río reventazón, turrialba, mcz 29268: 10 km. n río reventazón bridge, usc 7073; 5 km. sw río reventazón bridge on paraiso-orosi road, usc 669; turrialba, ummz 118405, usc 455, usnm 29936-9. =heredia=: puerto viejo, ku 36791. =guanacaste=: las cañas, usc 7164; santa cecilia, mcz 7924-5; tilarán, usc 7161 (5). =limón=: bambú, usc 7171 (2), 7183 (13); la lola, usc 820 (6), 6083-94, 8064, 8071; pandora, usc 7188 (7), 7189, 7190 (3), 7191 (5); pico blanco, usnm 30685-6; río larí, 14-16 km. sw amubre, usc 7179, 7180 (10); sipurio, usnm 30658-9; suretka, ku 36764, 36765 (skeleton), 36766-78. =puntarenas=: 6 km. n dominical, ku 91749-50, 91811 (young), 91812 (tadpoles); esparta, mcz 8028; 6 km. e golfito, ku 91718-41, 91809 (young), 91810 (tadpoles), 91816-9 (skeletons), usc 7103 (23); quebrada agua buena, 3 km. sw rincón de osa, usc 7236 (6); quebrada boruca, 22 km. e palmar norte, ku 64264; rincón de osa, camp seattle, ummz 123680-5, s-2792 (skeleton), usc 705 (5), 6023, 7254; río barranca, usc 7119 (2); río ceiba, 6 km. nw buenos aires, ku 91747-8, usc 7112 (7); río ciruelitas, 16 km. nw esparta, usc 7121 (3); río claro, 14.2 km. nw villa neily, usc 7110 (4); río ferruviosa, 7 km. s rincón de osa, usc 7235 (4); río lagarto at pan-american hwy. (guanacaste border), usc 7122 (4); río la vieja, 30 km. e palmar norte, ku 87684 (tadpoles), 91743-6, usc 7083 (2); río oro, 28.5 km. nw villa neily, ku 91742; río volcán, 10 km. w buenos aires, usc 7113; río zapote, 7 km. e palmar, usc 7100 (4); 3-5 km. w palmar, usc 7101 (18); 7 km. se palmar, ku 64261-3; 1.2 km. nw villa neily, usc 8032; 3 km. nw villa neily, usc 7109 (20); 5 km. nw villa neily, usc 6176, 8035. =san josé=: bajos de jorco, ku 91813 (tadpoles); escazú, ku 34863, 34869-75, usc 813; between monrovia and la hondura, ± 0.5 km. n santa rosa, usc 302 (2); paso ancho, río jorco, ummz 122649 (6), usc 530 (3); río jorco, near desamparados, ku 91757-65, 91796-7, 91820-3 (skeletons), usc 228, 513, 7117 (7); río peje, 10 km. sse san isidro el general, usc 7115 (3); río tiriví, mcz 7972; san isidro el general, cnhm 101096, ku 28201, 32989, ummz 72024; 15 km. wsw san isidro el general, ku 64245-56, 68473 (tadpoles), 68474 (young), 68475 (tadpoles), 86516, 91754-6, 91793-5, usc 7097 (6); 17.1 km. wsw san isidro el general, usc 6047; 18 km. wsw san isidro el general, usc 689; 20 km. wsw san isidro el general, ku 64257-9, 64260 (skeleton), 68468 (young), 68469 (tadpoles), 68470 (young), 68471-2 (tadpoles), 68476 (young), 68633-4 (skeletons), 91751-3; san josé, amnh 7501-4, usc 298; santa rosa, río virilla, usc 7145. panama: =chiriquí=: río jacu, 5.8 km. ese paso canoas, ku 91905. "veraguas," zmb 3141 (2). analysis of morphological characters osteology in attempting to assay the taxonomic significance of skeletal differences we are faced with a dearth of data on the skeletons of frogs in general and hylids in particular. recent reviews by brattstrom (1957) and hecht (1962, 1963) have been concerned with general salientian classification and phylogeny, principally at the family level. savage and carvalho (1953), griffiths (1959), and baldauf (1959) used osteological characters in determining the taxonomic status of the families pseudidae, brachycephalidae, and bufonidae, respectively. carvalho (1954) presented osteological evidence for the generic separation of new world microhylids. zweifel (1956) and tihen (1962) used osteological characters at the levels of the species-group and species in their respective studies on _scaphiopus_ and _bufo_. little has been recorded about the skeletons of the hylids. goin (1961) mentioned dentigerous elements and cranial co-ossification in his synopsis of the genera of hylids. copland (1957) in his review of the _hyla_ of australia, funkhouser (1957) in her revision of _phyllomedusa_, and zweifel (1958) in his review of _nyctimystes_ did not consider skeletal characters. some osteological studies on hylids have yielded worthwhile information. mittleman and list (1953) used osteological characters in defining the genus _limnaoedus_: starrett (1960) used cranial characters in combination with jaw musculature in defining the genus _smilisca_, and duellman (1964) used cranial characters in delimiting the _hyla bistincta_ group. brief descriptions of cranial structure were given for _phrynohyas_ (duellman, 1956) and _ptychohyla_ (duellman, 1963a); specific and sexual differences in the skulls of _hyla chaneque_ and _hyla taeniopus_ were pointed out by duellman (1965). stokely and list (1954) described early cranial development in the hylid _pseudacris triseriata triseriata_. because our knowledge of the skeleton in hylids is so incomplete, we are not attempting to place _smilisca_ in the general scheme of hylid phylogeny on the basis of skeletal characters. instead, our purposes are to describe the skeleton and its ontogenetic development in one member of the genus (_s. baudini_), and to make comparisons that show taxonomic differences in osteological characters among species of _smilisca_. the study of 68 dried skeletons and 25 cleared and stained preparations, including an ontogenetic series of _s. baudini_, has resulted in an understanding of the progressive development of skeletal elements and a knowledge of interspecific and intraspecific variation in these elements. furthermore, investigations of the osteology have provided correlations between some cranial characters and certain aspects of external morphology. _descriptive osteology of smilisca baudini_ the following description is based primarily on an adult female (ku 68184): _skull._--the skull is large, solid, and broader than long; the greatest width is between the sutures of quadratojugal and maxillary on either side of the skull (pls. 2-3). the maxillaries bear well-developed dorsal flanges, curve gently, join the moderately convex premaxillaries anteriorly and form a slightly truncate snout. the combined premaxillary width is about one-fourth the width of the skull. the premaxillaries are separated medially, and laterally from the maxillaries by sutures. each premaxillary bears a dorsomedial alary process, which is anteriorly convex and four times as high as the depth of the lateral wing of premaxillary; each premaxillary also has a ventromedial palatine process that projects dorsally from the lingual edge of the premaxillary. the septomaxillaries are closely associated dorsally with the premaxillaries immediately lateral to the prenasal processes. the nasals are large, widest anteriorly and narrowing posteriorly, parallel to maxillaries, and not separated from the ethmoid by cartilage. the nasals bear long, delicate maxillary processes extending nearly to the maxillaries. anteriorly, the nasals are widely separated by the partially ossified internasal septum, which is in contact with the premaxillaries between the prenasal processes; the anterior points of the nasals lie approximately one-half the distance between the anterior ends of the ethmoid and the premaxillaries. the ethmoid is large and completely ossified; the margins are smooth. the trunate anterior edge lies between the nasals and is in contact with the internasal septum. the frontoparietals are large, smooth-margined, and bear large supraorbital flanges curving posterolaterally at the rear of the orbit. a small, oval foramen involves the posterior part of the ethmoid and anterior portion of frontoparietals; continued ossification in older specimens fills in the foramen, thereby resulting in a solidly roofed cranium. the auditory regions are relatively massive and bear narrow tegmen tympani; the distal ends of the tegmen tympani are medial to the lateral edge of the pterygoids in dorsal view. the squamosals are large; the long anterior arm is separated from the maxillary by a suture. the delicate, spindle-shaped columellae lie ventral to the tegmen tympani and squamosals, are spatulate distally, and have a broad basal attachment to the auditory region. the vomers are moderately large and are in contact anteriorly with the premaxillaries and posteriorly with the ethmoid. each vomer has two wide serrated flanges laterally. the tooth-bearing parts of the vomers are widely separated and at a slight angle to one another; the vomers terminate medially in two pointed processes on the ethmoid. the palatines are edentate, but bear strong ridges throughout their lengths. they are broadly in contact with the maxillary, are narrow medially, and are attached by pointed processes to the medial part of the ethmoid. the pterygoids are large, attached to the maxillaries immediately anterior and medial to the squamosal-maxillary connection, bear well-developed pedicles, which are broadly attached to the proötic, and a wide wing is in contact posteriorly with the distal two-thirds of the quadrate. the angular makes up most of the lower jaw, bears a broad articular surface posteriorly, and has a small coronoid process on the lingual edge; anteriorly the angular is separated from the dentary and mentomecklian by meckel's cartilage. the dentary lies external to the angular and extends from the mentomecklian to approximately the mid-length of the angular. the mentomecklians are ossified, but separated by cartilage medially. _hyoid._--the hyoid plate is curved, thin, and mostly cartilaginous, but calcined posteriorly (fig. 6). the anterior cornua are slender, cartilaginous, and curve anteromedially from the hyoid plate and thence laterally and posteriorly, to attach to the posterior surface of the proötics. the lateral cornua are broad, flat, cartilaginous lateral extensions from the bases of the anterior cornua. the posterior cornua are bony, except distally. [illustration: fig. 6. ventral view of hyoid apparatus of an adult male _smilisca baudini_ showing areas of muscle attachment: _gen. l._, attachment of geniohyoideus lateralis; _gen. m._, attachment of geniohyoideus medialis; _hyo._, attachment of hyoglossus; _omo._, attachment of omohyoideus; _pet._, petrohyoideus; _st._, attachment of sternohyoideus. ku 64220, ×5.] _vertebral column._--the atlas lacks transverse processes and a neural crest, whereas transverse processes are present on the other seven presacral vertebrae, and knoblike neural crests are present on the second, third, and fourth vertebrae; a faint neural ridge is visible on the fifth vertebra. the transverse processes are directed laterally on the second and sixth vertebrae, ventrolaterally on the third, posterolaterally on the fourth and fifth, and anterolaterally on the seventh and eighth. the processes are slightly expanded on the fourth, and more so on the fifth, vertebra. the sacral diapophyses are expanded and have a border of calcified cartilage laterally. there are two sacral condyles. the slender coccyx has a thin dorsal ridge on the anterior three-fourths of its length. _pectoral girdle._--the omosternum is large, ovoid, and cartilaginous; the sternum is a thin cartilaginous sheet deeply notched posteriorly and is not differentiated into episternal and xiphisternal elements. the coracoids are robust, twice as stout as the clavicles. the epicoracoidal cartilages overlap in the usual arciferal manner, except that they are fused anteriorly between the slender clavicles. the clavicles are strongly arched. the clavicle, coracoid, and scapula on each side form a bony articulation at the glenoid fossa. a bifurcation of the ventral end of the scapula results in a large glenoid foramen. the scapula is flat and expanded dorsally; the suprascapula is broad, flat, and calcified in large adults. in young specimens no distinct ossification of the cleithrum or ossification of endochondral centers are evident. _arm and hand._--the humerus is equally well-developed in both sexes and has a prominent lateral crest. the radius and ulna are completely fused. a bony prepollex is present in both sexes. the metacarpals are about equal in length. the phalangeal formula is 2-2-3-3; the terminal phalanges are claw-shaped. _pelvic girdle._--the ilia are long, slender, and slightly curved. a thin ridge projects laterally from the dorsal edge of the posterior one-half of each ilium. the ilial prominence is large and knoblike when viewed from above. the anterior edge of the ilial prominence is at the level of the anterior edge of the acetabular border. the dorsal acetabular expansion is small. the pubis is slender, and the ischium is elevated and robust. _leg and foot._--the slightly curved femur has a distinct crest proximally on the posterior surface. the nearly straight tibio-fibula is slightly longer than the femur. the tibial and fibial elements are completely fused but have a distinct cleft between them. a small foramen exists at the mid-length of the tibio-fibula. the fibulare (calcaneum) is much more robust than the tibiale (astragalus). the prehallux is large and flat. the metatarsals of the third, fourth, and fifth digits are equal in length; the metatarsal of the second is somewhat shorter, and that of the first is much shorter. the phalangeal formula is 2-2-3-4-3; the terminal phalanges are claw-shaped. _developmental cranial morphology of smilisca baudini_ the following description of development of the skull of _smilisca baudini_ is based on the examination of 12 cleared and stained specimens. in table 3 the cranial bones are listed in the left hand column in the approximate order of their appearance in the young frogs. across the top of the table selected specimens designated by developmental stage or snout-vent length are listed. it should be noted that although each individual, from left to right, has an increasing number of ossified bones, the correlation with increasing size is imperfect; the precise ages of the individuals are unknown. the first bones to appear are the septomaxillaries, frontoparietals, part of the exoccipital, and the parasphenoid in developmental stage 40. the frontoparietals are represented by two slender ossifications dorsomedial to the orbits; the septomaxillaries are present as small ossifications anterior to the nasal capsules (pl. 1a). the parasphenoid is present as a faint median ossification, and the exoccipital shows some ossification. table 3.--the order of occurrence of cranial ossifications in the skull of smilisca baudini. where numbers are divided by a slash mark, the left and right symbols correspond to the left and right sides of the skull, respectively. =====================+=====+=====+=====+=====+=====+=====+==== bone |stage|stage|12.6 |13.9 |32.0 |27.0 |20.1 | 40 | 44 | mm. | mm. | mm. | mm. | mm. ---------------------+-----+-----+-----+-----+-----+-----+--- frontoparietal | x | x | x | x | x | x | x parasphenoid | x | x | x | x | x | x | x septomaxillaries | x | x | x | x | x | x | x exoccipitals | x | x | x | x | x | x | x squamosals | - | x | x | x | x | x | x premaxillaries | - | x | x | x | x | x | x maxillaries | - | x | x | x | x | x | x nasals | - | - | x | x | x | x | x pterygoids | - | - | x | x | x | x | x vomers | - | - | - | x | x | x | x palatines | - | - | - | x | x | x | x quadratojugals | - | - | - | x | x | x | x ethmoid | - | - | - | - | x | x | x columellas | - | - | - | - | x | x | x supraorbital flanges | - | - | - | - | - | x | x proötics | - | - | - | - | - | - | x vomerine teeth | - | - | 1/1 | 4/3 | 5/5 | 3/3 | 5/4 maxillary teeth | - | 0/7 | 3/5 | 6/5 |30/31|30/26|37/36 premaxillary teeth | - | 2/4 | 3/3 | 5/5 | 7/6 | 8/6 | 8/7 ---------------------+-----+-----+-----+-----+-----+-----+---the dentigerous bones are among the most rapidly developed, although not the first to appear. they are present in developmental stage 44 before metamorphosis is completed. the maxillaries bear a few teeth anteriorly and are ossified posteriorly to a point one-third of the distance from the anterior to the posterior edge of the orbit. ossification lengthens the posterior termini of the maxillaries to the posterior edge of the orbit. in front of the anterior margin of the orbit, bone is proliferated dorsal to the main axes of the maxillaries and forms moderate dorsal maxillary flanges. the premaxillaries appear simultaneously with the maxillaries. initially they are widely separated medially from each other, and laterally from the developing maxillaries; each bears two or three teeth, large dorsally blunt alary processes, and small palatine processes. the median and lateral edges of the prenasal processes lengthen heterochronously, causing the median edges to be longest and to lie slightly dorsal to the level of the septomaxillaries. after the maxillaries and premaxillaries develop, the vomers appear as small horizontal ossifications anterior to the parasphenoid. ossification begins in the lateral flanges, then in the prevomerine processes, and lastly in the posterior dentigerous parts of the bones; the prevomerine processes are the last parts of the vomers to ossify completely. initially the frontoparietals are present as thin rods of ossification dorsomedial to the orbits; the