Bulletin Olf THE Illinois State Laboratory OF Natural History Urbana, Illinois, U. S. A. STEPHEN A. FORBES, Ph.D., L,L.D., Director Vol. IX. August, 1913 Article XII. I SOME NEW ILEINOIS ENCHYTR^ID^ BY Prank Smith, A.M., and Paul S. Wklch, Ph.D. ERRATA AND ADDENDA Page 54, lines 3 and 2 from bottom, and elsewhere in Article III. for Cassia chainaechrista read Cassia chamaccrista. Page 62, between lines 4 and 5 from bottom of table insert Erigeron annuus. Page loi, table, after Croloii glandulosus read var. septentrionalis: and for Eijuisettim laeinyatum read Eqnisetum bycmale var. inlermedium. Page 131. line 3, for coerulea read caerulca. Page 138, last line, for Zi::a read Zizia. Page 141, line 21 from bottom, dele Diodia teres. Page l6g. between lines 3 and 4, insert as follows : Erigeron annuus (L.) Pers. An interstitial in the liunch-grass association in the Hanover area. Page 177, line 5, for casti^'ard read iveslzvard. Page 209. line 3 from bottom, for copalina read copallina. Page 210, line 13 from bottom, for Diospyrus read Diospyro'i. Page 211, line 5, for Foresteria read Foresticra. Page 256, line 3 of table, for Dr. H. M. Pepoon read H. 5". Pepoon. Page 278, line 16, the fifth word should be in Roman type. Page 286, line 6 (second column), page 295, list of secondary species (second column), and page 353. line 8 from bottom, for hiematis or hiemale read hye- mnte. Page 313, line 4 from bottom (first column), for pedicularis read pcdicularia. Page 315, line 10, second column, for Apoeynum read Apocynum. Page 323, line 3 from bottom, for Cyperus read Scirpus. Page 330, line 14, for virginianum read inrginicum. Page 336. lines 3 and 2 from bottom, for virginicum read virginianum. Page 337. line 2 from bottom, for philadelphicum read philadelphicns. Page 339» in first list of invading species, for Rhus hiria read Rhus typhiua. Page 351. line 4 from bottom, for .verophtic read .verophytic. Page 355, above line 6 from bottom, insert Scirpus heterochaetus Chase. Page 356, line 14 from bottom, for Symlocarpus read Symplocarpus. Page 360, line 14, for Pirus read Pyrus. Page 362, after line 7, insert Acer saccharinum L. Page 363, line 2 from bottom, for quadiflorum read quadriflorum. Page 365, line 14, for Ihapus read thapsus. Page 369, last line, for Tanecelum read Tanaceium. Page 417, line i. dele the. Page 497, line 9 from bottom, for neglible read negligible, and in foot-note, for Auslall read Anstalt. Page 498, line 4 from bottom, for Lockport read Chillicothe. Page 500, line 13 from bottom, after up insert in. Page 5or, line 2 from bottom, for dissolving read dissolved. Page 504, line 23, for gryina read gyrina : line 17, for dentata read knickerbockeri. Page 506, line 11, for vernata read ternala. Page 507, line 3 from bottom, for Macon read tt'asoi. Page 513, line 19, for Nepa read Zaitha; line 18, and page 517, line 13 from bot- tom, page 520, line 12 from bottom, and page 532, line 4, read naid or naids for naiid or naiids. Page 517, line 6 from bottom, for pondiveed read pickcrel-iveed. Page 519. for first sentence of last paragraph read as follows: We have no exactly comparable chemical data for July; but analyses for August give percentages of saturation for Morris and Marseilles as follows : 20.4 per cent, at Morris on the nth and 11 per cent, at Marseilles on the 12th; 16.35 P<^r cent, at Morris on the 22d and 23d and 7.4 per cent, at Marseilles on the 24th and 25th. Page 521, line 6 from bottom, and page 529, line g, for chrysoleucas read cryso- leticas. Page 525, line 22, and page 536, lines 21 and 24, for Ekmann read Ekmnn. Page 532, line i, for Ancyclus read Ancylus. Page 551, line 7, for 00 read 572. Page 615, second line above foot-note, for 106 read 94. Page 616, line I, for the second Biindeln read Bilndel; line 2, for Biindeln read Biindcls; line 3, for ausscrn read ausscren; line 6, for sweierlie read sweierlei. Page 629, line 12, for kciii read kcincn. Page 634, line 9, for untcrnommcn read untcninntincnen; and in line 14 from bottom, after ;;/$ insert is fig. Plate III, Fig. i, after the word mixed in legend insert consocies of the. Plate IX, Fig. 2, dele the legend and read instead ; Root-system of Tephrosia virgiiiiana, exposed by blowing of the sand. Plate X, Fig. 2, dele the legend and read instead ; A blowout almost stabilized by bunch-grasses, especially Leptoloma cognatum. Plate XXXIX. for Calainogrostis read Calamagrostis. Plate LIV. exchange places of cuts, but not the legends. Plate LXXXV, for 7 read ye. Article XII. — Sonic Nnv Illinois Bnchytrccidcc* By Frank Smith and Paul S. Welch. Nothing has been published on the Bnchytrccidce of the Mississippi Valley with the exception of a brief description of Fridericia agilis by the senior author nearly twenty years ago ('95, p. 288) and the insufficient account of the problematic forms from Lake Superior by S. I. Smith and Verrill ('71). In this paper we first describe two new species of Fridericia from Urbana, Illinois. The major part of these descriptions was con- tained in a thesis offered by the junior author in 191 1 in partial ful- filment of the requirements for the master's degree from the Uni- versity of Illinois. We next extend the description of Fiidericia agilis Smith and add a numl>er of rather important characters not previously given. Finally, we describe a new species of Marionina from Urbana, Illinois, which is the first member of this genus re- ported from the United States. Descriptions of seven other new species of Bnchytrccidcc from the Mississippi Valley will soon appear in a paper by the junior author which is about to go to press. The drawings for figures 22 to 28 inclusive were made by Mr. S. Fred Prince, illustrator for the Illinois State Laboratory of Natural History. All other drawings were made by the junior author. FRIDERICIA Michaelsen About ninety species are assigned to this genus at the present time, making it the largest genus of the family. Of this large as- semblage of species only fourteen have been recorded from North America, and only one from Illinois. Four species from the vicinity of Philadelphia, Pennsylvania, one species from Havana, Illinois, and seven species from California constitute the list for the United States. So far as is known the members of this genus are all ter- restrial in habit, although they usually occur under somewhat moist conditions. The genus was established by Michaelsen in 1889, when he dis- covered that certain species then assigned to Bnchytrcciis, Neoenchy- trccHS, Mcscnchyfrccus and Archicnchytrcciis had definite characters in common which were of sufficient importance to justify the estab- lishing of a new genus. ?Ie defines ('00, p. 106) the genus as fol- lows : Contributions from the Zoological Laboratory, University of Illinois, un- der the direction of Henry B. Ward, No. 22. (il(5 "Borsten in 4 Biindeln, gerade, zu 2 im Biindeln und dann gleich lang oder zu mehreren und dann die inneren des