frontoparietals extend from the anterior to the posterior end of the orbit by developmental stage 44. the anterior ends of the bones remain thin and pointed; ossification progresses medially from the midpoint of the length of the orbit and posteriorly to the level of the exoccipital; a median center of ossification joins the frontoparietals posteriorly, thereby forming the posterior border of the frontoparietal fontanelle. the supraorbital flanges of the frontoparietals do not appear until all other cranial bones are ossified, or nearly so. the most rapid ossification begins laterally at the posterior edge of the orbit and decreases anteriorly over the posterior half of the orbit. this differential rate of proliferation of bone results in the pattern of development of the supraorbital flanges shown in figure 7. the nasals appear as thin slivers of bone half way between the anterior ends of the frontoparietals and the end of the snout. as ossification proceeds the nasals assume a triangular shape in dorsal view. the anterior ends are pointed; the lateral margins are parallel to the maxillaries. the posteromedial points do not reach the lateral margins of the ethmoid, and the maxillary processes extend about three-fourths the distance from the bodies of the nasals to the maxillaries. following the union of the frontoparietals posteriorly, the nasals widen anteriorly and are narrower at the midpoints of their long axes than anteriorly or posteriorly. with further ossification the maxillary processes extend to the maxillaries and form complete bony anterior margins to the orbits; the mid-parts of the nasals widen (pl. 1b). [illustration: fig. 7. developmental sequence of the frontoparietal fontanelle and associated bony elements in _smilisca baudird_: (a) ku 60026, ×5; (b) ku 85438, ×4; (c) ku 26328, ×3; (d) ku 68184, ×2.3.] the parasphenoid is the first of the palatal bones to appear. at metamorphosis the bone is well developed; the anterior tip is situated just in front of the anterior edge of the orbit, and posteriorly the lateral processes extend laterally beyond the ossified parts of the auditory region. the pterygoids do not appear until metamorphosis, when ossification is evident in only the mid-parts of the posterolateral arms. ossification follows in the mid-parts of the anterolateral arms and occurs last in the pterygoid pedicles. the palatines do not appear until all three arms of the pterygoids are at least partly ossified. ossification proceeds rapidly from the maxillaries medially to the unossified ethmoid, which is the last of the cranial bones to appear. initially it is extremely shallow; dorsally it is widely separated from the nasals, and ventrally the posterior margin meets the anterior point of the parasphenoid. in dorsal view, ossification proceeds anteriorly between the nasals and posteriorly, ventral to the frontoparietals; ventrally, ossification proceeds posteriorly dorsal to the parasphenoid. the ventral arms of the squamosal and the supraoccipital region of the exoccipital are the first occipital bones to appear. ossification follows in the regions of the semicircular canals and occipital condyles. the dorsal end of the ventral arm of the squamosal and the posterior arm of the squamosal ossify as a unit at the same time the quadratojugal appears. shortly thereafter the anterior arm of the squamosal ossifies, the distal part of the columella appears, and the anterior and lateral parts of the auditory region ossify. the angular and dentary of the lower jaw appear concurrently with the dentigerous bones. initially, the angular is short and broad; the articular surface is absent, and the anterior end is slightly overlapped by the dentary. the mentomecklians do not ossify until approximately the same time that the quadratojugal appears in the upper jaw. _comparative osteology_ the genus _smilisca_ is characterized by the following combination of cranial osteological characters: (1) a large amount of bone is involved in the skull and a minimal amount of cartilage and/or secondarily ossified cartilage; co-ossification is absent. (2) the skulls are uniformly broad with angular lateral margins, and truncate anteriorly. (3) an internasal septum and quadratojugals are present. (4) a well-developed squamosal minimally extends one-fourth the distance from the dorsal end of the quadrate to the maxillary, and maximally is separated from the maxillary by a suture. (5) the ethmoid is large; the distance between the anterior end of the ethmoid and the anterior edge of the premaxillary varies between 15 and 20 per cent of the total length of the skull. on the basis of cranial osteology two species-groups can be recognized within the genus _smilisca_. the _sordida_ group, comprising _s. sordida_ and _puma_, is characterized by a broad skull in which the lateral margins of the maxillaries are relatively straight anterior to the orbit. the moderate-sized nasals are rounded anteriorly, and bear relatively short, sometimes blunt, maxillary processes. the long axes of the nasals are not parallel to the maxillaries. the ethmoid is proportionately small in the _sordida_ group. the bony part of the ethmoid terminates near the anterior edge of the orbits and does not extend anteriorly between the nasals; the entire anterior margin of the ethmoid is separated from the nasals by cartilage. the squamosals are generally small. they are narrow in dorsal view, and minimally extend one-fourth the distance from the dorsal end of the quadrate to the maxillary, and maximally, two-thirds the distance. the tegmen tympani are relatively small (fig. 8). [illustration: fig. 8. dorsal views of the skulls of the species of _smilisca_: (a) _s. baudini_ (ku 68184); (b) _s. puma_ (ku 68636); (c) _s. phaeota_ (ku 41090); (d) _s. sila_ (ku 80625); (e) _s. cyanosticta_ (ku 55938), and (f) _s. sordida_ (ku 36765). ×1.5.] in contrast to the tendency for reduction of cranial parts in the _sordida_ group, the _baudini_ group, constituted by _s. cyanosticta_, _phaeota_, and _baudini_, is characterized by more ossification of the cranial elements. the skull is broad; the lateral margins are less angular and are gently curved, rather than straight as in the _sordida_ group. the nasals tend to be larger with the long axes parallel to the maxillary. anteriorly the nasals are pointed, and posteriorly they bear long, delicate palatine processes extending to the maxillary. the ethmoid is fully ossified, extends anteriorly between the nasals, and laterally is separated by a suture from the nasals if the latter are fully ossified. the squamosals are large, and wide in dorsal view. they minimally extend one-fourth the distance from the dorsal end of the quadrate to the maxillary, and maximally are sutured to the maxillary. the tegmen tympani are massive. _smilisca sila_ is intermediate between the two species-groups described. the skull is broad; the lateral margins are gently curved, and have a pronounced angularity just anterior to the palatines which results in a broad, truncate snout. the nasals are moderate in size; because of the anterior angularity of the lateral margins, the long axes of the nasals lie parallel to the maxillary. the nasals are only slightly pointed anteriorly, and posteriorly they bear short, blunt palatine processes and medial processes in contact with the lateral corners of the ethmoid. the ethmoid is fully ossified, but does not extend anteriorly between the nasals. the squamosals are moderate in size and extend one-fourth the distance from the dorsal end of the quadrate to the maxillary. the tegmen tympani are relatively large, but proportionately short. the cranial characters utilized in the analysis of species groups (general shape, nature of the nasals, ethmoid, squamosals, and tegmen tympani), together with other characters, such as the relative height and shape of the prenasal processes, the extent of the internasal septum, and the nature of the vomers, frontoparietals, maxillaries and pterygoids are useful in distinguishing the various species (table 4, fig. 8), as well as in establishing relationships within the species-groups. within the _sordida_ group, _s. sordida_ and _s. puma_ can be distinguished by the following characters: the bony part of the ethmoid terminates posterior to the anterior edge of the orbit and is thus widely separated from the nasals by cartilage in _s. puma_. in _s. sordida_ the bony part of the ethmoid always terminates at a level equal to, or slightly in front of the anterior edge of the orbit; therefore, less cartilage exists between the ethmoid and nasals in _s. sordida_ than in _s. puma_. the width of the premaxillary comprises about 30 per cent of the width of the skull in _s. sordida_ and 20 per cent in _s. puma_. the proportion of the length of the skull anterior to the bony part of the ethmoid in _s. sordida_ is approximately 21 per cent, as compared with about 29 per cent in _s. puma_. the prenasal processes are convex in _s. sordida_ and straight in _s. puma_. the marked ontogenetic variation in _s. sordida_ is considered in more detail in the account of that species, but it is pertinent to the present discussion to note that with respect to some features of the skull some young breeding specimens of _s. sordida_ are intermediate in appearance between large females of _s. sordida_ and adults of _s. puma_. in some breeding males (usually the smaller individuals) of _s. sordida_ the bony part of the ethmoid terminates at the anterior edge of the orbit and is widely separated from the nasals by cartilage. in small individuals _s. sordida_, especially in males, and in adults of _s. puma_ the tegmen tympani are relatively short, whereas in adult females of _s. sordida_ these elements are long and slender. in the smaller specimens of _s. sordida_ and in _s. puma_ the squamosal is small; it extends only about one-fourth of the distance to the maxillary in the smaller _s. sordida_ and about one-half the distance in _s. puma_. the more massive squamosal in large adult females of _s. sordida_ extends at least two-thirds of the distance to the maxillary. table 4.--comparative cranial osteology of smilisca. ===============+==============================+======================= character | _s. baudini_ | _s. cyanosticta_ ---------------+------------------------------+---------------------- | | alary processes| four times as high as | three times as high | lateral wing of premaxillary;| as lateral wing of | anteriorly | premaxillary; | convex. | anteriorly | | convex. | | nasals | long, wide anteriorly, | long, widest | narrowing posteriorly; | posteriorly; | attached to ethmoid. | attached to | | ethmoid. | | | | | | ethmoid | long; entirely ossified; | long, entirely | smooth margins. | ossified; | | smooth margins. | | frontoparietal | small, ovid fontanelle | large fontanelle, two | present or absent; | and one-half times as | long, pointed postorbital | long as wide; narrow | processes curving | supraorbital flanges | along posterior | with irregular margins. | border of orbit. | | | squamosal | large: anterior arm | large; anterior arm | in contact with maxillary. | in contact with | | maxillary. ---------------+------------------------------+----------------------- table 4 (continued) ===============+=============================+========================= character | _s. phaeota_ | _s. puma_ ---------------+-----------------------------+------------------------ | | alary processes| two and one-half | two times as high as | times as high as lateral | lateral wing of | wing of premaxillary; | premaxillary; | anteriorly convex. | straight. | | | | nasals | long, widest anteriorly | short, narrow, not | and posteriorly, | attached to ethmoid. | bearing posteromedial | | process; not attached | | to ethmoid. | | | | | ethmoid | long, entirely ossified; | short, about two-thirds | smooth margins. | ossified; irregular | | margins. | | frontoparietal | fontanelle absent; | keyhole-shaped fontanelle; | large supraorbital | smooth margins; | flanges having | flanges absent. | straight edges and extending| | posterolaterally. | | | | | squamosal | large; anterior arm | small; anterior arm | extending 1/2-2/3 way | extending 1/2 way to | to maxillary. | maxillary. ---------------+-----------------------------+------------------------- table 4 (continued) ===============+===========================+============================ character | _s. sila_ | _s. sordida_ ---------------+---------------------------+--------------------------- | | alary processes| one and one-half | two and one-half | times as high as lateral | times as high as lateral | wing of premaxillary; | wing of premaxillary; | straight. | slightly convex | | anteriorly. | | nasals | short, wide, bearing | moderately long narrowest | small posteromedial | anteriorly and | processes; not attached | posteriorly; not attached | to ethmoid. | to ethmoid. | | | | | | ethmoid | moderately long; entirely | short; one-half to entirely | ossified; smooth | ossified; irregular | margins. | margins. | | frontoparietal | large, ovoid fontanelle; | large, elongate fontanelle; | smooth margins; | smooth margins; | flanges absent. | flanges absent. | | | | | | | | squamosal | moderately large; anterior| moderately small; anterior | arm extending | arm extending | 1/4 way to maxillary. | 1/4-2/3 way to maxillary. ---------------+---------------------------+---------------------------within the _baudini_ group, the skull of _s. cyanosticta_ is the most generalized of the three species; the cranial characters are intermediate between _s. phaeota_ and _s. baudini_. the lateral margins of the skull in _s. cyanosticta_ are gently curved, and have an angularity anterior to the palatine-maxillary suture; the anterior margins are less angular in _s. phaeota_, which has a broader snout. posteriorly in _s. baudini_ the margins are slightly curved medially, and the greatest width of the skull is between the quadratojugal-maxillary sutures on either side of the skull. the frontoparietals of _s. cyanosticta_ bear slightly irregular lateral margins and a large fontanelle. there is a tendency for obliteration of the fontanelle with increasing age in both _s. baudini_ and _s. cyanosticta_; the lateral margins of the frontoparietals bear large supraorbital flanges in both of these species. in _s. phaeota_ the flanges are most prominent; they extend posterolaterally with straight margins along two-thirds of the length of the orbit and terminate in rather blunt points. the broad interorbital flanges result in a relatively broad external interorbital distance. in _s. baudini_ the flanges are curved posterolaterally around the orbit and terminate in sharp, thin points. the tegmen tympani of all three species are massive. in _s. cyanosticta_ the proötics slope posteriorly, whereas they slope anteriorly in _s. baudini_ and _s. phaeota_. the skulls of _s. cyanosticta_ and _s. baudini_ are alike in certain respects. the squamosals of both species are large and connected to the maxillary by a bony connection; the squamosals of _s. phaeota_ are large, but extend only two-thirds of the distance from the dorsal end of the quadrate to the maxillary. in _s. baudini_ and _s. cyanosticta_ the nasals are separated throughout their lengths from the ethmoid, whereas the nasals of _s. phaeota_ are separated from the ethmoid by cartilage. the latter separation is due to an incomplete ossification of the nasals in _s. phaeota_. the bony part of each nasal is constricted in the middle of the long axis of the bone, and the nasals are widest anteriorly; posteriorly each nasal bears a medial process, which is narrowly separated from the lateral edge of the ethmoid. table 5.