Biindeln mehr oder weniger regelmassig paarweise und Stufenweise als die aussern. Riickenporen mit Verschlusszellen meist vom 7., selten vom 6. Segni. an vorhanden. Kopfporus meist klein, dorsal zwischen Kopflappen und I. Segm. Lyniphkorper von zweierlie Gestalt. Peptonephridien stets vorhanden. Der Oesophagus geht alhnahHch in den Mitteldarm iiber. Das Riickengefass entspringt meist postclitellial. Blut farblos. Nephridien meist mit grossem Anteseptale, in dem der Flimmerkanal schon Windungen beschreibt. Samenleiter lang. Samentaschen meist mit dem Darm kommunizierend, einfach oder mit Divertikeln." Eisen ('05, p. 106) extended the definition by adding the follow- ing characters: "Penial bulb without interior muscular strands.". . . "The intestine in the vicinity of the clitellum contains specialized chy- lus cells." It remains for future investigation to show whether or not these characters are worthy of generic rank, since in the majority of the species of the genus the penial bulb and the chylus cells have not been studied. PRIDERICIA FIRMA n. Sp. Definition.—Length, 24-33 "T"- Somites, 62-67. Color, whit- I ish yellow. Prostomium blunt. First dorsal pore in VII. Setre, [' 4-7 per bundle in anterior part ; 4-5 in middle part ; 2 in posterior part. Clitellum on XII-XIII. Lymiphocytes elliptical, abundant. Brain 14 longer than wide ; anterior margin concave ; posterior mar- gin convex. Peptonephridia branching dendritically. Anteseptal part of nephridium slightly smaller than postseptal part; efferent duct arises from mid-ventral surface of latter, near septum. Spermiducal funnel 1.3 to 1.5 times longer than diameter; duct long, slender, and much contorted, and confined to XII. Spemiathecte each with pear- shaped ampulla communicating with digestive tract in V; each am- pulla with 3-4 sessile, unequal, lobular diverticula; duct longer than ampulla, slender, and without glands at ectal opening. I The characters of the penial bulb and the chylus cells are not in- cluded in the above definition but will be discussed in another part of the paper (pp. 618, 619). Described from ten sexually mature specimens. Type and para- types in the collection of the junior author and paratypes in the col- lection of the senior author. The specimens which are the basis of this description were found with many others near Urbana, Illinois, in November, 1910, in moist soil under decaying leaves in undisturbed forest land. They were abundant and most of them were sexually mature. (•)I7 Afijnitics.—This species is easily distinguished from the other species of the genus. It resembles F. agilis Smith ('95, p. 288), F. longa Moore ("95, p. 341), and F. ratzcli Eisen ('72, p. 123) more closely than it does other memljers of the genus, but it differs from F. agilis in the number of set?e per bundle, the number of diverticula on each spermatheca, and in the characters of the ne- phridia. It differs from F. longa in the characters of the setae, pep- tonephridia, and spermathecje ; and from F. rat::cli in size, and in the characters of the brain and the spermathecee. EXTERNAL CHARACTERS The body is long, slender, and cylindrical. The average length is about 30 mm., the extremes being 24 and 33 mm. The diaineter is greatest at the clitellum (0.6 mm.), gradually decreasing both caudad and cephalad. The intersegmental grooves are rather ob- scure except in the region just posterior to the prostomium, where a few are quite distinct. No secondary annulations are present. The number of somites varies from 62 to 67. The prostomium (PI. XCVIII, Fig. I ) is blunt and rounded. The seta Ixindles in the an- terior part of the body each contain 6-7 setje; those in the posterior part 2-4 each. The proximal ends (PI. XCVIII, Pig. 2), especially of the more fully developed setce, are strongly bent. INTERNAL CHARACTERS Body Wall.—The cuticula, though comparatively thin, is tough and firm, and offers the chief difficulty in the dissection of alcoholic specimens ; but when once broken, it can be easily stripped from the body. It has a luster, in both surface and sectional views, which gives it a glistening appearance. It is of about the same thickness throughout the length of the body. It is reflected into the mouth and lines the digestive tract as far as the openings of the peptonephridia, being continued into the latter for a short distance. It lines the canal of the head pore for most of its length, but is merely perforated at the dorsal pores. The comparative thinness of the cuticula is probably attributable to the fact that this species lives in moist earth. According to Vejdovsky ("79, p. 11) such forms as Aiiachcrta, which live in dry surroundings, have a \'ery thick cuticula, while in those living in water or in very moist earth it is thin and delicate. It was noticed that the mortality in this species was very high when the earth surrounding the specimens approached dryness, or when they were exposed for a short time to the open air. 618 Tile clitelluni is on XII and XIII and is distinctly developed. The cells (PI. XCVIII, Fig. 3) are all of the same kind. The first dorsal pore is in VII, and each pore has a mid-dorsal position in the middle of its somite lengthwise. Brain,.—The brain (PI. XCVIII, Pig. 4) is in the anterior part of II but projects slightly into I. The length is about one and one fourth times the width. The anterior margin is moderately concave, while the posterior margin is strongly convex. The lateral margins are nearly parallel. Two pairs of supporting strands extend from the lateral margins to the body wall. Pcptoncphridia (Salk'ary Glands).—A pair of these organs open into the ventral side of the digestive tract in the anterior part of IV, close to the septum. They extend caudad through IV and V and a short distance into VI. Each gives off a number of branches at irregular intervals throughout its length. Chylns Cells.—The chylus cell region is that part of the intestine included in XIII-5<3 XVIII. Here the intestine is lined with the layer of ental epithelial cells, which are ciliated and approximately rectangular in section, and each of these cells contains a large spheri- cal nucleus. The chylus cells lie deeper in the intestinal wallj and open between the cells of the ental layer into the lumen of the intes- tine. They are long, rather narrow, and broader at the bases than at the apices. The intracellular canal (PI. XCVIII, Figs. 6, 7) is ciliated and nearly straight in the apical part, but elsewhere it is non- ciliated and somewhat tortuous, especially in the broad basal part. It is surrounded by a definite layer of cytoplasm of uniform thickness, which is more hyaline and stains more intensely than the finely granu- lar and deeply staining adjacent part. The two kinds of cytoplasm are sharply differentiated. The elliptical nucleus lies in the base of the cell in the angle formed at the chief bend of the canal. The cell walls are indistinct, and it is somewhat difficult to determine the exact line of demarcation between adjacent cells. However, suffi- cient details of structure can be made out to prove that these cells are definite units and that each canal is confined entirely within the limits of a single cell. There is some variation in the form of the chylus cells, since near septa and sometimes in the lateral walls of the intestine they are rather short, while between septa they may be rather long and slender. However, in all cases the general structure and the arrangement of parts are uniform. Interstitial cells are ab- sent. Certain considerations relating to the chylus cells will be taken up in another part of the paper (p. 626). Ncphridia.