--variation in the number of teeth in the species of smilisca. (all are males; n = number of jaws, or twice the number of individuals; means are given in parentheses after the observed ranges.) =================+====+==============+==============+=========== species | n | maxillary | premaxillary | vomerine -----------------+----+--------------+--------------+---------- _s. baudini_ | 20 | 49-65 (56.0) | 9-16 (13.6) | 5-9 (7.2) _s. cyanosticta_ | 8 | 50-64 (57.9) | 10-12 (10.8) | 4-11 (7.1) _s. phaeota_ | 20 | 50-68 (58.1) | 10-15 (12.1) | 5-9 (7.3) _s. puma_ | 6 | 60-67 (63.6) | 11-13 (12.0) | 4-7 (5.3) _s. sila_ | 8 | 48-60 (52.9) | 10-14 (11.3) | 5-7 (5.7) _s. sordida_ | 12 | 39-55 (44.2) | 7-11 (9.3) | 4-6 (5.2) -----------------+----+--------------+--------------+----------the teeth of all species of _smilisca_ are spatulate and bifid. the numbers of maxillary, premaxillary, and vomerine teeth are summarized in table 5. smaller and presumably younger specimens of all species of _smilisca_ have fewer teeth than do larger specimens of the same species. this correlation between size and number of teeth does not exist as an interspecific trend within the genus; for example, the smallest species in the genus, _s. puma_, has the highest number of maxillary teeth. in small specimens of a given species wide gaps are present between the maxillary teeth posteriorly; in large specimens the gaps are filled by teeth, beginning anteriorly and progressing posteriorly, until the maxillary dentition is continuous. musculature no extensive study of the muscular system was undertaken, but certain muscles know to be of taxonomic importance were studied. _jaw musculature._--starrett (1960) pointed out the unique jaw musculature in _smilisca_. in this genus m. depressor mandibulae consists of two parts, one arising from the dorsal fascia and one from the posterior arm of the squamosal. two muscles arise from the anterior arm of the squamosal and insert on the lateral face of the mandible. of these muscles, m. adductor mandibulae posterior subexternus lies medial to the mandibular branch of the trigeminal nerve; the other, m. adductor mandibulae externus superficialis, lies lateral to the same nerve (fig. 9). in most other hylids the latter muscle is absent. no significant variation in the position of the muscles was noted in the various species of _smilisca_, though m. adductor mandibulae originate somewhat more anteriorly in _s. baudini_ and _s. cyanosticta_ than in the other members of the genus, all of which have a shorter anterior arm of the squamosal that does not reach the maxillary. the two separate parts of m. depressor mandibulae are not so widely separated in members of the _sordida_ group as in the _baudini_ group. [illustration: fig. 9. lateral view of the left jaw of _smilisca baudini_; _a. m. e. s._, adductor mandibulae externus superficialis; _a. m. p. s._, adductor mandibulae posterior subexternus; _col._, columella; _d. m._ depressor mandibulae; _m. b. t. n._, mandibular branch trigeminal nerve; _sq._, squamosal. ku 64214, ×5.] [illustration: fig. 10. ventral view of throat musculature in an adult male _smilisca baudini_ (superficial musculature on left, deep musculature on right); _a. c._ anterior cornua of hyoid; _gen. l._, geniohyoideus lateralis; _gen. m._, geniohyoideus medialis; _hyo._, hyoglossus; _omo._, omosternum; _pet._, petrohyoideus; _s._, submentalis; _sm._, submaxillaris; _st._, sternohyoideus; _v. s._, vocal sac. ku 64220, × 2.5.] _throat musculature._--the frogs that comprise the genus _smilisca_ are characterized by paired subgular vocal sacs, essentially the same as those in _triprion_ (duellman and klaas, 1964). the following description is based on _smilisca baudini_ (fig. 10). m. submentalis lies in the anterior angle of the lower jaw, is thick, and consists of transverse fibers extending between the dentaries. m. submaxillaris is thin and arises from the whole of the inner surface of the lower jaw, except for the anterior angle occupied by m. submentalis. anteriorly m. submaxillaris is broadly attached by fascia to m. hyoglossus and m. geniohyoideus, which lie dorsal to m. submaxillaris. medially this attachment continues posteriorly for about one-half the length of the hyoglossus. posteriorly m. submaxillaris is folded and attached to m. sternoradialis of the pectoral girdle. the vocal sacs are formed by a pair of posterolateral evaginations of m. submaxillaris; a broad connection between the pouches allows free passage of air between the pouches. the deeper throat musculature is essentially the same as that described for _phrynohyas spilomma_ by duellman (1956), except for slight differences in the place of attachment on the hyoid. skin _structure_ the skin of _smilisca_ is typical of that of most hylids in organization and structure. _smilisca sila_ is distinguished from other members of the genus by the presence of small wartlike protrusions and peculiar white, pustular spots on the dorsum. the wartlike structures are composed of three or four epidermal cells, which protrude from the surface of the epidermis; the structures are covered by a slightly thickened layer of keratin. the white pustules are slightly elevated above the surrounding skin. internally they consist of aggregations of swollen, granular, pigment-cells (perhaps lipophores) lying between the epidermis and the melanophores. _biochemical variations_ dried skins of all species of _smilisca_ were sent to josé m. cei, instituto nacional de cuyo, mendoza, argentina, for biochemical screening by means of the chromatographic techniques described by erspamer and cei (1963). the species in the _baudini_ group have detectable amounts of penta-hydroxi-trypatamine, whereas only a trace is present in the other species. furthermore, species in the _baudini_ group differ from _s. sila_ and the _sordida_ group in lacking, or having only a trace of, tryptophan-containing polypeptides. these superficial biochemical tests support the arrangement of species as ascertained by conventional taxonomic characters. external morphological characters the features of external morphology that were studied in connection with the taxonomy of the genus _smilisca_ are discussed below. _size and proportions_ the frogs of the genus _smilisca___ are medium to large tree frogs. the three species comprising the _baudini_ group (_s. baudini_, _cyanosticta_, and _phaeota_) are notably larger than _s. puma_, _sila_, and _sordida_ (table 6). the largest specimen that we examined is a female of _s. baudini_ having a snout-vent length of 90 mm. _smilisca puma_ is the smallest species; the largest male has a snout-vent length of 38 mm. and the largest female, 46 mm. table 6.--comparison of sizes and certain proportions of the species of smilisca. (means in parentheses below observed ranges; data for males only.) ================+====+===========+=============+===========+ | | snout-vent|tibia length/| tympanum/ | species | n | length | snout-vent | eye | ----------------+----+-----------+-------------+-----------+ | | | | | _s. baudini_ |140 | 47.3-75.9 | 42.1-53.6 | 56.1-94.4 | | | (58.7) | (47.8) | (73.5) | | | | | | _s. cyanosticta_| 40 | 44.6-56.8 | 51.9-59.7 | 62.7-88.4 | | | (50.7) | (56.0) | (71.4) | | | | | | _s. phaeota_ | 50 | 40.8-65.5 | 50.9-60.2 | 62.7-85.5 | | | (53.9) | (55.5) | (76.6) | | | | | | _s. puma_ | 20 | 31.9-38.1 | 48.2-53.1 | 52.1-72.2 | | | (34.7) | (51.3) | (64.9) | | | | | | _s. sila_ | 33 | 31.6-44.8 | 49.7-58.1 | 47.6-58.3 | | | (37.7) | (54.8) | (53.2) | | | | | | _s. sordida_ | 55 | 31.9-44.6 | 50.5-57.1 | 46.5-57.1 | | | (37.9) | (53.4) | (49.1) | ----------------+----+-----------+-------------+-----------+ no outstanding differences in proportions exist between species, although certain proportions are sufficiently different in some species to warrant mention. _smilisca baudini_ is a more squat and stocky frog than other members of the genus; this is reflected in the somewhat shorter hind legs (table 6). the size of the tympanum relative to that of the eye is highly variable within samples of a given species. even so, noticeable differences in the tympanum/eye ratio are apparent. members of the _baudini_ group have the largest tympani, whereas _s. sila_ and _sordida_ have the smallest, and _s. puma_ is intermediate (table 6). _shape of snout_ although all members of the genus have rather truncate snouts, subtle differences exist among the species (pl. 12). _smilisca sila_ has the shortest snout; that of _s. baudini_ is only slightly longer. the snouts of _s. cyanosticta_ and _puma_ are nearly square in lateral profile, whereas those of _s. phaeota_ and _sordida_ are slightly inclined. the shape of the snout is relatively uniform within each species and displays no noticeable sexual dimorphism, except in _s. sordida_, in which there are sexual differences and geographic variation (see p. 324). _hands and feet_ the characters of the hands and feet are among the most taxonomically important external features in _smilisca_. consistent differences exist in relative lengths of the digits, size of subarticular tubercles, size and number of supernumerary tubercles, size and shape of the inner metatarsal tubercle, and amount of webbing (pls. 4 and 5). in the _baudini_ group the series of species (_baudini-phaeota-cyanosticta_) show a progressive increase in amount of webbing in the hand and a decrease in number, and corresponding increase in size, of supernumerary tubercles. the amount of webbing in the feet of _s. baudini_ and _phaeota_ is about the same, but the webbing is slightly more extensive in _s. cyanosticta_. _smilisca puma_ is unique in the genus in lacking webbing in the hand; furthermore, this species is distinctive in having many large subarticular tubercles on the hand and a relatively small inner metatarsal tubercle. the two stream-inhabitants, _s. sila_ and _sordida_, have shorter and stouter fingers than the other species. the webbing is most extensive in both the hands and feet of these species, which also are distinctive in having many small supernumerary tubercles on the feet. _ontogenetic changes_ minor ontogenetic changes in structure involve the shape of the snout, relative size of the eye, development of the tympanum, and amount of webbing in the hand. in recently metamorphosed young the snout is more rounded than in adults; the canthus and loreal concavity are not evident. usually the tympanum is not differentiated in recently metamorphosed young, and the eye is proportionately large. the webbing in the feet is completely developed at metamorphosis, but young individuals have noticeably less webbing in the hand than do adults of the same species. coloration some of the most distinctive characters of the species of _smilisca_ are color and pattern of the living frogs. although many chromatic features are lost or subdued in preserved specimens, the patterns usually persist. _metachrosis_ change in color, well known in frogs, is common in hylids, especially in species having green dorsal surfaces (_phyllomedusa_ is a notable exception). the non-green _smilisca_ (_puma_, _sila_, and _sordida_) changes color, but this mostly is a change in intensity of color. in these species the markings usually are most distinct at night; frequently by day the frogs become pallid. the most striking examples of metachrosis in _smilisca_ are found in the _baudini_ group, in which the dorsal ground-color changes from green to tan; correlated with the change in ground-color may be a corresponding change in the dorsal markings, but the dorsal markings may change to the opposite color. chromosomes chromosomes of all six species of _smilisca_ were studied by means of the propriono-orcein squash technique described by duellman and cole (1965). karyotype analysis was attempted for several species by means of intraperitoneal injections of colchicine, which affected the mitotic cells as desired, but the testes examined contained too few mitotic cells to allow accurate determination of karyotypes. haploid (_n_) chromosome numbers were determined from cells in diakinesis, metaphase i, and metaphase ii of meiosis. diploid (2_n_) chromosome numbers were determined from cells in late prophase and metaphase of mitosis. chromosome counts from as few as 23 meiotic cells of _s. phaeota_ and as many as 80 cells of _s. sordida_ reveal a constant haploid (_n_) number of 12; counts of chromosomes in one to five mitotic cells in all species, except _s. sila_, reveal that the diploid (2_n_) number is 24. natural history breeding like most hylid frogs _smilisca_ is most readily collected and observed when individuals congregate for breeding. _time of breeding_ _smilisca_ breeds primarily in quiet water and reaches its height of breeding activity at times of plentiful rainfall,--usually from may through october. through most of its range _smilisca baudini_ breeds in those months, but in some places where abundant rain falls in other seasons, the species breeds at those times. for example, in southern el petén and northern alta verapaz, guatemala, _smilisca baudini_ has been found breeding in february and march. the other pond-breeding species (_s. cyanosticta_, _phaeota_, and _puma_) live in regions lacking a prolonged dry season, and possibly they breed throughout the year, but breeding activity seems to be greatest in the rainiest months. the two stream-breeding species (_s. sila_ and _sordida_) breed in the dry season when the streams are low and clear, principally in december through april. at high elevations the species sometimes breed in the rainy season; also, individuals sometimes breed in the short dry season (summer canicula) in july and august. at several localities species have been found breeding at different times of the year: _s. baudini_ in march and july at chinajá, guatemala; _s. phaeota_ in april and august at palmar sur, costa rica; _s. puma_ in february and july at puerto viejo, costa rica; and _s. sila_ in february, april, and august at el volcan, panamá. these observations indicate only that the population breeds at more than one time in the year, but do not provide any evidence on the breeding cycles of the individual frogs. this is one important aspect of the natural history of _smilisca_ for which we lack data. _breeding sites_ all members of the genus _smilisca_ presumably deposit their eggs in water. _smilisca baudini_ usually breeds in temporary rain pools; often these are nothing more than shallow, muddy puddles. in other instances the sites are extensive ditches or large flooded areas (pl. 8, fig. 1). this species is an opportunistic breeder, and males gather at any of a wide variety of suitable breeding sites that are formed by torrential rains in the early part of the rainy season. _smilisca baudini_ nearly always breeds in open pools having bare earthen edges. frequently congregations of _s. baudini_ are found at such small pools, but are absent from nearby large ponds surrounded by vegetation. little is known of the breeding habits of _s. cyanosticta_, which inhabits humid forests on foothills and lowlands. apparently its breeding sites are not unlike those of _s. phaeota_, which usually are pools surrounded by vegetation (pl. 8, fig. 2), although sometimes males of _s. cyanosticta_ call from open muddy puddles. in uplands, where standing water is uncommon, this species breeds in quiet pools in streams. _smilisca puma_ breeds in grass-choked ponds and marshes, where the males call from bases of dense clumps of grass in the water (pl. 9, fig. 1). _smilisca sila_ and _s. sordida_ differ noticeably from other species in the genus by breeding exclusively in streams, where males usually call from rocks or gravel bars in or at the edges of streams (pl. 9, fig. 2); sometimes individuals perch on bushes overhanging streams. in the streams, or parts of streams, utilized by these frogs the water is clear, shallow, and has a slow gradient; occasional males have been found calling along cascading mountain streams. breeding choruses composed of ten or more species of frogs are not uncommon in middle america, but _smilisca_ usually breeds alone or with one or two other species and at the most five others. this tendency towards solitary breeding possibly is the result of selection of breeding sites that are unsuitable to many other species of frogs. nevertheless, many other species of frogs have been found at the breeding sites with the various species of _smilisca_; these breeding associates (table 7) are most numerous for _s. baudini_, which has a broad geographic range, including a variety of habitats. _breeding behavior_ _calling sites._--all species of _smilisca_ usually call from the ground, including rocks and gravel bars; some individuals sit in shallow water near the edge of the pool or stream. sometimes males of _s. baudini_, _sila_, and _sordida_ call from low bushes or trees near the breeding site. one _s. baudini_ was observed calling while it was floating on the surface of a pond. _smilisca cyanosticta_, _phaeota_, and _puma_ call from secluded places at the edge of the water or in the water, whereas _s. baudini_, _sila_ and _sordida_ call from open situations. table 7.--breeding associates of the various species of smilisca. ==============================+========+============+========+=====+=====+======== associate |_s. |_s. |_s. |_s. |_s. |_s. |baudini_|cyanosticta_|phaeota_|puma_|sila_|sordida_ ------------------------------+--------+------------+--------+-----+-----+------- _rhinophrynus dorsalis_ | x | | | | | _leptodactylus bolivianus_ | | | x | | | _leptodactylus labialis_ | x | | x | | | _leptodactylus melanonotus_ | x | | x | x | x | _leptodactylus occidentalis_ | x | | | | | _leptodactylus quadrivittatus_| | | x | | | _leptodactylus pentadactylus_ | | | x | x | | x _engystomops pustulosus_ | x | | x | | | _bufo canaliferus_ | x | | | | | _bufo cavifrons_ | | x | | | | _bufo coccifer_ | x | | | | | _bufo coniferus_ | | | x | | | _bufo cristatus_ | | x | | | | _bufo gemmifer_ | x | | | | | _bufo haematiticus_ | | | x | | x | x _bufo kellogi_ | x | | | | | _bufo luetkeni_ | x | | | | | _bufo marinus_ | x | | x | x | x | x _bufo marmoreus_ | x | | | | | _bufo mazatlanensis_ | x | | | | | _bufo melanochloris_ | | | x | | x | x _bufo perplexus_ | x | | | | | _bufo typhonius_ | | | x | | x | _atelopus varius_ | | | | | x | x _diaglena reticulata_ | x | | | | | _diaglena spatulata_ | x | | | | | ------------------------------+--------+------------+--------+-----+-----+------- table 7.--_continued_ ==============================+========+============+========+=====+=====+======== associate |_s. |_s. |_s. |_s. |_s. |_s. |baudini_|cyanosticta_|phaeota_|puma_|sila_|sordida_ ------------------------------+--------+------------+--------+-----+-----+------- _hyla boulengeri_ | | | x | | | _hyla colymba_ | | | | | x | _hyla ebraccata_ | x | | x | | | _hyla elaeochroa_ | | | x | x | | _hyla eximia_ | x | | | | | _hyla legleri_ | | | | | | x _hyla microcephala_ | x | | x | | | _hyla phlebodes_ | | | x | x | | _hyla picta_ | x | | | | | _hyla robertmertensi_ | x | | | | | _hyla rosenbergi_ | | | x | | | _hyla rufioculis_ | | | | | | x _hyla smithi_ | x | | | | | _hyla staufferi_ | x | | | | | _hyla walkeri_ | x | | | | | _phrynohyas inflata_ | x | | | | | _phrynohyas spilomma_ | x | | | | | _phrynohyas venulosa_ | x | | | | | _phyllomedusa callidryas_ | x | | x | | | _phyllomedusa dacnicolor_ | x | | | | | _phyllomedusa moreleti_ | x | x | | | | _pternohyla fodiens_ | x | | | | | _smilisca baudini_ | x | x | | | | _smilisca cyanosticta_ | x | x | | | | _smilisca phaeota_ | | | x | | | _smilisca puma_ | | | | x | | ------------------------------+--------+------------+--------+-----+-----+------- table 7.--_concluded_ ==============================+========+============+========+=====+=====+======== associate |_s. |_s. |_s. |_s. |_s. |_s. |baudini_|cyanosticta_|phaeota_|puma_|sila_|sordida_ ------------------------------+--------+------------+--------+-----+-----+------- _smilisca sila_ | | | | | x | x _smilisca sordida_ | | | x | | x | x _triprion petasatus_ | x | | | | | _cochranella fleischmanni_ | | | | | x | x _centrolene prosoblepon_ | | | | | x | _gastrophryne elegans_ | x | | | | | _gastrophryne olivacea_ | x | | | | | _gastrophryne usta_ | x | | | | | _hypopachus alboventer_ | x | | | | | _hypopachus caprimimus_ | x | | | | | _hypopachus inguinalis_ | x | | | | | _hypopachus maculatus_ | x | | | | | _hypopachus oxyrrhinus_ | x | | | | | _hypopachus variolosus_ | x | | | | | _rana palmipes_ | x | | x | x | | _rana pipiens_ | x | | | | | _rana warschewitschi_ | | | x | | x | x ------------------------------+--------+------------+--------+-----+-----+-------_chorus structure._--limited observations on some of the species of _smilisca_ show a definite organization of the calling behavior of individuals. _smilisca baudini_ and _s. phaeota_ call in duets. this is especially noticeable in _s. baudini_, in which the members of a duet often call from sites separated by only a few centimeters. the call of _s. baudini_ consists of a series of like notes (see description of call in following section); the duration of each note is about equal to the interval between notes. normally one individual utters one note, pauses, and utters a single note again, or series of two or three notes. if there is no response, the first individual often waits several seconds or even several minutes and then repeats the call. the second individual usually responds after the first or second note of the sequence. the notes of the second individual usually are spaced so that they are emitted in the intervals between the notes of the first individual. this can be shown diagrammatically by having the figure "1" represent notes of the first individual and figure "2," the notes of the second; an empty interval is represented by "0": 1-0-1-2-1-2-1-2-1-2-1-2 usually a chorus is initiated by one duet and is quickly picked up by other individuals also calling in duets. a numerical representation of a chorus of eight frogs would approximate the following organization: 1-0-1-2-1-2-1-2-1-2-1-2-1-2 3-0-3-4-3-4-3-4-3-4-3-4-3 5-6-5-6-5-6-5-6-5-6-5-6 7-8-7-8-7-8-7-8-7-8-7-8 after the first one or two duets are initiated, the second individuals in the following duets usually call immediately after their respective partners have given the first notes. the other noteworthy aspect about the organization is that the entire chorus usually stops abruptly. normally the first duet stops calling shortly before the others, but this is not invariable. often one duet or one individual will emit several notes after the rest of the frogs have become silent. an interval of several minutes sometimes elapses before the chorus begins again. successive choruses apparently are initiated by the same duet. responses can be initiated artificially by imitating the call, and sometimes any loud noise will start a chorus. similar duets have been observed in _s. phaeota_. in this species the intervals are often much longer than the notes, and if two males are calling in close proximity, their calls can be mistaken for those of one individual. _smilisca phaeota_ does not congregate in large numbers; usually only two males call from one restricted site. _smilisca sila_ has a call consisting of a primary note followed by one or more secondary notes. males often call in duets, but not necessarily so. in a duet, the first male usually utters only primary notes until the second individual responds; then each individual produces a rapid series of secondary notes. _smilisca puma_ also produces primary and secondary notes. although individuals sometimes call alone, duets, trios, or quartets were more common. the chorus is initiated by one individual uttering primary notes until joined by the second, third, and fourth frogs. in one quartet in a marsh 7.5 kilometers west of puerto viejo, costa rica, on february 19, 1965, the same individual initiated four consecutive choruses. each time the second member of the chorus was the same; the third and fourth frogs joined the chorus nearly simultaneously. individuals of _s. sordida_ are usually irregularly situated along a stream. no duets or other combinations of individuals are apparent in the chorus structure, but once an individual calls, a frog nearby calls almost immediately; then a frog near the second individual calls, and so on. the resulting series of calls gives the impression that the sound is moving along the stream as successive individuals join the chorus and the first callers become quiet. it is not known if the same individual initiates successive choruses or if the order of calling is the same in subsequent choruses. these limited observations on chorus structure in _smilisca_ show the presence of behavioral organization. the methods of establishing the organization and the significance of the call-order in breeding have yet to be discovered. calling males of _s. baudini_ are often close together; some individuals have been observed almost touching one another, but no indication of territoriality or aggressive behavior has been witnessed. the more distant spacing of the stream-breeding species _s. sila_ and _s. sordida_ may be a function of calling-territories, but no direct evidence is available to substantiate this supposition. _sex recognition and amplexus._--observations on _smilisca baudini_ indicate that the calls of males attract females. at tehuantepec, oaxaca, méxico, a female was first observed about two meters away from a male calling at the edge of a rain pool; in a series of short hops she progressed directly towards the male, although vegetation obscured him until she was less than a meter away. when she approached to within about 20 centimeters of the male, he took notice of her, moved to her, and clasped her. at chinajá, alta verapaz, guatemala, a female swam directly across a pool about three meters wide to a calling male. her line of movement took her within a few centimeters of a silent male, to whom she paid no attention. she stopped just in front of the calling male, which immediately clasped her. at a large muddy pond 4 kilometers west-northwest of esparta, puntarenas, costa rica, a female was observed swimming toward a small submerged tree; a male was calling from a branch about one meter above the water. the female climbed to a branch about 20 centimeters below the male, which upon seeing her there immediately jumped down and clasped her. these few observations of _s. baudini_ show that in this species females are capable of locating calling males by means of phono-orientation; visual reception on the part of females seems to be secondary. contrariwise, males apparently become aware of the proximity of females by seeing them; once a male sees a female he usually tries to clasp her. possibly the males receive stimuli by means of chemo-reception, but in each observed instance the male obviously looked at the female. amplexus is axillary in all members of the genus. normally amplexing males hunch their backs and press their chins to the females' backs. clasping pairs are usually found at the edge of the water, but sometimes amplexus takes place in trees or bushes. _egg deposition._--oviposition has been observed only in _smilisca baudini_. on the night of june 28, 1961, at chinajá, alta verapaz, guatemala, a clasping pair was observed at the edge of a shallow rain pool. after sitting for several minutes in shallow water, the female (with male on her back) swam part way across the pool and grasped an emergent stick with one hand. the female's body was nearly level with the surface of the water, and her hind legs were outstretched as deposition commenced; eggs were extruded rapidly. after a few seconds the female moved slowly to another twig a few centimeters away and deposited more eggs. this process was repeated until the female was spent. the spawn resulted in a surface film covering roughly one square meter. it is doubtful if this type of egg deposition occurs in any other species in the genus, especially those that lay their eggs in streams. _breeding call_ the breeding calls of the six species of _smilisca_ are alike in their explosive nature. calls are emitted quickly with a short burst of air filling the vocal sac, which immediately deflates. phonetically the calls can be described as a single "wonk" or series of such notes in _s. baudini_ and _s. cyanosticta_, a low growl in _s. phaeota_, a relatively high pitched rattle in _s. sordida_, and a low squawk usually followed by one or more rattling secondary notes in _s. puma_ and _s. sila_. quantitatively, the calls of the six species differ in number of notes, duration of notes, and in pitch (table 8, pls. 10 and 11). although no measurements were taken on the intensity of the calls, we observed in the field that each of the species has a loud voice. the call of _s. baudini_ seems to carry farther than any of the others. table 8.--comparison of breeding calls in smilisca. (observed range given in parentheses below mean. in species having primary and secondary notes, only the primary notes are analyzed here.) ==============+====+=======+===========+=========+=========+=======================+ | | notes | | | funda | major | species | | per | duration | pulses | mental | frequencies (cps) | | n | call | of note | per |frequency+-----------+-----------+ | | group | (seconds) | second | (cps) | lower | upper | --------------+----+-------+-----------+---------+---------+-----------+-----------+ _s. | 20 | 8.0 | 0.11 | 174.7 | 166.2 | 351 | 2507 | baudini_ | | (2-15)|(0.09-0.13)|(140-195)|(135-190)| (175-495) |(2400-2725)| | | | | | | | | _s. | 10 | 1.2 | 0.38 | 147.0 | 145.1 | 841 | 1894 | cyanosticta_| | (1-2) |(0.25-0.45)|(110-180)|(135-160)| (480-975) |(1600-2100)| | | | | | | | | _s. phaeota_ | 10 | 1.6 | 0.31 | 116.0 | 143.0 | 372 | - | | | (1-2) |(0.10-0.45)|(100-130)|(110-165)| (330-495) | - | | | | | | | | | _s. puma_ | 28 | 3.7 | 0.13 | 208.2 | 145.6 | 743 | 1868 | | | (2-10)|(0.06-0.35)|(187-240)|(125-200)| (495-980) |(1456-2240)| | | | | | | | | _s. sila_ | 15 | 2.4 | 0.16 | 108.5 | 103.0 | 899 | 2218 | | | (1-6) |(0.06-0.28)| (97-120)| (90-115)| (665-1180)|(1980-2700)| | | | | | | | | _s. sordida_ | 19 | 1.7 | 0.29 | 104.7 | 123.1 | 1216 | 2694 | | | (1-6) |(0.18-0.45)| (78-135)| (90-140)|(1150-1540)|(2340-2990)| --------------+----+-------+-----------+---------+---------+-----------+-----------+ _call rate._--the rate at which call-groups are produced varies from one every few seconds to one in several minutes. in _s. baudini_, _cyanosticta_, _phaeota_, and _sordida_, call-groups are produced as frequently as every 12 seconds, but usually more time elapses between call groups. in _s. sordida_, five or more minutes sometimes elapse between call-groups. the interval is somewhat less in _s. phaeota_. calls are repeated at much shorter intervals in _s. puma_ (5-55 seconds) and _s. sila_ (4-20 seconds). _notes per call-group._--except for _s. puma_ and _s. sila_, the series of notes produced in any given call of a species of _smilisca_ is essentially the same; there is no differentiation into primary and secondary notes. _smilisca cyanosticta_ and _s. phaeota_ emit only one or two relatively long notes per call-group, whereas _s. baudini_ and _s. sordida_ produce as many as 15 and 6 notes, respectively. males of _s. puma_ and _s. sila_ often produce only the primary note; sometimes this is done several times before the secondary notes are produced. for example, one _s. puma_ (ku 91711; tape no. 379) produced the following number of notes in consecutive call-groups: 1, 1, 1, 1, 2, 2, 3, 1, 4; secondary notes are present in only four of the nine call-groups. a typical series of consecutive call-groups in _s. sila_ (ku 91852; tape no. 385) has 1, 1, 1, 2, 4, 2 notes per call-group; secondary notes are present in only half of the call-groups. _smilisca puma_ apparently always produces at least two primary notes before emitting secondary notes; sometimes only primary notes are produced in one series of calls. the number of secondary notes following a given primary varies from one to nine; the modal number is one, and the mean is three in 27 call-groups. _smilisca sila_ frequently begins a series of calls with two or more primary notes, but sometimes the first primary note is followed immediately by two or more secondary notes. the number of secondary notes following a given primary varies from one to five; the modal number is one, and the average is two in 13 call-groups. _duration._--the average duration of call-groups consisting of two or more notes is 1.18 seconds in _s. baudini_; 1.02 in _cyanosticta_, 0.91 in _phaeota_, 1.32 in _puma_, 1.48 in _sila_, and 1.29 in _sordida_. although there is considerable variation in the lengths of the notes (only primary notes in _s. puma_ and _sila_ are considered here), _s. cyanosticta_, _phaeota_, and _sordida_ have noticeably longer notes than do the other species (table 8). the secondary notes are longer than the primary notes in _s. puma_ (average 0.27 secs. as compared with 0.13 secs.) and in _s. sila_ (average 0.25 secs., as compared with 0.16 secs.). _note repetition rate._--the rate at which notes in call-groups containing two or more notes are produced varies in _s. baudini_ from 2.5 to 7.1 (average, 3.7) calls per second; _cyanosticta_, 1.8-2.1 (1.9); _phaeota_, 2.0-2.4 (2.2); _puma_, 1.9-2.9 (2.2); _sila_, 1.3-2.4 (1.8); and _sordida_, 1.5-2.6 (2.1). _smilisca baudini_, which has notes of short duration (0.09 to 0.13 seconds), has the fastest note-repetition rate. although the individual notes of _s. cyanosticta_ and _s. phaeota_ are relatively long (average, 0.38 and 0.31 seconds, respectively), the intervals between the notes is short; consequently, their note-repetition rates do not differ greatly from those of _s. puma_ and _s. sila_, which have shorter notes (average, 0.13 and 0.16 seconds, respectively) but longer intervals between notes. _pulse rate._--pulses vary in frequency from 78 to 240 per second in the calls analyzed (only primary notes in _s. puma_ and _s. sila_), but the variation in any given species is much less than that in the entire genus (table 8). _smilisca puma_ is outstanding in having a high pulse rate, which is approached only by that of _s. baudini_. even in the species having the lowest pulse rates, the pulsations are not audible. the secondary notes produced by _s. puma_ and _s. sila_ have a slower pulse rate than the primary notes; often the pulses are audible. in _s. puma_ the pulse rate of secondary notes is sometimes as low as 48 pulses per second, and in _s. sila_ still lower (as low as 40 pulses per second). the upper limits of pulse rate in the secondary notes in these species merge imperceptibly with the rates of the primary note; consequently, on the basis of pulse rate alone it is not always possible to distinguish primary from secondary notes. _frequency._--_smilisca_ produces noisy (as opposed to more musical) calls, and the energy is distributed throughout the frequency spectrum; the calls are poorly modulated, except in _s. sordida_, in which