—The first pair of nephridia are related to VI/VII. The anteseptal part (PI. XCVIII, Fig. 5) of each nephridium is 019 slightly smaller than the postseptal part. The efferent duct arises from the ventral surface of the latter close to the septum. The lumen is more or less contorted in both anteseptal and postseptal parts. Spermarics.—The spermaries are in the usual position in XI. In most of the preparations the germ cells show various stages of mitotic division. In sexually mature specimens a large mass of developing sperm cells nearly fills the somite, often pushing the septa into con- tact with septa IX/X and XII/XIII. Sperm Ducts.—The spermiducal fvmnel (PI. XCVIII, Fig. 8) is nearly cylindrical, and the length is approximately one and one halt times the diameter. It is situated in the posterior part of XI with the base in close proximity to the ventral part of XI/XII, and the long axis is directed obliquely dorsad. The anterior end has a well- jj differentiated reflected collar, distinctly set off from the body of the [: funnel by a constriction. The sperm duct is rather long, contorted, '| and confined to XII. I Penial Bulb.—The penial bulbs (PI. XOIX, Figs. 9, 10) are in ) close relation to the penial pores* and each is involved in, a very mate- rial modification of the body wall. At each penial pore there is a deep j invagination and an interruption of the muscle layers. The character \. of the hypodennis is abruptly altered, but the cuticula, slightly re- || duced in thickness, is reflected into the invagination as a lining. The \ body of the bulb is composed of two kinds of cells: ( i) large, glandu- lar cells which occupy all of the dorsal and peripheral parts of the bulb and have large spherical nuclei; and (2) long, narrow, columnar cells which are situated in the interior of the bulb and are arranged radially around the penial lumen* for its entire length. Cells of the second type contain small ovoid nuclei at their inner ends and stain very lightly. They inerge gradually into ordinary hypodermal cells at the ventral side of the bulb, which, in turn, give place to the clitellar cells present in the mid-ventral region. The bulb is covered by a well-developed musculature which is a continuation of the circular muscle layer of the body wall. The bulb is a typical lumbricillid bulb as defined by Eisen ('05, p. 8). He claims that this organ has taxonomic importance and can be used in the characterization of species, genera, and subfamilies. The structure of the bulb agrees also with his diagnosis of the genus *We shall not undertake a discussion of the homologies of the penial bulb, but for the purposes of this paper shall designate the ectal opening of its lumen as the penial pore (PI. XCIX, Fig. 10, pen. po.) instead of the spermi- ducal pore, and the lumen itself (Fig 10, pen. hiin.) as the penial lumen. This latter is the same as Eisen's "extension of the sperm duct" and "elongation of the sperm duct" ('05, pp. 108 and 8). 020 Fridericia and the subfamily Liniibvicillincc, based on the character of the penial bulb. The discussion of the importance of this organ in classification is taken up in detail in. another ])ai>er to be pub- lished in the near future. Spcnuathcccr.—Each spermatheca (PI. XCIX, Fig. ii) consists of three distinct parts; an ampulla, diverticula, and the duct. The am- pulla, the largest part of the spermatheca, is a pear-shaped organ, of which the smaller end is united with the digestive tract in the pos- terior part of V, and the larger end is connected with the body wall by means of the duct. The cavity within conforms somewhat to the external shape of the organ, being rather spacious in the region of the diverticula, and gradually narrowing to the point where it com- municates with the lumen of the digestive tract. The communica- tions of the two spermatheca? with the digestive tract are separate. Three or four sessile lobular diverticula are present on the ectal end of the ampulla. They are somewhat similar in shape but vary in size. The duct originates at one side of the ectal end of the ampulla from a cone-shaped expansion. It is uniform in diameter, about three times as long as the ampulla, and is lined with cuticula through- out its entire length. The ectal opening is lateral in position and near the intersegmental groove IV/V. No accessory glands are present at the ectal opening. The hypodermis of IV and V, with the exception of the dorsal third, is strongly thickened, the thickening being greatest in the vicinity of the spermathecal pores. Fridesicia TENERA n. sp. Deftnition.—Length, 9-17 mm. Somites, 52-39. Color, whitish. Prostomium rounded. First dorsal pore in VII. Setse, 4-6 per bundle in anterior region of body ; 2-4, usually 2, in posterior re- gion. Clitellum on XII and XIII. Lymphocytes elliptical. Brain about one fifth longer than wide; anterior margin concave, posterior margin convex, lateral margins converge cephalad. Peptonephridia each with several branches originating from common base, and some secondary branches. Anteseptal and postseptal parts of nephridia about equal in size; efferent duct arises from mid-ventral surface of postseptal part near septum. Spermiducal funnel about twice as long as wide ; duct long, much contorted, and confined to XII. Spermathec?e each with barrel-shaped ampulla communicating with digestive tract in V, and each bearing about seven globular sessile diverticula; two glands at ectal opening of spermathecal duct. For discussion of penial bulb and chylus cells see pages 622, 623. 021 Described from seventeen sexually mature specimens. Type and paratypes in the collection of the junior author; paratypes in the collection of the senior author. The specimens which fonn the basis of this description were foiuid in a compost pile in the forest grounds of the University of Illinois and were collected in October and November, 1910. Abilities.—This species seems to approach several other species, namely, F. lobifcra Vejdovsky ('79, p. 57), F. udei Bretscher ('99, p. 411), F. bcddardi Bretscher ('00, p. 29), F. macgregori Eisen ('05, p. 118), and F. californica Eisen ('05, p. 119). Because of brevity, use of general terms, and indefinite statements, the descrip- tions of F. udei and F. beddardi are somewhat puzzling and compari- sons troublesome, but the aggregate of differences is such that it is not difficult to separate F. tciicra from them. It differs from F. lobifcra in length, and in the characters of the brain, peptonephridia, and spermathec?e ; from F. udei in length, and in the characters of the peptonephridia, spermathecje, and nephridia ; from F. bcddardi in the characters of the peptonephridia, brain, and spermathecae ; from F. macgregori in length, munber of somites, and in the characters of the brain and spennathecae ; and from F. californica in length, num- ber of somites, and the characters of the brain and spermiducal fun- nel. In connection with the above comparison a certain discrepancy which appears in Eisen's description of F. macgregori should be noted. In his description of the brain, he says, "Brain anteriorly much convex; posteriorly slightly so." His text figure of the brain shows that the anterior margin is less convex than the posterior. It seems probable that the figure is correct. EXTERN.-\I, CHARACTERS The worms are relatively slender and delicate, having a length of 9-17 mm., and a maximum diameter of only about 0.4 mm. The number of somites varies from 52 to 59. The living worms are opaque and whitish in appearance. The clitellum is on XII and XIII and is only moderately developed. It is made up of transverse rows (PI. XCIX, Fig. 13) in which glandular cells alternate with clear one.s. The intersegmental grooves are obscure, except the first four or five. The setae are of the typical Fridericia type and are relatively large. The distal ends are sharply pointed, and the proximal ones (PI. XCIX, Fig. 14) are strongly recurved. In the anterior part o! the body the dorsal bundles contain 4-5 setae, the ventral ones usually 6; in the middle region both sets of bundles contain 4 setae; while in the posterior part, 2 is the predominating number. 622 INTERNAI, CHARACTERS Body ]VaU.—The hypoderniis contains unicellular flask-shaped gland cells (PI. XCIX, Fig. i6) which are numerous and generally distributed. They occur among the ordinary hypodermal cells, and each opens at the surface through the distal narrowed neck region. They are much larger than the hypodermal cells though shorter. They are approximately uniform in shape except in the thickened region of the first somite and the prostomium, where they are more elongate. The contents show a certain degree of polarity. In the ectal part of the cell the contents are of such a nature that they stain only slightly, while the opposite end is filled with material which stains deeply. The latter also contains a conspicuous nucleus. Brain.—The brain (PI. XCIX, Fig. 15) lies entirely in II and is about one fifth longer than wide. The anterior margin is concave ; the posterior margin is convex; and the lateral margins are gradually convergent anteriorly. Two pairs of supporting strands connect the lateral margins of the brain with the body wall. Peptonephrid'ia.—A pair of these organs open into the ventral side of the digestive tract in IV. Each is composed of several branches which arise from a common base, and most of which are directed caudad and terminate in VI. Secondary branching exists to some extent. Chylus Cells.—The chylus cell region is that part of the intestine included in XV-XVII. The chylus cells (PI. C, Fig. 18) are flask- shaped, and each cell contains the characteristic intracellular canal, which has a somewhat sinuous course, especially in the ectal end of the cell. The canal is ciliated throughout almost its entire length. There seems to be no special modification of the cytoplasmic layer surrounding the canal, such as exists in F. firma and some other species. Each cell contains a conspicuous nucleus, which usually lies in the chief bend of the canal. Between the apical ends of the chylus cells are fitted the wedge-shaped ental epithelial cells. The location of the chylus cells in XV—XVII is different from that in any other species in which they have been described and may, in accordance with the general view of Eisen, be a valid specific character. Nephridia.—The first pair of nephridia are connected with V/VI. The anteseptal and postseptal parts (PI. C, Fig. 17) are about equal in size. The efferent duct arises from the mid-ventral surface of the postseptal part, near the septum, and opens to the exterior slightly anterior to the corresponding ventral setae. The lumen is tortuous throughout its whole length. G23 Penial Bulb.—In many respects the structure of the penial bulb (PI. C, Fig. 20) in this species is similar to that of F. Urma, although a number of distinct differences are apparent. Unlike that of the latter species the body of the Ijulb is composed of cells of but one kind. They are large and glandular, and each contains a large, con- spicuous nucleus at the peripheral end. Each cell has a prolongation which extends to the penial lumen. The peripheral part of the cell stains deeply but the prolongation stains only very slightly. The sperm duct enters the bulb at the anterior end and extends oblicpiely ventrad, opening into the penial lumen. Figure 20 shows the struc- tural detail of this organ as it appears in a transverse section of the worm. This bulb is clearly of the lumbricillid type as defined by Eisen. It differs from that of F. firiiia in lacking the inner bulb cells which surround the penial lumen. Spermatheca.—Each spermatheca (PI. C, Fig. 21) is differen- tiated into ampulla, duct, and several diverticula. The ampulla is somewhat elongated and inflated in the middle. The ectal end bears a circle of globular sessile diverticula, usually seven in number and slightly dissimilar in shape and size. The duct arises from the center of this circle of diverticula and extends, with few curves, to its ectal opening in the lateral wall of the bod}^ slightly posterior to IV/V. Two pear-shaped glands are present at the ectal opening. The cuticula is reflected into the lumen of the duct and lines it for its entire length. Fridericia .\gilis Smith At the time that this species was described*, certain organs of the EnchytrccidcE were not considered of as much systematic importance as at present, and, accordingly, they were ignored, or received scant attention in the description of F. agilis. In view of the large niunber of species of the genus already known and of the large number that are almost sure to be made kno\\^^ in the future, it is important that descriptions should be quite detailed. For this reason an examina- tion of additional material and a further study of old material have been made and a more extended description prepared, no attempt Ijeing made to distinguish the old material from the new. Definition.—Length, 25-30 mm. Diameter, 0.63-0.82 mm. Somites, 57-66, average, 62. Color, whitish. Prostomium blunt and rounded. First dorsal pore in VII. Set?e, 2-4, usually 2, per bundle. Clitellum on XII-XIII. Lymphocytes numerous, broadly elliptical. •In 189S, in the BuUetin of the Illinois State Laboratory of Natural His- tory, Vol. IV, Art. VIII, pp. 288-289. 