two usually discrete bands of frequency are present (pl. 11c). for the most part the calls of _smilisca_ consist of little modified energy of the fundamental frequency and of its harmonics, some of which are emphasized. the upper frequency range varies within each species and even within the calls of one individual. _smilisca phaeota_ has the lowest upper frequencies; no calls ranged above 4400 cycles per second (cps.), and half of the calls never exceeded 3000 cps. _smilisca cyanosticta_ produces calls in which the upper frequency is below 7000 cps. and usually below 6000 cps. likewise, _s. puma_ produces calls that are below 7000 cps., whereas _s. sila_ has frequencies of up to 8400 cps. in both _s. baudini_ and _s. sordida_, the highest frequencies attained are about 9100 cps. variation in the highest frequencies in a series of consecutive calls by one individual frog was noted in all species. such variation is especially prevalent in _s. puma_; for example one individual (ku 87771; tape no. 376) recorded at a temperature of 24° c. at 7.5 kilometers west of puerto viejo, heredia province, costa rica, on july 31, 1964, produced three consecutive primary notes having upper frequencies of about 6000, 4000, and 4000 cps., respectively. apparently in a given species the production of the higher frequencies in some notes and not in others is correlated with the amount of distention of the vocal sac and is not dependent upon the structure or tension of the vocal cords. although the dominant frequency in _s. sordida_ is lower than that in _s. baudini_ and _s. cyanosticta_, the call of the former is audibly higher-pitched. this is due primarily to the emphasis on certain harmonics at a high frequency (sometimes as high as 9000 cps.) in _s. sordida_, whereas in _s. baudini_ and other species, if harmonics are present at those frequencies, they are not emphasized. the fundamental frequencies are as low as 90 cps. in _s. sila_ and _s. sordida_ and as high as 200 cps. in _s. puma_ (table 8). the fundamental frequency seemingly is relatively unimportant in determining the general pitch of the call, a characteristic most dependent on the dominant frequency and emphasized harmonics in the higher-frequency spectrum. in none of the species is the fundamental the dominant frequency. in the low-pitched call of _s. phaeota_ the dominant frequency is the third harmonic (the second harmonic above the fundamental frequency, which is the first harmonic). in all other species a much higher harmonic is dominant; for examples, in _s. cyanosticta_ harmonics from 10 to 15 are dominant; in _s. baudini_, 15-19; and _s. sila_, 20-30. a glance at the audiospectrographs and their accompanying sections (pls. 10 and 11) reveals the presence of two emphasized bands of frequency in all species except _s. phaeota_, in which only the lower band is present. these two bands of emphasized harmonics are part of a continuous, or nearly continuous, spread of energy throughout the frequency spectrum, except in _s. sordida_ in which the bands are usually distinct. as shown in the sections, certain harmonics in each of the bands are emphasized with nearly equal intensity. therefore, with the exception of _s. phaeota_, the calls of _smilisca_ are characterized by two major frequencies, one of which is the dominant frequency and the other is a subdominant frequency (table 8). the upper major frequency is dominant in all calls in _s. baudini_ and _s. cyanosticta_, but either major frequency may be dominant in other species. the upper major frequency is dominant in 65 per cent of calls by _s. puma_, 87 per cent in _s. sila_, and 68 per cent in _s. sordida_. individuals of these three species sometimes produce a series of calls in which the dominant frequency changes from one of the major frequencies to the other. four consecutive notes emitted by an individual of _s. sordida_ recorded 13 kilometers east-northeast of golfito, puntarenas province, costa rica, had dominant frequencies of 910, 1950, and 750 cps., respectively. in each case, an alternation of major frequencies took place in respect to dominance. an individual of _s. puma_ from 7.5 kilometers west of puerto viejo, costa rica, produced a primary note followed by one secondary note; each note had major frequencies at 600 and 1800 cps.; the dominant frequency of the primary note was at 1800 cps., whereas in the secondary note the dominant frequency was at 600 cps. the difference in emphasis on the major frequencies is so slight that shift in dominance is not audible. _effect of temperature on calls._--the present data are insufficient to test statistically the correlation between temperature and variation within certain components of the calls in _smilisca_, but even a crude graph shows some general correlations. the widest range of temperatures is associated with the recordings of _s. baudini_. three individuals recorded at a temperature of 30° c. at tehuantepec, oaxaca, had pulse rates of 180 pulses per second and fundamental frequencies of 160-180 cps., as compared with an individual recorded at a temperature of 17° c., which had a pulse rate of 140 and a fundamental frequency of 135 cps. all individuals of _s. baudini_ recorded at higher temperatures had faster pulse rates and higher fundamental frequencies. pulse rates differ in the other species in the genus but less strikingly (probably owing to narrower ranges of temperatures at which recordings were made). in five recordings of _s. sordida_ made at 20° c. the pulse rate is 80-90, as compared with four recordings made at 25° c. having pulse rates of 120-135. thirteen recordings of _s. sila_ made at 17° c. have pulse rates of 97-112 (average 105); one individual recorded at 26° c. has 120 pulses per second. seemingly no correlation exists between temperature and other characteristics of the calls, such as duration and rate of note-repetition. _the breeding call as an isolating mechanism._--blair (1958), bogert (1960), duellman (1963a), fouquette (1960), johnson (1959), and others have provided evidence that the breeding calls of male hylids (and other anurans) serve as isolating mechanisms in sympatric species. in summarizing this discussion of the breeding calls of _smilisca_ we want to point out what seem to be important differences in the calls that may prevent interspecific hybridization in sympatric species of _smilisca_. the genus is readily divided into two species-groups on morphological characters; this division is supported by the breeding calls. in the species of the _baudini_ group the calls are unmodulated and lack secondary notes. in the _sordida_ group the calls either have secondary notes or are modulated. _smilisca baudini_ occurs sympatrically with _s. cyanosticta_ and _s. phaeota_; where they occur together, both species sometimes breed in like places at the same time. we are not aware of these species breeding synchronously at exactly the same site, although _s. baudini_ and _s. cyanosticta_ were calling on the same nights and less than 100 meters apart in oaxaca in june, 1964. regardless of their respective breeding habits, sympatric species have calls that differ notably. except for the higher fundamental and dominant frequencies, the calls of _s. cyanosticta_ and _s. phaeota_ closely resemble one another, but the calls of both species differ markedly from that of _s. baudini_. the geographic ranges of _s. cyanosticta_ and _s. phaeota_ are widely separated. the calls of the allopatric species _s. puma_ and _s. sila_ are not greatly different. _smilisca sordida_ has a distinctive call and occurs sympatrically with _s. puma_ and _s. sila_. in the streams in southern costa rica _s. sordida_ and _s. sila_ breed synchronously, but the high-pitched modulated call of the former is notably different from the lower, unmodulated call of _s. sila_. the data indicate that the calls of related sympatric species differ more than the calls of related allopatric species. we postulate that these differences evolved to support the reproductive isolation of the sympatric species. the data are insufficient to determine geographic variation in the calls and to determine if differences in the calls are enhanced in areas of sympatry as compared with the allopatric parts of the ranges. _other calls._--as stated previously, there is no direct evidence of territoriality in _smilisca_; we have heard no calls that can be definitely identified as territorial. single notes of _s. baudini_, _phaeota_, and _sila_ have been heard by day, just prior to rains, or during, or immediately after rains. such calls can be interpreted as "rain calls," which are well known in _hyla eximia_ and _hyla squirella_. distress calls are known in several species of _rana_ and in _leptodactylus pentadactylus_; such calls result from the rapid expulsion of air over the vocal cords and with the mouth open. distress calls have been heard from _s. baudini_. at charapendo, michoacán, méxico, a male that had one hind limb engulfed by a _leptodeira maculata_ emitted several long, high-pitched cries. a clasping pair of _s. baudini_ was found in a bush at the edge of a marshy stream 2 kilometers northeast of las cañas, guanacaste province, costa rica. when the pair was grasped, the female emitted a distress call. eggs eggs of _s. baudini_, _cyanosticta_, and _phaeota_ have been found in the field, and eggs of _s. sila_ have been observed in the laboratory. the eggs of _s. puma_ and _sordida_ are unknown. insofar as known, _smilisca baudini_ is unique in the genus in depositing the eggs in a surface film. each egg is encased in a vitelline membrane, but individual outer envelopes are lacking. the eggs are small; the diameter of recently-deposited eggs is about 1.3 mm. and that of the vitelline membrane is about 1.5 mm. the eggs of _s. cyanosticta_ and _phaeota_ are deposited in clumps, and the eggs are larger than those of _s. baudini_. diameters of eggs of _s. cyanosticta_ are about 2.3 mm., and those of the outer envelopes are about 4.0 mm. artificially fertilized eggs of _s. sila_ raised in the laboratory have diameters of about 2.4 mm.; the diameter of the outer envelopes is about 4.9 mm. in order to determine the reproductive potential of the six species, ovulated eggs were removed from females and counted. the numbers of eggs recorded are: 3 _s. baudini_--2620, 2940, 3320; 1 _s. cyanosticta_--910; 3 _s. phaeota_--1665, 1870, 2010; 1 _s. puma_--518; 3 _s. sila_--369, 390, 473; 3 _s. sordida_--524, 702, 856. these limited data indicate that the large species (_s. baudini_, _cyanosticta_, and _phaeota_) have more eggs than do the smaller species. the stream-breeding species (_s. sila_ and _sordida_) have relatively few eggs by comparison with the pond-breeders. possibly this is a function of size of eggs rather than a correlation with the site of egg-deposition. tadpoles the acquisition of tadpoles of all of the species of _smilisca_ has made possible the use of larval characters in erecting a classification and in estimating the phylogenetic relations of the several species. furthermore, developmental series of tadpoles of four species allow a comparison of the growth and development in these species. throughout the discussion of tadpoles we have referred to the various developmental stages by the stage numbers proposed by gosner (1960). _general structure_ tadpoles of the genus _smilisca_ are of a generalized hylid type, having 2/3 tooth-rows, unspecialized beaks, mouth partly or completely bordered by papillae, lateral fold present in the lips, spiracle sinistral, anal tube dextral, and caudal musculature extending nearly to tip of caudal fin. although minor differences exist in coloration, proportions, and mouthparts, no great modifications of the basic structure are present. _comparison of species_ the larval characters of the species of _smilisca_ are compared below and illustrated in figures 11-15. _shape and proportions._--the bodies of _s. baudini_, _cyanosticta_, _phaeota_, and _puma_ are rounded and about as wide as deep; the eyes are moderately large and directed dorsolaterally, and the nostrils are about midway between the bluntly rounded snout and the eyes. the mouths are medium-sized and directed anteroventrally. the bodies of tadpoles of _s. sila_ and _sordida_ are slightly compressed dorso-ventrally. the snout is moderately long and sloping; the eyes are larger and directed more dorsally than in the other species, and the nostrils are closer to the eyes than the snout. the mouths are moderately large and directed ventrally. the tail is about half again as long as the body in _s. baudini_, _cyanosticta_, _phaeota_, and _puma_; in these species the caudal musculature is moderately heavy, and the caudal fins are deep. the caudal musculature is upturned distally in _s. baudini_ and _phaeota_, and the dorsal fin extends anteriorly onto the body in these two species and in _s. puma_. the tail is about twice as long as the body in _s. sila_ and _sordida_. in both species the caudal fins are shallow in comparison with the depth of the caudal musculature, especially in _s. sordida_ (fig. 14); in neither species does the dorsal fin extend anteriorly onto the body. [illustration: fig. 11. tadpoles of _smilisca baudini_: (a) stage 21 (ku 62155) × 10; (b) stage 25 (ku 68467) × 5; (c) stage 30 (ku 60018) × 4; (d) stage 41 (ku 60018) × 3.] _mouthparts._--the mouth of _s. sordida_ is completely bordered by two rows of papillae, whereas in the other species the median part of the upper lip is devoid of papillae. _smilisca baudini_ and _puma_ have two rows of papillae; _s. sila_ has one complete row (except medially on the upper lip) and one incomplete row, and _s. cyanosticta_ and _phaeota_ have only one row (fig. 15). all species have numerous papillae in the lateral fold; the fewest lateral papillae are found in _s. cyanosticta_ and _phaeota_. although all species have two rows of teeth in the upper jaw and three rows in the lower jaw, specific differences in the nature of the rows exist between certain species. the second upper tooth-row is narrowly interrupted medially in _s. sila_ and _sordida_ and broadly interrupted in the other species. the first upper row is strongly arched in _s. puma_, moderately arched in _s. baudini_ and _sila_, and weakly arched in the other species. in all species the third lower tooth-row is the shortest, only slightly so in _s. sila_ and _sordida_, but only about half the length of the second lower row in _s. puma_. [illustration: fig. 12. tadpoles of _smilisca cyanosticta_: (a) stage 21 (ku 87648) (b) stage 25 (ku 87651) × 5; (c) stage 30 (ku 87652) × 4; (d) stage 40 (ku 87650) × 3.] the beaks are well developed and finely serrate in all species. the lower, broadly v-shaped, beak is slender in _s. puma_, rather robust in _s. baudini_ and _sila_, and moderately heavy in the other species. the lateral processes of the upper beak are shortest in _s. puma_ and longest in _s. baudini_ and _sordida_. in the latter the inner margin of the upper beak and lateral process have the form of a shallow s, whereas in the other species the inner margin of the upper beak forms a continuous arch with the lateral processes (fig. 15). [illustration: fig. 13. tadpoles of _smilisca phaeota_: (a) stage 21 (ku 68479) × 14; (b) stage 25 (ku 68480) × 5; (c) stage 30 (ku 68482) × 4; (d) stage 40 (ku 68483) × 3.] _coloration._--the tadpoles of _smilisca_ lack the bright colors or bold markings characteristic of some hylid tadpoles; even so, the subdued colors and arrangement of pigments provide some distinctive markings by which the species can be distinguished from one another. the species comprising the _baudini_ group (_s. baudini_, _cyanosticta_, and _phaeota_) are alike in having the body brown or grayish brown dorsally and transparent with scattered brown pigment ventrally. a cream-colored, crescent-shaped mark is present on the posterior edge of the body; this mark is usually most noticeable in _s. baudini_ and least so in _s. cyanosticta_. other differences in coloration in members of the _baudini_ group are relative and subtle. _smilisca phaeota_ usually is more pallid than _baudini_, and _cyanosticta_ usually is darker than _baudini_; both species have larger dark markings on the tail than does _s. phaeota_. _smilisca baudini_ has a dark streak on the middle of the anterior one-fourth of the tail (figs. 11-13). _smilisca puma_ is distinctive in having a grayish brown body and dark gray reticulations on the tail. _smilisca sila_ and _sordida_ are distinctive in having pairs (sometimes interconnected) of dark marks on the dorsal surfaces of the caudal musculature, and in dorsal view the tail appears to be marked with dark and pale creamy tan transverse bars. these dark marks, as well as the small flecks on the tail, are brown in _s. sila_ and red in _sordida_. _smilisca sila_ has dark brown flecks on the dorsal surface of the body and small white flecks laterally; these markings are absent in _s. sordida_ (fig. 14). descriptions of the coloration of living tadpoles are given in the accounts of the species. [illustration: fig. 14. tadpoles of _smilisca_; (a) _s. puma_, stage 30 (ku 91807); (b) _s. sila_, stage 25 (ku 80260); _s. sordida_, stage 30 (ku 68475). all × 3.5.] _growth and development_ information on the growth and development of middle american hylids is scanty. adequate descriptions have been published for _phyllomedusa annae_ (duellman, 1963b), _phrynohyas venulosa_ (zweifel, 1964), and _triprion petasatus_ (duellman and klaas, 1964). material is available for adequate descriptions of the developmental stages of four species of _smilisca_ (tables 9-12, figs. 11-13). because none of the tadpoles was raised from hatching to metamorphosis, the rate of growth and duration of the larval stages are unknown. [illustration: fig. 15. mouthparts of tadpoles of _smilisca_; (a) _s. baudini_ (ku 60018); (b) _s. puma_ (ku 91807); (c) _s. cyanosticta_ (ku 87625); (d) _s. sila_ (ku 80620); (e) _s. phaeota_ (ku 68482); (f) _s. sordida_ (ku 68475). all ×17.] table 9.