624 Brain one half longer than greatest width; anterior margin slightly concave, posterior margin quite convex. Peptonephridia large, very much branched, opening into digestive tract in posterior part of III. Dorsal vessel arises in XIX. Nephridia with anteseptal part equal- ing postseptal part in size; efferent duct arises from posterior end of latter. Spermiducal funnel twice as long as its diameter. Sper- mathecre each with duct, ampulla, and diverticula; duct about three times as long as ampulla, with a few very small unicellular glands at ectal opening ; ampulla with about nine similar, globular diverticula. For discussion of penial bulb and chylus cells see pages 625, 626. Described from twelve sexually mature specimens. Type in the collection of the Illinois State Laboratory of Natural History. Para- tyi>es in the collections of each of the authors. The specimens on which the description of this species is based were collected at Havana, Illinois, in April and May of 1895. They were found abundant in the wooded banks and bottom-lands of the Illinois River, under logs and in the damp rich soil, mingled with decaying vegetation. EXTERNAI, CHARACTERS The body of the worm is smooth, rather robust, cylindrical, taper- ing very gradually towards the two extremities. The length of mature specimens is 25-30 mm. The diameter is greatest in the region of the clitellum, where, in alcoholic specimens, it is 0.63- 0.82 mm. The number of somites varies from 57 to 66. The inter- segmental grooves, except the first four or five, are indistinct. The intersegmental groove IV/V differs from the adjacent ones in being broader and more shallow. The prostomium is blunt, rounded, and smooth. The color of the living worm is whitish. The clitellum, which is on XII-XIII, is moderately developed and composed of cells of but one kind. The setae vary from 2 to 4 per bundle, the more usual number being 2. Each seta is rather strong, acute at the distal extremity, and distinctly bent at the proximal end. INTERNAL CH.\RACTERS Lymphocytes.—The lymphocytes are abundant in most parts of the body. They are broadly elliptical in outline, and the long axis of each averages about 0.025 "''"''• Brain.—The brain (PI. C, Pig. 22) is in I and II. The anterior margin is slightly concave, the posterior margin is quite convex, and the lateral margins are almost parallel. In transverse section it is ovoid. A pair of supporting strands extending from the latero- 625 posterior parts of the brain in a latero-caudal direction attach it to the body wall. Pcptoucphridia.—A pair of these organs connect independently with the ventral side of the digestive tract in the posterior part of III. The apparent absence of septum III/IV 'makes it difficult to determine the exact point of separation between III and IV. These glands open into the digestive tract verj' close to III/IV, apparently in the ix)sterior part of III. They are directed caudad and do not extend beyond IV/V. Each branches profusely and in an irregular, dendritic fashion. Chyltis Cells.—Chylus cells (PI. C, Fig. 23) are present in the walls of the intestine in yi XII-XVI. They are somewhat flask- shaped, the ectal ends being broader than the ental ones. They vary in fomi considerably, the length of some being nearly three times their diameter, while in others it is but little greater. It is possible that these ratios are subject to change in the same cell, owing to the different states of contraction of the intestinal wall. The ental part of the intracellular canal is straight; the basal part is sinuous and somewhat branched. Cilia are present throughout the greater part of the ental portion of the canal, and are directed toward the lumen of the intestine. The canal is lined by a specialized layer of cyto- plasm, which is everywhere uniform in thickness and structure. The perivisceral blood sinus comes into contact with the basal portion of each chylus cell. The cells of the ental epithelial layer are wedge- shaped, and usually occur singly between the apices of the chylus cells. The ental surface of these cells is thickly covered with long cilia. Interstitial cells are absent. Eisen ('05) made careful studies of these peculiar cells in a number of species which he described from the west coast of North America and was convinced that their location, form, and size furnish good specific characters. Since other students of Fridcricia have not described these structures in any detail, Eisen's conclusions were necessarily based on an investigation of but a small number of the many known species, and more extended studies must be made be- fore his views can be justly weighed. The characters of the chylus cells in F. finiia, F. tenera, and F. agilis do not coincide with those of any other species in which such cells have been described, but appear to present distinct specific differences. Ncpliridia.—The anteseptal and i^ostseptal parts (PI. CI, Fig. 24) of each nephridium are api>roximately equal in size, and both are well developed. The efferent duct arises from the posterior end of the postseptal part. The true origin (PI. CI, Fig. 25) of the duct is 026 occasionally obscured by a peculiar modification of the nephridium in which the posterior half of the postseptal part is bent ventrad and cephalad, and is in such close relation to the ventral side of the an- terior half that the origin seems to be near the septum. Spermidncal Funnel.—The spermiducal funnel (PI. CI, Fig. 26) is very large, and the two fumiels occupy the greater part of the coelom in XL They are barrel-shaped and about twice as long as their diaineter. Each has a distinct reflected collar set off from the body of the funnel by a constriction. The diameter of the collar is less than the maximum diameter of the funnel. The sperm duct is long, slendeT, and confined to XII. Penial Bulb.—The structure of the penial bulb (PI. CI, Fig. 27) so closely resembles that already described for F. Urma that it seems unnecessary to give a detailed description. The shape, relative size, musculature, and relation to the body wall are all about the same as in that species. Similarly, also, the body of the bulb is composed of two distinct sets of cells, viz., the inner bulb cells, which immediately surround the penial lumen, and the peripheral bulb cells, which com- prise the greater part of the mass of the organ. The relation of the bulb to the sperm duct is also similar to that described for F. Urnia. Spennatheccc.—Each spermatheca (PI. CI, Fig. 28) has a dis- tinctly differentiated duct and ampulla, the latter bearing diverticula. The duct is slender and about three limes as long as the ampulla. It unites with the ectal end of the latter at the middle of the ring of diverticula and extends, with few curves, to the ectal opening, which is laterad and near IV/V. A few inconspicuous gland cells are pres- ent about the ectal opening. The ampulla is barrel-shaped and thick- walled. It bears a ring of about nine similar lobular diverticula at the ectal end. The walls of the diverticula are comparatively thin, and the lumina are in direct communication with the lumen of the ampulla. The spemiathecae communicate independently with the digestive tract on its dorsal side in the posterior part of V. The spermathecfe are quite large and occupy a considerable part of the space in V. The Function of the Chylus Cells in Fridericia The homologies and functions of these peculiar cells are, indeed, quite problematical. The intimate connection of the cells with the digestive tract and their close contact with the blood vascular system indicate that they have an. important role in some metabolic process. Michaelsen ('86, p. 296) has maintained that they are organs of ab- sorption. Eisen ('05, p. 107) also favored this interpretation, and at- 627 tempted to support it by the assumption that the presence of the cilia in the canal indicates that the canal itself is simply an invagination of the ciliated apical surface of the cell, and that a means is thus provided for bringing the nutritive fluid into close relation with the blood sinuses at the base of the cell. He further held that the prob- able function of the cilia in the apical end of the canal is to facilitate the introduction of the nutritive fluid, and that the canal is the means whereby greater intestinal surface and rapid absorption are insured without consequent diminution or weakening of the intestinal wall. Vejdovsky ('06, p. 65) and Cejka ('10, p. 17) assign a secretory function to these cells, and assume that the cilia aid in the passage of the secretion from the canal into the lumen of the alimentary tract. They also claim that the cells are produced by modifications of the ] deeper-lying cells of the epithelium and that, in sections, intermediate 3 stages in such a development can be seen. ; We have made no observations which enable us to contribute J directly to the determination either of the function or of the mode J of origin of these cells, but have made some observations on the ar- ; rangement of the cilia of the canals and offer some suggestions as to their probable function. i{ We have numerous sections of specimens from each of the species • of Fridericia herein described which show clearly that the cilia of the canal are directed towards the opening at the apical end as repn resented by Vejdovsky, and have found no instance in which they extend in the opposite direction as figured by Eisen ('05, PI. XVIT and XX). We believe that the cilia may have a function quite dif- ferent from the one previously assumed. Experiments and observations by Cuenot, although not dealing directly with this subject, throw light on the problem in an interest- ing way. He has shown ('97, p. 105) that the vibratile cilia in the nephrostome of the nephridium of Oligochccta can not introduce the cccloiiiic fluid into the interior of the nephridium except when inequali- ties in the fluid pressure in the lumen will permit. Furthermore, he has shown that solid Ixjdies in suspension in the coelomic fluid can not pass through the orifice of the nephrostome because the thickly set cilia play the role of a sieve or filter and form an impassable bar- rier to any solid particles, except possibly those which are excessively minute. In view of these facts it seems probable that the prime function of the cilia of the nephrostome is the prevention of the passage of solid particles into the nephridial lumen rather than the introduction of fluids into the lumen as has been assumed. It .seems, then, reasonable to infer that the cilia in the apical part of each chylus cell canal also function as a filter, preventing the in- 628 troduction of solid particles from the intestine, and that they do not induce a fioAV of fluid into the lumen as claimed. No experimental evidence has yet been secured to substantiate this inferential view but it seems plausible. For some reasons there is even a better basis for ascribing this filtering function to the cilia of the chylus cells than to those of the nephridia. In the first place, the canals of the chylus cells end blindly, while the nephridial canals have external openings and in consequence the nephridial lumen may sometimes be almost or entirely empty, when, as claimed by Cuenot, it is possi- ble for the cilia of the nephrostome to aid in the introduction of the fluids. On the other hand, the chylus cells occur in a part of the intestine where, under normal conditions, the nutritive fluid is con- stantly present. Since the intracellular canal of each cell has but one opening, the canal is presumably at all times filled with fluid, the amount of which is far more dependent on pressure conditions in neighboring parts than on any action of the cilia. The powerful and frequently changing muscular contractions in various parts of the body are accompanied by more or less violent disturbances of pressure and consequent movements in the fluid contents. It seems reasonable to assume that such disturbances would more profoundly affect the extensive digestive tract and its contents than the diminu- tive nephridium, which is to such a considerable extent freely suspended in fluid. In the second place, the apical ends of the chylus cells are exposed to great quantities of solid particles, since the con- tents of the digestive tract must pass by them, and yet none of the preparations examined by us have shown any indication of the pres- ence of such material in the canals. Unless there were some provi- sions for the prevention of the entrance of solid matter these blind canals would be almost sure to contain such material and their func- tional efficiency be correspondingly reduced. MARIONINA Michaelsen During the autumn of 1895 a study of the animal life in the waterworks reservoir of Urbana, Illinois, was made by Miss Bertha V. H. Forbes, a student in the University of Illinois, and in her collections were a moderate number of small enchytrseid worms. They were turned over to the senior author of this paper and were partly worked up at that time, but circumstances prevented the com- pletion of the work, and nothing further was done with the worms until the present year, when their further study was undertaken by the junior author. They proved to be an undescribed species of Marionina. 