--growth and development of tadpoles of smilisca baudini. (means are given in parentheses after the observed ranges.) ==================================================================== stage | n | total length | body length | tail length ------------------------------------------------------------------- 21 |10 | 5.1-5.4 (5.22) | 2.6-2.7 (2.54) | 2.5-2.7 (2.58) 24 |10 | 6.0-6.5 (6.20) | 2.3-2.6 (2.45) | 3.5-3.9 (3.69) 25 |10 | 7.2-8.3 (7.78) | 3.0-3.3 (3.14) | 4.2-5.0 (4.64) 27 |10 | 18.5-21.5 (20.22) | 8.0-9.0 (8.38) |10.4-13.0 (11.84) 29 |10 | 21.5-24.5 (22.60) | 8.5-10.0 (9.25) |12.5-14.5 (13.35) 37 | 3 | 28.5-31.0 (30.00) |11.0-12.5 (11.67) |17.5-19.0 (18.00) 38 |10 | 35.0-37.5 (35.50) |12.0-13.5 (12.80) |21.5-24.0 (22.70) 40 | 2 | 34.0-37.0 (35.50) |12.5-13.5 (13.00) |21.5-23.5 (22.50) 41 |10 | 34.0-37.0 (35.50) |12.5-13.5 (13.00) |21.5-23.5 (22.50) 42 | 3 | 24.0-30.0 (27.00) |12.5-13.0 (12.67) |11.5-17.0 (14.33) 45 | 6 | 14.0-24.0 (17.58) |12.5-14.0 (13.37) | 1.5-10.0 (4.17) 46 |23 | --- |12.0-15.5 (13.34) | --- ------------------------------------------------------------------- table 10.--growth and development of tadpoles of smilisca cyanosticta. (means are given in parentheses after the observed ranges.) ====================================================================== stage | n | total length | body length | tail length --------------------------------------------------------------------- 21 | 10| 5.8-6.5 (6.28) | 2.8-3.1 (3.00) | 3.0-3.5 (3.28) 25 | 10| 7.9-9.2 (8.44) | 2.7-3.2 (2.96) | 4.8-6.0 (5.48) 30 | 7| 22.5-25.0 (23.50) | 8.5-9.5 (9.00) |14.0-15.5 (14.57) 36 | 10| 27.0-30.0 (28.75) | 9.5-11.5 (10.80) |17.0-18.5 (17.95) 42 | 2| 26.0-27.0 (26.50) | 10.00 |16.0-17.0 (16.50) 46 | 2| - | 14.00 | - ---------------------------------------------------------------------hatchlings of three species (_s. baudini_, _cyanosticta_, and _phaeota_) are available. these larvae have non-functional eyes and large oral suckers. by the time the larvae have developed to stage 21, external gills are present, the caudal musculature and caudal fin have been differentiated, and the head is distinguishable from the body. in stage 21 oral suckers and a large amount of yolk are still present. the developmental data on the four species show no significant variations; consequently, we will describe the development of only one species, _smilisca phaeota_ (table 11, figs. 13 and 16). _stage 21._--bulging cream-colored yolk mass, transparent cornea, and moderately long, unbranched filamentous gills, and oral suckers present; mouth having irregular papillae on lower lip; teeth and beaks absent; caudal myomeres distinct; pigmentation uniform over body and caudal musculature; caudal fin transparent with scattered small flecks. [illustration: fig. 16. relative rate of growth in tadpoles of _smilisca phaeota_ as correlated with developmental stages. formulas for the limb bud refer to its length (l) in relation to basal diameter (d).] _stage 25._--operculum complete; gills absent; sinistral spiracle apparently functional; cloacal tail-piece, nasal capsules, and external nares present; gut partly formed; mouth bordered by single row of papillae, except medially; small papillae present in lateral fold of lips; two upper and three lower tooth-rows present, but not fully developed; beaks apparently fully developed; depth of dorsal and ventral fins less than depth of caudal musculature: tip of tail upturned; pigment on body most dense on dorsum and sides; faint, nearly pigmentless crescent-shaped mark on posterior edge of body; concentrations of pigment forming small spots on tail. _stage 28._--mouthparts complete; limb bud about half as long as thick; other structural features and coloration closely resemble those in stage 25. _stage 30._--limb bud approximately twice as long as thick; body as deep as wide; dorsal fin deepest just posterior to body; ventral fin deeper than caudal musculature; tail sharply upturned distally; anal tube dextral; brown pigment sparse on flanks. table 11.--growth and development of tadpoles of smilisca phaeota. (means are given in parentheses after the observed ranges.) ==================================================================== stage | n | total length | body length | tail length ------------------------------------------------------------------- 15 | 10| - | 1.9-2.1 (1.97) | - 16 | 8| - | 2.0-2.2 (2.07) | - 18 | 4| - | 2.2-2.6 (2.31) | - 21 | 3| 7.9-8.6 (8.21) | 4.1-4.5 (4.31) | 3.8-4.1 (3.92) 25 | 10| 8.7-10.6 (9.69) | 4.5-4.8 (4.64) | 4.3-6.0 (5.05) 26 | 11| 12.3-16.1 (14.01) | 4.2-6.3 (5.60) | 6.7-9.8 (8.41) 27 | 10| 13.0-15.7 (14.28) | 4.9-6.2 (5.40) | 7.7-10.5 (8.88) 28 | 13| 13.9-20.9 (15.62) | 5.2-8.3 (5.75) | 8.5-12.6 (9.85) 29 | 8| 17.8-22.3 (19.79) | 6.3-8.4 (7.19) | 11.5-14.0 (12.60) 30 | 9| 20.3-24.8 (22.85) | 8.1-10.5 (9.32) | 10.5-15.5 (13.53) 31 | 5| 24.1-28.5 (26.61) | 9.4-11.2 (10.59) | 14.7-17.3 (16.02) 34 | 5| 24.8-29.4 (27.31) | 9.2-11.6 (10.73) | 15.6-18.5 (16.80) 36 | 3| 30.0-30.1 (30.07) |10.1-12.2 (11.15) | 18.9-20.0 (19.44) 37 | 4| 28.9-34.1 (31.75) |11.5-12.4 (11.88) | 17.4-22.5 (19.88) 38 | 1| 28.98 |12.88 | 16.10 39 | 2| 35.6-36.9 (36.25) |14.00 | 21.6-22.9 (22.25) 40 | 2| 32.3-39.8 (36.05) |14.00 | 18.3-21.8 (20.05) 43 | 2| 21.5-23.0 (22.25) |14.2-14.8 (14.45) | 6.8-8.8 (7.80) 44 | 4| - |14.5-15.6 (15.08) | - 46 | 11| - |12.7-16.7 (14.26) | - ------------------------------------------------------------------- table 12.--growth and development of tadpoles of smilisca sordida. (means are given in parentheses after the observed ranges.) =======+===+==================+==================+================= stage | n | total length | body length | tail length -------+---+------------------+------------------+---------------- 25 | 6 | 25.5-28.0 (26.1) | 9.0-9.5 (9.3) | 16.2-18.5 (16.7) 33 | 2 | 28.5-30.0 (29.3) | 10.2-10.5 (10.4) | 18.0-19.8 (18.9) 36 | 8 | 29.5-34.5 (32.3) | 10.2-11.7 (10.8) | 19.3-23.0 (21.5) 37 | 7 | 31.6-37.5 (34.6) | 11.0-12.5 (11.5) | 21.6-25.0 (23.2) 41 | 3 | 33.0-37.2 (35.2) | 11.6-12.2 (11.9) | 21.4-25.2 (23.2) 43 | 1 | --- | 12.4 | --- 46 | 9 | --- | 13.1-15.7 (14.9) | --- -------+---+------------------+------------------+----------------_stages 34_, _36_, _37_, and _38_.--stage 34, foot paddle-shaped with four toe buds; stage 36, five toe buds; stages 37 and 38, lengthening of toes. in all four stages, spiracle persistent, and pigmentation resembling that of early stages. _stage 39._--metatarsal tubercle present; greatest total length (36.9 mm.) attained. _stage 40._--subarticular tubercles prominent; skin over forelimbs transparent; cloacal tail-piece and spiracle absent; outer tooth-rows degenerating; caudal fins shallower than in preceding stages; distal part of tail nearly straight; size of dark markings on tail decreased; pigment present on hind limb. _stage 43._--forelimbs erupted; larval mouthparts absent; corner of mouth between nostril and eye; transverse bands present on hind limbs; tail greatly reduced (about 8 mm. in length). _stage 44._--sacral hump barely noticeable; tail reduced to a stub; corner of mouth at level of pupil of eye; dorsal surfaces pale olive-green; venter white. changes proceed in a definite pattern during the growth and development of tadpoles. larval teeth are absent in hatchlings; the inner tooth-rows develop first, and the third lower row last. at metamorphosis the third lower row is the first to be lost. the tail increases gradually in length relative to the body. in stage 25 the tail is 52.1 per cent of the total length, and in stage 36, 64.6 per cent. in later stages the tail becomes relatively shorter through resorption. duellman and klaas (1964:320) noted a great size-variation in _triprion_ tadpoles in stage 25. no such variation is apparent in any stage of any of the species of _smilisca_ studied. the growth and development of the other species of _smilisca_ do not differ significantly from that of _s. phaeota_. the tadpoles of _s. sila_ and _sordida_ from streams have relatively longer tails at hatching. for example, in tadpoles of _s. sordida_ the average length of tail is 64.0 per cent of the body-length in stage 25, and in stage 37, 67.0 per cent. _behavior_ the tadpoles of _s. baudini_, _cyanosticta_, _phaeota_, and _puma_ are pelagic inhabitants of shallow ponds. early stages of _s. baudini_ in which external gills are present have been observed to hang vertically with the gills spread out at the surface of the water, a behavior noted by zweifel (1964:206) in tadpoles of _phrynohyas venulosa_, which also develop in warm, standing water having a relatively low oxygen-tension. when disturbed the pelagic tadpoles usually dive and seek shelter amidst vegetation or in mud on the bottom. this behavior was observed in _s. baudini_, _cyanosticta_, and _phaeota_ by day and at night. no tadpoles of _s. puma_ were observed by day; those seen at night were near the surface of small water-filled depressions in a grassy marsh; they responded to light by taking refuge in the dense grass. perhaps tadpoles of this species are negatively phototactic and remain hidden by day. the stream-inhabiting tadpoles of _s. sila_ and _sordida_ live in clear pools in rocky streams, where they were observed to cling by their mouths to rocks in the stream and to seek shelter amidst pebbles or beneath rocks and leaves on the bottom. these tadpoles are not found in shallow riffles. we have not found tadpoles of two species of _smilisca_ in the same body of water and therefore cannot offer observations on ecological relationships in sympatric situations. phylogenetic relationships identifiable hylid remains are known from the miocene to the recent, but these fossils are mostly fragmentary and provide little useful information regarding the phylogenetic relationships of living genera. frogs of the genus _smilisca_ are generalized and show no striking adaptations, either in their structure or in their modes of life history. interspecific relationships in attempting to understand the relationships of the species of _smilisca_ we have emphasized osteological characters. the phylogeny suggested by these characters is supported by other lines of evidence, including external morphology, tadpoles, and breeding calls. our concept of the prototype of the genus _smilisca_ is a moderate-sized hylid having: (1) a well-developed frontoparietal fontanelle, (2) frontoparietal lacking lateral processes, (3) no bony squamosal-maxillary arch, (4) a fully ossified ethmoid, (5) paired subgular vocal sac, (6) moderately webbed fingers and toes, (7) relatively few supernumerary tubercles on the digits, (8) eggs deposited in clumps in ponds, (9) anteroventral mouth in tadpoles bordered by one row of labial papillae, but median part of upper lip bare, (10) tail relatively short and deep in tadpoles, and (11) a breeding call consisting of a series of like notes. two phyletic lines evolved from this prototype. the first of these was the stock that gave rise to the _baudini_ group. the evolutionary changes that took place in this line included increase in size, development of a lateral curvature of the maxillary, and an increased amount of cranial ossification, especially in the dermal roofing bones. this phyletic line retained the larval characters and breeding call of the prototype. the second phyletic line gave rise to the _sordida_ group and diverged from the prototype in the development of an angular maxillary and a breeding call consisting of a primary note followed by secondary notes. the frogs in this phyletic line retained the moderate size of the prototype and did not develop additional dermal bone. our concept of the phylogenetic relationships is shown graphically in figure 17. within the _baudini_ group one stock retained separate nasals and did not develop a bony squamosal-maxillary arch, but broad lateral processes developed on the frontoparietals. the tadpoles remained unchanged from the primitive type. this stock evolved into _s. phaeota_. in the other stock the nasals became fully ossified and a bony squamosal-maxillary arch developed. one branch of this second stock retained tadpoles having only one row of labial papillae and did not develop lateral processes on the frontoparietals; this branch evolved into _s. cyanosticta_. the other branch diverged and gave rise to _s. baudini_ by developing relatively shorter hind legs, large lateral processes on the frontoparietals, and tadpoles having two rows of labial papillae. within the _sordida_ group the cranial features remained unchanged in one line, which gave rise to _s. sila_, whereas in a second line the nasals were reduced, and their long axes shifted with the result that they are not parallel to the maxillaries; the amount of ossification of the ethmoid was reduced, and the tadpoles developed two rows of labial papillae. in this second line one branch retained the pond-breeding habits and gave rise to _s. puma_, whereas a second branch became adapted to stream-breeding and gave rise to _s. sordida_. _baudini_ \ _cyanosticta_ \ / + _phaeota_ _sordida_ \ / / + puma_ / \ \/ _sila_ \ /_____/ \ / \ / \ / | | prototype [illustration: fig. 17. hypothesized phylogenetic relationships of the species of _smilisca_.] certain aspects of this proposed phylogeny warrant further comment. features such as the deposition of additional bone that roofs the skull or that forms lateral projections from the frontoparietals, like those in _s. baudini_ and _phaeota_, are minor alterations of dermal elements and not basic modifications of the architecture of the skull. consequently, we hypothesize the independent development of these dermal changes in _s. baudini_ and _phaeota_. similar kinds of dermal modifications have evolved independently in many diverse groups of frogs. likewise, we propose the parallel development of stream-adapted tadpoles in _s. sordida_ and _sila_; in both cases the tadpoles adapted to changing environmental conditions (see following section on evolutionary history). tadpoles of _s. sordida_ already had two rows of labial papillae before entering the streams; subsequently the tadpoles developed complete rows of papillae, ventral mouths and long tails having low fins. possibly the tadpoles of _s. sila_ had two rows of labial papillae prior to their adapting to stream conditions; in the process of adapting they developed ventral mouths and long tails having low fins. similar modifications in tadpoles have occurred in many diverse groups of middle american hylids, such as _plectrohyla_, _ptychohyla_, the _hyla uranochroa_ group, and the _hyla taeniopus_ group. our lack of concern about coloration is due to the fact that, with the exception of the blue spots on the flanks and posterior surfaces of the thighs in some species, the coloration of _smilisca_, consisting of a pattern of irregular dark marks on a paler dorsum and dark transverse bars on the limbs, is not much different from that of many other neotropical hylids. blue is a structural color, rare among amphibia, which is achieved by the absence of lipophores above the guanophores. thus, the incident light rays at the blue end of the spectrum are reflected by the guanophores without interference by an overlying yellow lipophore screen. according to noble (1931), lipophores are capable of amoeboid movement that permits shifts in their positions, between or beneath the guanophores. we do not know whether this behavior of lipophores is widespread and is effected in response to environmental changes, or whether it is a genetically controlled attribute that is restricted in appearance. if the latter is the case we must assume that the prototype of _smilisca_ possessed such an attribute which was lost in _s. baudini_, _phaeota_, and _puma_. the development of blue spots is not constant in _s. sordida_ and _s. sila_; in _s. cyanosticta_ the spots range in color