629 This is not only the first record of a species of Marionina for Illinois, but it is also the first time a representative of this genus has been reported from the United States, although Eisen ('05) has described two new species from Alaska. The genus Marionina was established by Michaelsen in 1889. Previous to that time forms belonging to this genus were scattered among a number of the older genera, the old genus Pachydriliis (Liiinbricillus) receiving the larger part. Michaelsen ('00, p. 73) defines the genus Marionina as follows: "Borsten S-formig gebogen. Riickenporen fehlen; Kopfporus klein, zwischen Kopflappen und i. Segm. Das Riickengefass entspringt postclitellial und besitzt kein Herzkorper. Peptonephridien fehlen. Hoden massig. Samenleiter lang. Samentaschen ohne Divertikel." Eisen ('05, p. 90) has extended this definition somewhat: "Setae sigmoid, as in Lumbricillus. Head pore small, l>etween the prosto- mium and somite I. No dorsal pores. Blood red or yellow. Dorsal vessel rises posterior to clitellum. No cardiac gland. No peptone- phridia. Spemi-ducts comparatively long and narrow. Penial bulb without interior muscular strands. Testes undivided, each covered with a small sperm-sac. Ventral glands present or ab.sent. Nephridia with entire postseptal and with comparatively large head-like ante- septal." The close similarity between the definition of Marionina and that of Lunibricilhts is apparent, the chief diiTerence being the presence of multilobed testes in the latter. Eisen ('05, p. 90) thinks that an- other difference may be derived from the nephridium, "which in Marionina seems to be characterized by a long head-like anteseptal, while in Lumbricillus the anteseptal consists of merely the nephro- stome." There are insufficient grounds for giving such a character generic rank, since the anteseptal part of the nephridium in Marionina is quite variable. In M. werthi Mchlsn., M. falclandica Mchlsn., and in several other species, the anteseptal part consists of the nephro- stome only. Marionina is one of the smaller genera of the family Enchy- trccidcc. At present twenty-six apparently valid species are assigned to it, of which only two have been recorded for North America. They are M. alaskcr Eisen and M. anicrica)ta Eisen, both from Port Clarence, Alaska. Marionina forees-^ n. sp. Definition.—Length, 5-6 mm. Diameter, 0.221-0.238 mm. Somites, 25-28. Setre slightly sigmoid; those of a bundle equal in size; 2-4 per I)undle. Clitellum on XII-XIII; developed only on 630 dorsal and lateral surfaces. Prostomium slightly pointed. Head pore on o/I. Cuticula very thick and resistant. Lymphocytes few in number, oblong, nucleated, cytoplasm heavily granular in appear- ance. Brain about twice as long as wide; anterior margin conical, posterior margin emarginated, lateral margins convergent anteriorly. Peptonephridia lacking. No ventral glands. Septal glands in IV-- VI. Dorsal vessel arises in XIII. Blood yellowish. Anteseptal part of nephridium a mere nephrostome; postseptal part large, flattened laterally, efferent duct arising from its ventral side near posterior end. Spermatheca with distinct duct and ampulla; ampulla pear- shaped; not connected with the intestine and with no diverticula; duct narrow, rather short, with no glands at ectal opening. Length of spermiducal funnel about twice its diameter. Testes undivided; sperm sacs lacking. For a discussion of the penial bulb see page 632. Described from five sexually mature specimens, although a num- ber of others were examined in connection with the study on the living fomis. The type and a paratype are in the collection of the junior author, and paratypes in that of the senior author. The above specimens which fonn the basis of the description of this species were found in the mud and settlings in the bottom of the waterworks reservoir of Urbana, Illinois. At the time of col- lection they occurred only in moderate numbers. The dates of col- lection range from October 23 to November 26, 1895. The species name is given in recognition of the discoverer. Miss Bertha V. H. Forbes. EXTERNAL CHARACTERS The specimens are small, their length being only 5-6 mm. The figures given are for the living specimens and apply to all the sexually mature individuals. As measurements of alcoholic material gave al- most exactly the same result, it appears that in this species of the BncJiytrccidcr authentic data regarding length can be secured from the examination of preserved material. This fact may be due to the exceptionally thick and resistant cuticula, which may perhaps also account for the fact that living specimens have but a slight range in contraction and extension. The number of somites varies from 25 to 28. The body is cylindrical, and the diameter is approximately uniform for almost its entire length, only the first somite and the last two or three showing gradual increase in size. The diameter varies from 0.22 to 0.24 mm. The clitellum is on XII-XIII and is slightly developed dorsad and laterad but is interrupted on the ventral sur- face of the body. The prostomium (PI. CI, Fig. 29) is rounded, yet fiSl slightly pointed. The liead pore is present at of. The anterior four or ti\'e intersegmental grooves are distinct, Init beyond this region they are obscure. Three_to eight secondary trans\erse grooves occur on all of the somites posterior to II or III with the exception of XII and XIII. A conspicuous feature of living specimens is the presence of definitely arranged spots on the external surface. They are disposed in transverse rows, 2-3 per stmiite, each row containing 2-4 spots. One row is slightly anterior to the .setie, another slightly posterior to them, and a third caudad to the latter. These markings do not appear in alcoholic specimens. Sctcc.—The setse are slightly sigmoid, but in an examination of the living material or of specimens mounted in toto it is very easy to overlook their slightly sigmoid form and to mistake them for straight set;e. Trans\'erse sections of the worm (PI. CII, Fig. 31) re- veal the curves distinctly. The setK are arranged in four bundles ]>er somite, two ventral and two lateral. The number of setae per bundle varies from 2 to 4. In the anterior region the numbers 4 and 3 seem to predominate, while in the posterior region there arc usually 3 per bundle, and but 2 in the last one or two somites. The distal extremities of the setse of each bundle are all curved in the same direction, those of the lateral bundles curving ventrad and those of the ventral bundles curving dorsad. The proximal ends of the setae also show a slight cur\-ature in a direction opposite to that of the distal portion. Ciiticiila.—There is a very thick resistant cuticula of which the thickness is about equal to that of the combined thickness of the hypodermis and the muscular layers of the IxKly wall. This cuticula is so resistant that in some cases it caused considerable trouble in sectioning. It appears to be approximately uniform in thickness throughout the length of the body. INTERN.VL CII.\R.\CTlCRS L.yiiij^hocytcs.—The lymphocytes are scattered sjjaringly through- out the greater part of the crelom. They are nucleated and the cytoplasm is granular. The shajjc varies from an oval to a decidedly oblong form, and the length of some lymphocytes is about twice that of others (PI. CI, Fig. 30). Brain.