from blue to pale green. the coloration of the tadpoles is not distinctive, except for the presence of dorsal blotches on the tails of _s. sila_ and _sordida_. however, the similarity in pattern cannot be interpreted as indicating close relationships because nearly identical patterns are present in _hyla legleri_ and some species of _prostherapis_. this disruptive coloration seems to be directly associated with the pebble-bottom, stream-inhabiting tadpoles. in the _baudini_ group, _s. phaeota_ and _cyanosticta_ are allopatric, whereas _s. baudini_ occurs sympatrically with both of those species. the call of _s. baudini_ differs notably from the calls of _s. phaeota_ and _cyanosticta_, which are more nearly alike. although in the phylogenetic scheme proposed here _s. sila_ is considered to be more distantly related to _s. puma_ than is _s. sordida_, the calls of _s. sila_ and _puma_ more closely resemble one another than either resembles that of _s. sordida_. _smilisca sila_ and _puma_ are allopatric, whereas _s. sordida_ is broadly sympatric with both of those species. we assume that in their respective phyletic lines the differentiation of both _s. baudini_ and _sordida_ was the result of genetic changes in geographically isolated populations. subsequently, each species dispersed into areas inhabited by other members of their respective groups. selection for differences in the breeding calls helped to reinforce other differences in the populations and thereby aided in maintaining specificity. evolutionary history with respect to temporal and spatial aspects of evolution in _smilisca_, we have tried to correlate the phylogenetic evidence on _smilisca_ with the geologic data on middle america presented by lloyd (1963), vinson and brineman (1963), guzmán and cserna (1963), maldonado-koerdell (1964), and whitmore and stewart (1965). likewise, we have borne in mind the evidence for, and ideas about, the evolution of the middle american herpetofauna given by dunn (1931b), schmidt (1943), stuart (1950, 1964) duellman (1958, ms), and savage (ms). according to stuart's (1950) historical arrangement of the herpetofauna, _smilisca_ is a member of the autochthonous middle american faunal element, and according to savage's (ms) arrangement the genus belongs to the middle american element, a fauna which was derived from a generalized tropical american unit that was isolated in tropical north america by the inundation of the isthmian link in early tertiary, that developed _in situ_ in tropical north america, and that was restricted to middle america by climatic change in the late cenozoic. savage (ms) relied on the paleogeographic maps of lloyd (1963) to hypothesize the extent and centers of differentiation of the middle american faunal element. according to lloyd's concept, middle america in the miocene consisted of a broad peninsula extending southeastward to about central nicaragua, separated from the panamanian spur of continental south america by shallow seas. a large island, the talamanca range, and remnants of the guanarivas ridge formed an archipelago in the shallow sea. the recent discovery of remains of mammals having definite north american affinities in the miocene of the canal zone (whitmore and stewart, 1965) provides substantial evidence that at least a peninsula was continuous southeastward from nuclear central america to the area of the present canal zone in early mid-miocene time. south america was isolated from central america by the bolivar trough until late mid-pliocene. thus, in the mid-tertiary the broad peninsula of nuclear central america, which consisted of low and moderately uplifted regions having a tropical mesic climate, provided the site for the evolution of _smilisca_. it is not possible to determine when the genus evolved, but to explain the differentiation of the species it is unnecessary to have the ancestral _smilisca_ present prior to the miocene. we view the miocene _smilisca_ as the prototype described in the preceding section, and suppose that it lived in the mesic tropical environment of the eastern part of the central american peninsula (in what is now costa rica and western panamá). two stocks differentiated, probably in middle miocene times; one of these, the ancestral stock of the _baudini_ group, was widespread on the caribbean lowlands from the nicaraguan depression to the bolivar trough, and the other, the ancestral stock of the _sordida_ group, was restricted to the pacific lowlands of the same region. in late miocene time the ancestral stock of the _baudini_ group dispersed northwestward around the deep embayment in the nicaraguan depression into upper central america (in what is now honduras and guatemala) and thence into southern méxico. apparently differentiation took place on each side of the nicaraguan depression; the frogs to the south of the depression evolved into _s. phaeota_, whereas those to the north of the depression represented the stock from which _s. baudini_ and _cyanosticta_ arose. prior to the uplift of the mountains in the late miocene and the pliocene the _baudini-cyanosticta_ stock probably was widespread in northwestern central america. the elevation of the mountains resulted in notable climatic changes, principally the development of sub-humid environments on the pacific lowlands. the frogs living on the pacific lowlands became adapted to sub-humid conditions and developed into _s. baudini_. the stock on the caribbean lowlands remained in mesic environments and evolved into _s. cyanosticta_. possibly in the middle miocene before the talamanca range in costa rica and western panamá was greatly uplifted, the ancestral stock of the _sordida_ group invaded the caribbean lowlands of what is now costa rica. the subsequent elevation of the talamanca range in the pliocene effectively isolated the ancestral stock of _s. sila_ on the pacific lowlands from the _puma-sordida_ stock on the caribbean lowlands. the former was subjected to the sub-humid conditions which developed on the pacific lowlands when the talamanca range was uplifted. it adapted to the sub-humid environment by living along streams and evolving stream-adapted tadpoles. on the caribbean side of the talamanca range the _puma-sordida_ stock inhabited mesic environments. the stock that evolved into _s. puma_ remained in the lowlands as a pond-breeding frog, whereas those frogs living on the slopes of the newly elevated mountains became adapted for their montane existence by developing stream-adapted tadpoles and thus differentiated into _s. sordida_. probably the six species of _smilisca_ were in existence by the end of the pliocene; at that time a continuous land connection existed from central america to south america. the climatic fluctuations in the pleistocene, and the post-wisconsin development of present climatic and vegetational patterns in middle america, brought about the present patterns of distribution of the species. from its place of origin on the caribbean lowlands of lower central america, _s. phaeota_ dispersed northward into nicaragua and southward along the pacific slopes of northwestern south america. perhaps in the late pleistocene or in post-wisconsin time when mesic conditions were more widespread than now, _s. phaeota_ moved onto the pacific lowlands of costa rica. its route could have been through the arenal depression. subsequent aridity restricted its range on the pacific lowlands to the golfo dulce region. climatic fluctuation in northern central america restricted the distribution of _s. cyanosticta_ to mesic habitats on the slopes of the mexican and guatemalan highlands and to certain humid areas on the lowlands. _smilisca baudini_ was well adapted to sub-humid conditions, and the species dispersed northward to the rio grande embayment and to the edge of the sonoran desert and southward into costa rica. in southern méxico and central america the species invaded mesic habitats. consequently, in some areas it is sympatric with _s. cyanosticta_ and _phaeota_. _smilisca puma_ dispersed northward onto the caribbean lowlands of southern nicaragua. its southward movements probably were limited by the ridges of the talamanca range that extend to the caribbean coast in the area of punta cahuita in costa rica. _smilisca sila_ dispersed along the pacific lowlands and slopes of the mountains from eastern costa rica and western panamá through eastern panamá to northern colombia. climatic fluctuation in the pleistocene evidently provided sufficient altitudinal shifts in environments in the talamanca range to permit _s. sordida_ to move onto the pacific slopes. from its upland distribution the species followed streams down to both the caribbean and pacific lowlands, where it is sympatric with _s. puma_ on the caribbean lowlands and _s. sila_ on the pacific lowlands. the evolution of the species-groups of _smilisca_ was effected through isolation by physical barriers in the cenozoic; the differentiation of the species was initiated by further isolation of populations by changes in physiography and climate. present patterns of distribution resulted from pleistocene and post-wisconsin climatic changes. today, sympatric species have different breeding habits and breeding calls which reinforce the differences in morphology. summary and conclusions the genus _smilisca_ is composed of six species of tree frogs; each species is defined on the basis of adult morphology, larval characters, and breeding behavior. keys are provided to aid in the identification of adults and of tadpoles. analysis of the characters and examination of type specimens indicates that several currently-recognized taxa are synonymous, as follows: 1. _hyla beltrani_ taylor, 1942 = _smilisca baudini_. 2. _hyla gabbi_ cope, 1876 = _smilisca sordida_. 3. _hyla manisorum_ taylor, 1954 = _smilisca baudini_. 4. _hyla nigripes_ cope, 1876 = _smilisca sordida_. 5. _hyla wellmanorum_ taylor, 1952 = _smilisca puma_. _smilisca phaeota cyanosticta_ smith, 1953 is elevated to specific rank, and one new species, _smilisca sila_, is named and described. the skeletal system of developmental stages and the adult of _smilisca baudini_ is described, and the skull is compared with that of other members of the genus. the tadpoles are described, compared, and illustrated; the larval development of _smilisca phaeota_ is described. breeding behavior and breeding calls are described and compared. some species of _smilisca_ have breeding choruses. two species, _s. sila_ and _sordida_, breed in streams, whereas the others breed in ponds. the genus is considered to be part of the middle american faunal element; the species are thought to have differentiated in response to ecological diversity and historical opportunities provided by cenozoic changes in physiography and climate. literature cited baird, s. f. 1854. descriptions of new genera and species of north american frogs. proc. acad. nat. sci. philadelphia, 7:59-62. april 27. 1859. reptiles of the boundary. united states and mexican boundary survey. washington, d. c., p. 35, pl. 41. baldauf, r. j. 1959. morphological criteria and their use in showing bufonid phylogeny. jour. morph., 104:527-560. may. barbour, t. 1923. notes on reptiles and amphibians from panama. occas. papers mus. zool. univ. michigan, 129:1-16. january 25. blair, w. f. 1959. call structure and species groups in u. s. treefrogs (_hyla_). southwest. nat., 3:77-89. june 1, 1959. 1962. non-morphological data in anuran classification. syst. zool., 11:72-84. june. 1963. evolutionary relationships of north american toads of the genus bufo: a progress report. evolution, 17:1-16. march. bogert, c. m. 1960. the influence of sound on the behavior of amphibians and reptiles. _in_ lanyon, w. e. and tavolga, w. n. (eds.) animal sounds and communication, pp. 137-320. brattstrom, b. h. 1957. the phylogeny of the salientia based on skeletal morphology. syst. zool., 6:70-74. june. breder, c. m., jr. 1946. amphibians and reptiles of the rio chucunaque drainage, darien, panama, with notes on their life histories and habits. bull. amer. mus. nat. hist., 86:375-436, pls. 42-60. august 26. brocchi, p. 1877. note sur quelques batrachiens hylaeformes recuilles au mexique et au guatemala. bull. soc. philom. paris, ser. 7, 1:122-132. carvalho, a. l. 1954. a preliminary synopsis of the genera of american microhylid frogs. occas. papers mus. zool. univ. michigan, 555:1-19, pl. 1. july 16. cochran, d. m. 1961. type specimens of reptiles and amphibians in the united states national museum. bull. u. s. nat. mus., 220:xv + 291 pp. cope, e. d. 1862. catalogues of the reptiles obtained during the explorations of the parana, paraguay, vermejo and uraguay rivers.... proc. acad. nat. sci. philadelphia, 14, pt. 9:346-359. 1865. third contribution to the herpetology of tropical america. proc. acad. nat. sci. philadelphia, 17:185-198. october. 1871. ninth contribution to the herpetology of tropical america. proc. acad. nat. sci. philadelphia, 23, pt. 2:200-224. copland, s. j. 1957. australian tree frogs of the genus _hyla_. proc. linnean soc. new south wales, 82, pt. 1:9-108. september. duellman, w. e. 1956. the frogs of the hylid genus _phrynohyas_ fitzinger, 1843. misc. publ. mus. zool. univ. michigan, 96:1-47, pls. 1-6. february 21. 1958. a monographic study of the colubrid snake genus _leptodeira_. bull. amer. mus. nat. hist., 114:1-152, pls. 1-31. february 24. 1963a. a review of the middle american tree frogs of the genus ptychohyla. univ. kansas publ. mus. nat. hist., 15:297-349, pls. 11-18. october 18. 1963b. a new species of tree frog, genus _phyllomedusa_, from costa rica. rev. biol. trop., 11(1):1-23. october. 1964. a review of the frogs of the hyla bistincta group. univ. kansas publ. mus. nat. hist., 15:469-491. march 2. 1965. frogs of the _hyla taeniopus_ group. copeia, no. 2:159-168. june 25. duellman, w. e. and klaas, l. t. 1964. the biology of the hylid frog _triprion petasatus_. copeia, no. 2:308-321. june 30. duméril, a. m. c. and bibron, g. 1841. erpétologie générale ou histoire naturelle complète des reptiles, vol. 8, 792 pp. dunn, e. r. 1931a. the amphibians of barro colorado island. occas. papers boston soc. nat. hist., 5:403-421. october 10. 1931b. the herpetological fauna of the americas. copeia, no. 3:106-119. october 30. 1944. herpetology of the bogotá area. rev. acad. colombiana cien. exact., fis. nat., 6:68-81. fouquette, m. j., jr. 1960. isolating mechanisms in three sympatric tree frogs in the canal zone. evolution, 14:484-497. december. funkhouser, anne 1957. a review of the neotropical tree-frogs of the genus _phyllomedusa_. occas. papers nat. hist. mus. stanford univ., 5:1-89. april 1. gadow, h. 1908. through southern mexico. london, witherby and co. xvi + 527 pp. gaige, h. t., hartweg, n. and stuart, l. c. 1937. notes on a collection of amphibians and reptiles from eastern nicaragua. occas. papers mus. zool. univ. michigan, 357:1-18. october 26. goin, c. j. 1961. synopsis of the genera of hylid frogs. ann. carnegie mus., 36:5-18. july 14. gosner, k. l. 1960. a simplified table for staging anuran embryos and larvae with notes on identification. herpetologica, 16:183-190. september 23. griffiths, i. 1959. the phylogeny of _sminthillus limbatus_ and the status of the brachycephalidae (amphibia, salientia). proc. zool. soc. london, 132:457-487, pls. 1-4. guzmán, e. j. and cserna, z. 1963. tectonic history of mexico. amer. assoc. petrol. geol., mem. 2:113-129. hecht, m. k. 1962. a reevaluation of the early history of the frogs. part i. syst. zool., 11:39-44. march. 1963. a reevaluation of the early history of the frogs. part ii. syst. zool., 12:20-35. march. johnson, c. 1959. genetic incompatibility in the call races of _hyla versicolor_ le conte in texas. copeia, no. 4:327-335. december 30. lloyd, j. j. 1963. tectonic history of the south central-american orogen. amer. assoc. petrol. geol., mem. 2:88-100. maldonado-koerdell, m. 1964. geohistory and paleogeography of middle america, _in_ wauchope, r. and west, r. c. (eds.). handbook of middle american indians, vol. 1, univ. texas press, austin, 570 pp. maslin, t. p. 1963. notes on some anuran tadpoles from yucatán, méxico. herpetologica, 19:122-128. july 3. mittleman, m. b. and list, j. c. 1963. the generic differentiation of the swamp treefrogs. copeia, no. 2:80-83. may 29. noble, g. k. 1931. the biology of the amphibia. mcgraw hill, new york, 577 pp. orton, g. l. 1957. the bearing of larval evolution on some problems in frog classification. syst. zool., 6:79-86. june. peters, w. 1863. mittheilungen uber neue batrachier. monats. konigl. akad. wiss. berlin, pp. 445-471. 1873. uber eine neue schildkrötenart, _cinosternon effeldtii_ und einige andere neue oder weniger bekannte amphibien. monats. konigl. akad. wiss. berlin, pp. 603-618, pl. 5. october 16. rivero, j. a. 1961. salientia of venezuela. bull. comp. zool., 126:1-207. november. savage, j. m. and carvalho, a. l. 1953. the family position of neotropical frogs currently referred to the genus _pseudis_. zoologica, 38:193-200. schmidt, k. p. 1941. the amphibians and reptiles of british honduras. zool. ser. field mus. nat. hist., 22:475-510. december 30. 1943. corollary and commentary for "climate and evolution." amer. midl. nat., 30:241-253. july. schmidt, o. 1857. diagnosen neuer frösche des zoologischen cabinets zu krakau. sitzungb. konigl. akad. wiss. math.-natur. cl., 24(1):10-15. march. 1858. deliciae herpetogicae musei zoologici cracoviensis. denkschr. k. k. akad. wiss. math.-natur. cl., 14(2):237-258, pls. 1-3. smith, h. m. 1953. a new subspecies of the treefrog _hyla phaeota_ cope of central america. herpetologica, 8:150-152. january 30. smith, h. m. and taylor, e. h. 1950. type localities of mexican reptiles and amphibians. univ. kansas sci. bull., 33:313-380. march 20. starrett, p. 1960. a redefinition of the genus _smilisca_. copeia, no. 4:300-304. december 30. stebbins, r. c. and hendrickson, j. r. 1959. field studies of amphibians in colombia, south america. univ. california publ. zool., 56:497-540. february 17. stokely, p. s. and list, j. c. 1954. the progress of ossification in the skull of the cricketfrog _pseudacris nigrita triseriata_. copeia, no. 3:211-217. july 29. stuart, l. c. 1935. a contribution to a knowledge of the herpetology of a portion of the savanna region of central petén, guatemala. misc. publ. mus. zool. univ. michigan, 29:1-56, pls. 1-4, 1 map. october 1. 1948. the amphibians and reptiles of alta verapaz, guatemala. misc. publ. mus. zool. univ. michigan, 69:1-109. june 12. 1950. a geographic study of the herpetofauna of alta verapaz, guatemala. contr. lab. vert. biol., 45:1-77, pls. 1-9, 1 map. may. 1954. herpetofauna of the southeastern highlands of guatemala. contr. lab. vert. biol., 68:1-65, pls. 1-4. november. 1958. a study of the herpetofauna of the uaxactun-tikal area of northern el petén, guatemala. contr. lab. vert. biol., 75:1-30. june. 1961. some observations on the natural history of tadpoles of _rhinophrynus dorsalis_ duméril and bibron. herpetologica, 17:73-79. july 11. 1964. fauna of middle america, _in_ wauchope, r. and west, r. c. (eds.). handbook of middle american indians, vol. 1, univ. texas press, austin, 570 pp. taylor, e. h. 1942. new caudata and salientia from méxico. univ. kansas sci. bull., 28:295-323. november 15. 1952. the frogs and toads of costa rica. univ. kansas sci. bull., 35:577-942. july 1. 1954. additions to the known herpetological fauna of costa rica with comments on other species. no. i. univ. kansas sci. bull., 36:597-639. june 1. taylor, e. h. and smith, h. m. 1945. summary of the collections of amphibians made in méxico under the walter rathbone bacon traveling scholarship. proc. u. s. natl. mus., 95:521-613, pls. 18-32. june 30. tihen, j. a. 1962. osteological observations on new world bufo. amer. midl. nat., 67:157-183. january. 1965. evolutionary trends in frogs. amer. zoologist, 5:309-318. vinson, g. l. and brineman, j. h. 1963. nuclear central america, hub of antillean transverse belt. amer. assoc. petrol. geol., mem. 2:101-112. whitmore, f. c., jr. and stewart, r. h. 1965. miocene mammals and central american seaways. science, 148:180-185. april 9. zweifel, r. g. 1956. two pelobatid frogs from the tertiary of north america and their relationships to fossil and recent forms. amer. mus. novitates, 1762:1-45. april 6. 1958. results of the archbold expeditions. no. 78. frogs of the papuan hylid genus _nyctimystes_. amer. mus. novitates, 1896:1-51. july 22. 1964. life history of _phrynohyas venulosa_ (salientia: hylidae) in panamá. copeia, no. 1:201-208. march 26. _transmitted march 14, 1966._ [] 31-3430 university of kansas publications museum of natural history institutional libraries interested in publications exchange may obtain this series by addressing the exchange librarian, university of kansas library, lawrence, kansas. copies for individuals, persons working in a particular field of study, may be obtained by addressing instead the museum of natural history, university of kansas, lawrence, kansas. when individuals request copies from the museum, 25 cents should be included, for each 100 pages or part thereof, for the purpose of defraying the costs of wrapping and mailing. for certain longer papers an additional amount, indicated below, toward some of the costs of production, is to be included. * an asterisk designates those numbers of which the museum's supply is exhausted. vol. 1. nos. 1-26 and index. pp. 1-638, 1946-1950. *vol. 2. (complete) mammals of washington. by walter w. dalquest. pp. 1-444, 140 figures in text. april 9, 1948. vol. 3. *1. the avifauna of micronesia, its origin, evolution, and distribution. by rollin h. baker. pp. 1-359, 16 figures in text. june 12, 1951. *2. a quantitative study of the nocturnal migration of birds. by george h. lowery, jr. pp. 361-472, 47 figures in text. june 29, 1951. 3. phylogeny of the waxwings and allied birds. by m. dale arvey. pp. 473-530, 49 figures in text, 13 tables. october 10, 1951. *4. birds from the state of veracruz, mexico. by george h. lowery, jr. and walter w. dalquest. pp. 531-649, 7 figures in text, 2 tables. october 10, 1951. index. pp. 651-681. *vol. 4. (complete) american weasels. by e. raymond hall. pp. 1-466, 41 plates, 31 figures in text. december 27, 1951. vol. 5. nos. 1-37 and index. pp. 1-676, 1951-1953. *vol. 6. (complete) mammals of utah, _taxonomy and distribution_. by stephen d. durrant. pp. 1-549, 91 figures in text, 30 tables. august 10, 1952. vol. 7. nos. 1-15 and index. pp. 1-651, 1952-1955. vol. 8. nos. 1-10 and index. pp. 1-675, 1954-1956. vol. 9. nos. 1-23 and index. pp. 1-690, 1955-1960. vol. 10. nos. 1-10 and index. pp. 1-626, 1956-1960. vol. 11. nos. 1-10 and index. pp. 1-703, 1958-1960. vol. 12. *1. functional morphology of three bats: eumops, myotis, macrotus. by terry a. vaughan. pp. 1-153, pls. 1-4, 24 figures in text. july 8, 1959. *2. the ancestry of modern amphibia: a review of the evidence. by theodore h. eaton, jr. pp. 155-180, 10 figures in text. july 10, 1959. 3. the baculum in microtine rodents. by sidney anderson. pp. 181-216, 49 figures in text. february 19, 1960. *4. a new order of fishlike amphibia from the pennsylvanian of kansas. by theodore h. eaton, jr., and peggy lou stewart. pp. 217-240, 12 figures in text. may 2, 1960. 5. natural history of the bell vireo, vireo bellii audubon. by jon c. barlow. pp. 241-296, 6 figures in text. march 7, 1962. 6. two new pelycosaurs from the lower permian of oklahoma. by richard c. fox. pp. 297-307, 6 figures in text. may 21, 1962. 7. vertebrates from the barrier island of tamaulipas, méxico. by robert k. selander, richard f. johnston, b. j. wilks, and gerald g. raun. pp. 309-345, pls. 5-8. june 18, 1962. 8. teeth of edestid sharks. by theodore h. eaton, jr. pp. 347-362, 10 figures in text. october 1, 1962. 9. variation in the muscles and nerves of the leg in two genera of grouse (tympanuchus and pedioecetes). by e. bruce holmes. pp. 363-474, 20 figures in text. october 25, 1963. $1.00. 10. a new genus of pennsylvanian fish (crossopterygii, coelacanthiformes) from kansas. by joan echols. pp. 475-501, 7 figures in text. october 25, 1963. 11. observations on the mississippi kite in southwestern kansas. by henry s. fitch. pp. 503-519. october 25, 1963. 12. jaw musculature of the mourning and white-winged doves. by robert l. merz. pp. 521-551, 22 figures in text. october 25, 1963. 13. thoracic and coracoid arteries in two families of birds, columbidae and hirundinidae. by marion anne jenkinson. pp. 553-573, 7 figures in text. march 2, 1964. 14. the breeding birds of kansas. by richard f. johnston. pp. 575-655, 10 figures in text. may 18, 1964. 75 cents. 15. the adductor muscles of the jaw in some primitive reptiles. by richard c. fox. pp. 657-680, 11 figures in text. may 18, 1964. index. pp. 681-694. vol. 13. 1. five natural hybrid combinations in minnows (cyprinidae). by frank b. cross and w. l. minckley. pp. 1-18. june 1, 1960. 2. a distributional study of the amphibians of the isthmus of tehuantepec, méxico. by william e. duellman. pp. 19-72, pls. 1-8, 3 figures in text. august 16, 1960. 50 cents. 3. a new subspecies of the slider turtle (pseudemys scripta) from coahuila, méxico. by john m. legler. pp. 73-84, pls. 9-12, 3 figures in text. august 16, 1960. *4. autecology of the copperhead. by henry s. fitch. pp. 85-288, pls. 13-20, 26 figures in text. november 30, 1960. 5. occurrence of the garter snake, thamnophis sirtalis, in the great plains and rocky mountains. by henry s. fitch and t. paul maslin. pp. 289-308. 4 figures in text. february 10, 1961. 6. fishes of the wakarusa river in kansas. by james e. deacon and artie l. metcalf. pp. 309-322, 1 figure in text. february 10, 1961. 7. geographic variation in the north american cyprinid fish, hybopsis gracilis. by leonard j. olund and frank b. cross. pp. 323-348, pls. 21-24, 2 figures in text. february 10, 1961. 8. descriptions of two species of frogs, genus ptychohyla; studies of american hylid frogs, v. by william e. duellman. pp. 349-357, pl. 25, 2 figures in text. april 27, 1961. 9. fish populations, following a drought, in the neosho and marais des cygnes rivers of kansas. by james everett deacon. pp. 359-427, pls. 26-30, 3 figures in text. august 11, 1961. 75 cents. 10. north american recent soft-shelled turtles (family trionychidae). by robert g. webb. pp. 429-611, pls. 31-54, 24 figures in text. february 16, 1962. $2.00. index. pp. 613-624. vol. 14. 1. neotropical bats from western méxico. by sydney anderson. pp. 1-8. october 24, 1960. 2. geographical variation in the harvest mouse, reithrodontomys megalotis, on the central great plains and in adjacent regions. by j. knox jones, jr., and b. mursaloglu. pp. 9-27, 1 figure in text. july 24, 1961. 3. mammals of mesa verde national park, colorado. by sydney anderson. pp. 29-67, pls. 1 and 2, 3 figures in text. july 24, 1961. 4. a new subspecies of the black myotis (bat) from eastern méxico. by e. raymond hall and ticul alvarez. pp. 69-72, 1 figure in text. december 29, 1961. 5. north american yellow bats, "dasypterus," and a list of the named kinds of the genus lasiurus gray. by e. raymond hall and j. knox jones, jr. pp. 73-98, 4 figures in text. december 29, 1961. 6. natural history of the brush mouse (peromyscus boylii) in kansas with description of a new subspecies. by charles a. long. pp. 99-110, 1 figure in text. december 29, 1961. 7. taxonomic status of some mice of the peromyscus boylii group in eastern méxico, with description of a new subspecies. by ticul alvarez. pp. 111-120, 1 figure in text. december 29, 1961. 8. a new subspecies of ground squirrel (spermophilus spilosoma) from tamaulipas, mexico. by ticul alvarez. pp. 121-124. march 7, 1962. 9. taxonomic status of the free-tailed bat, tadarida yucatanica miller. by j. knox jones, jr., and ticul alvarez. pp. 125-133, 1 figure in text. march 7, 1962. 10. a new doglike carnivore, genus cynarctus, from the clarendonian, pliocene, of texas. by e. raymond hall and walter w. dalquest. pp. 135-138. 2 figures in text. april 30, 1962. 11. a new subspecies of wood rat (neotoma) from northeastern mexico. by ticul alvarez. pp. 139-143. april 30, 1962. 12. noteworthy mammals from sinaloa, mexico. by j. knox jones, jr., ticul alvarez, and m. raymond lee. pp. 145-159, 1 figure in text. may 18, 1962. 13. a new bat (myotis) from mexico. by e. raymond hall. pp. 161-164, 1 figure in text. may 21, 1962. *14. the mammals of veracruz. by e. raymond hall and walter w. dalquest. pp. 165-362, 2 figures in text. may 20, 1963. $2.00. 15. the recent mammals of tamaulipas, méxico. by ticul alvarez. pp. 363-473, 5 figures in text. may 20, 1963. $1.00. 16. a new subspecies of the fruit-eating bat, sturnira ludovici, from western mexico. by j. knox jones, jr., and gary l. phillips. pp. 475-481, 1 figure in text. march 2, 1964. 17. records of the fossil mammal sinclairella, family apatemyidae, from the chadronian and orellan. by william a. clemens, jr. pp. 483-491, 2 figures in text. march 2, 1964. 18. the mammals of wyoming. by charles a. long. pp. 493-758, 82 figures in text. july 6, 1965. $3.00. index. pp. 759-784. vol. 15. 1. the amphibians and reptiles of michoacán, méxico. by william e. duellman. pp. 1-148, pls. 1-6, 11 figures in text. december 20, 1961. $1.50. 2. some reptiles and amphibians from korea. by robert g. webb, j. knox jones, jr., and george w. byers. pp. 149-173. january 31, 1962. 3. a new species of frog (genus tomodactylus) from western méxico. by robert g. webb. pp. 175-181, 1 figure in text. march 7, 1962. 4. type specimens of amphibians and reptiles in the museum of natural history, the university of kansas. by william e. duellman and barbara berg. pp. 183-204. october 26, 1962. 5. amphibians and reptiles of the rainforests of southern el petén, guatemala. by william e. duellman. pp. 205-249, pls. 7-10, 6 figures in text. october 4, 1963. 6. a revision of snakes of the genus conophis (family colubridae, from middle america). by john wellman. pp. 251-295, 9 figures in text. october 4, 1963. 7. a review of the middle american tree frogs of the genus ptychohyla. by william e. duellman. pp. 297-349, pls. 11-18, 7 figures in text. october 18, 1963. 50 cents. 8. natural history of the racer coluber constrictor. by henry s. fitch. pp. 351-468, pls. 19-22, 20 figures in text. december 30, 1963. $1.00. 9. a review of the frogs of the hyla bistincta group. by william e. duellman. pp. 469-491, 4 figures in text. march 2, 1964. 10. an ecological study of the garter snake, thamnophis sirtalis. by henry s. fitch. pp. 493-564, pls. 23-25, 14 figures in text. may 17, 1965. 11. breeding cycle in the ground skink, lygosoma laterale. by henry s. fitch and harry w. greene. pp. 565-575, 3 figures in text. may 17, 1965. 12. amphibians and reptiles from the yucatan peninsula, méxico. by william e. duellman. pp. 577-614, 1 figure in text. june 22, 1965. 13. a new species of turtle, genus kinosternon, from central america. by john m. legler. pp. 615-625, pls. 26-28, 2 figures in text. june 20, 1965. 14. a biogeographic account of the herpetofauna of michoacán, méxico. by william e. duellman. pp. 627-709, pls. 29-36, 5 figures in text. december 30, 1965. 15. amphibians and reptiles of mesa verde national park, colorado. by charles l. douglas. pp. 711-744, pls. 37 and 38, 6 figures in text. march 7, 1966. index in preparation. vol. 16. 1. distribution and taxonomy of mammals of nebraska. by j. knox jones, jr. pp. 1-356, plates 1-4, 82 figures in text. october 1, 1964. $3.50. 2. synopsis of the lagomorphs and rodents of korea. by j. knox jones, jr., and david h. johnson. pp. 357-407. february 12, 1965. 3. mammals from isla cozumel, mexico, with description of a new species of harvest mouse. by j. knox jones, jr. and timothy e. lawlor. pp. 409-419, 1 figure in text. april 13, 1965. 4. the yucatan deer mouse, peromyscus yucatanicus. by timothy e. lawlor. pp. 421-438, 2 figures in text. july 20, 1965. 5. bats from gautemala. by j. knox jones, jr. pp. 439-472. april 18, 1966. more numbers will appear in volume 16. vol. 17. 1. localities of fossil vertebrates obtained from the niobrara formation (cretaceous) of kansas. by david bardack. pp. 1-14. january 22, 1965. 2. chorda tympani branch of the facial nerve in the middle ear of tetrapods. by richard c. fox. pp. 15-21. june 22, 1965. 3. fishes of the kansas river system in relation to zoogeography of the great plains. by artie l. metcalf. pp. 23-189, 4 figures in text, 51 maps. march 24, 1966. 4. factors affecting growth and production of channel catfish, ictalurus punctatus. by bill a. simco and frank b. cross. pp. 191-256, 13 figures in text. june 6, 1966. 5. a new species of fringe-limbed tree frog, genus hyla, from darién, panamá. by william e. duellman. pp. 257-262, 1 figure in text. june 17, 1966. 6. taxonomic notes on some mexican and central american hylid frogs. by william e. duellman. pp. 263-279. june 17, 1966. 7. neotropical hylid frogs, genus smilisca. by william e. duellman and linda trueb. pp. 281-375, pls. 1-12, 17 figures in text. july 14, 1966. more numbers will appear in volume 17. transcriber's note with the exception of the corrections listed below and several minor corrections not listed, the text presented is that which appeared in the original printed version. the list of kansas university publications has been compiled at the end of the article. typographical corrections page correction ==== ================== 287 cleared => cleaned 292 data based of => data based on 298 cnmh => cnhm 299 acahuitzotla => acahuizotla 304 cyanostica => cyanosticta 305 quatemala => guatemala 307 cyanostica => cyanosticta 313 matagalapa => matagalpa 322 carribean => caribbean 323 centralia => centrali 336 proportionaely => proportionately 346 noticably => noticeably 362 fouquett => fouquette 372 carvaljo => carvalho 375 dumeril => duméril ii trutles => turtles 