—The brain lies in I, If, and III. chiefly in IT. Its length (PI. CTI, Fig. 32) is approximately twice the greatest width. The anterior margin is decidedly conical; the posterior margin is con- sp'cuously emarginatc ; and the lateral margins converge anteriorly. 632 In transverse section it is ovoid in shape. Studies on tlie living speci- mens revealed the fact that the depth of the posterior emargination is subject to some \ariation, dependi^ig upon certain changes in the states of contraction of the animal. Sometimes the emargination is rather shallow, although always distinct; at other times the emargina- tion is deep. Two pairs of strands connect the brain with the body wall, one pair arising from the lateral margins of the brain and the other pair from the posterior lobes. Blood Vascular System.—Studies of both living and preserved specimens show that the dorsal vessel arises in XIII. A distinct swelling of this vessel occurs in XIII, where the diameter exceeds that of any other region. X^o cardiac body is present. The peri- visceral sinus appears in the region of the clitellum, and in transverse sections of the worm has a distinctly beaded appearance entirely around the intestine. There is reason to believe that this appearance is due to the fact that there are membranous partitions which run lengthwise of the sinus, thus dividing it into a number of longitudi- nal tubes. These tubes appear to be distended with blood, and when sectioned transversely exhibit a more or less circular outline. This sinus extends posteriorly from the origin of the dorsal vessel, and shows a decided redtiction at the septa. The ventral vessel branches at IV/V. The blood is yellowish in the living specimens. NepJiridia.—The first nephridia are connected with VII/VIII. The anteseptal part (PI. CII, Fig. 33) is very small and consists merely of the nephrostome. The postseptal part is large, elongated pos- teriorly, and flattened laterally. The efferent duct arises from the ventral side of the postseptal part, slightly anterior to its posterior end. Spermiducal Funnel.—The spermiducal funnel is about twice as long as its average diameter. In some specimens the funnel is slightly flattened laterally. A set of typical measurements are as follows : length 0.07 mm; average diameter 0.039 "I'^i- The duct is very long, much contorted, and confined to XII. Pcnial Bulb.—This organ is well developed, and is rather con- spicuous in transverse sections. It is situated on a distinct invagina- tion of the body wall. It is globular in shape (PI. CII, Fig 34), partic- ularly in cases where it is extruded. In the retracted" condition it is ovoid. The organ as a whole is covered with a thin peritoneal layer, beneath which lies a comparatively thin layer of muscle tis- sue. The body of the bulb is composed of a large number of similar, more or less spindle-shaped cells, whose long axes extend towards the external surface. Each cell is conspicuously nucleated and usu- ally the cell walls are distinct. When retracted, the duct enters the bulb well down on the ental side and extends through it to the ex- terior opening at the edge of the invagination. In the everted condi- tion the lateral part of the bulb is depressed, the invaginated surface becoming superrtcial, the bulb becoming more globular in shape, the position of the entrance of the duct being shifted to the dorsal side of the bulb, and the duct extending flirectly to the ectal surface. Apparently the cells of the bulb are all of one kind, and all extend to the surface of the bulb, none haxing l)een found emptying into the duct. Speiiiiathcccr.—A pair of these organs is present in V. Each spermatheca (PI. CII, Pig. 35) consists of a well-defined duct and an ampulla. The ectal opening of the duct is in the intersegmental groove IV/V, and is latero-ventrad in position. Xo definite glands are present at the ectal opening. The wall of the duct is thick, and shows a number of large nuclei scattered throughout its length. The lumen is very fine. The duct is nearly straight, exitending directly to the dorsal side of the digestive tract, where it expands to form the ampulla. The ovoid ampulla forms a closed sac, having no con- nection with the digestive tract. Its walls are quite thin, and in all of the specimens exam'ned it was partially or entirely filled with sperm cells. 034 LITERATURE CITED Bretscher, K. '99. Beitrag zur Kenntnis der Oligocliaetenfauna der Schweiz. Rev. Suisse Zool., 6 : 369-426. 7 fig. '00. Mitteilungen iiber die Oligocliaetenfauna der Schweiz. Rev. Siiis.se Zool., 8:1-44. 3 pi. V Cejka, B. '10. Die Oligochaeten der Russischen in den Jahren 1900-1903 unternommen Nordpolarexpedition. I. Uber eine neiie Gat- tiing der Enchytraeiden (Hepatogaster). Mem. Acad. vSci. St. Petersb. (8), 29, No. 2: 1-29. 3 pi. Cuenot, L. '97. Etudes physiologiques sur les Oligochetes. .Arch. d. Biol., 15 : 79-124. 2 pi. Eiseu, G. '72. Om nagra arktiska Oligochaeter. Ofv. Ak. Forh., 1872: 1 19-124. I pi. '05. Enchytrasida? of the West Coast of North America. Harri- man Alaska Expedition, 12: i-i66. 20 pi. New York. Michaelsen, W. '86. Ueber Chylusgefasssysteme bei Enchytraeiden. Arch, f. mikr. Anat., 28:292-304. i pi. '89. Synopsis der Enchytraeiflen. Al)Ii. nat. Ver. Hamburg, IT : 1-59. I pi. '00. Oligochaeta. Das Tierreich, 10 Lief.: I-XXIX+i-375. Ber- lin. Moore, J. P. '95. Notes on American Enchytrreidrc. I—New Species of Eridericia from the Vicinity of Philadelphia. Proc. Acad. Nat. Sci. Phila., 1895:341-345. i pi. Smith, E. '95. Notes on Species of North American Oligochasta. Bull. ' 111. State Lab. Nat. Hist., 4: 285-297. Smith, S. I., and Verrill, A. E. '71. Notice of the Invertebrata dredged in Lake Superior in 1871, by the U. S. Lake Survey, under the direction of Gen. C. B. Com,stock, S. I. Smith, naturalist. Am. Journ. Sci. Arts, {?,), 2:449-451. fi35 Vejdovsky, F. '79. Beitrjige zur \-ergleichen(len Alorphologie der Anneliden. I. Monograpliie der Enclivtraeiden. 6j pp., 14 pi. Prag. '06. Zweiter Beitrag zur Hi'iniociiltlicoric. Zcit. 1. wiss. Zool., 85 : 4S-73. 2 pi. EXPL.\N.\T10N OF PLATKS Ai!iiuF.vi.\'rioNS />/. .f.Jilnod sinus. w., musculature. (•/(/. (-'/..rliloragon- cells. n.,nucleus. f /( v. <'/., cliylus eel!. />., l)eritoneuni. ri'/., cilia. /i.fc.j., perivisceral lilond sinus. cl. c'/.,elitellar cells. /j^)i.fc.i.,penial bulb invagination. c«<.,cuticula. I'en. /M)H.,pcnial lumen. 01. r/>.r7.,ental epitbelial cell. />(». /"o., penial pore. /jji/).,hypodermis. />(,'r.g/.f'/., peripheral gland cells. III. h. ('/., inner bulb cells. r.)»., retractor nniscle. ill. c'/.c'ii.. intracellular canal. j/i.rf., sperm duct. /.fV^, lining layer of cytoplasm. Pt.ATE XCVTTT Pridcric'm firiiia Outline of anterior ciul, lateral view. Seta. Superficial section of tbe clitellar cells. Outline of brain, dorsal view. Outline of nephridium. Part of transverse section of intestine in cbylus cell region. Chylus cell, greatly enlarged. Outline of spermiducal funnel. Fig. 0. Longitudinal section, oblif|ue to sagittal plane, through lower part of In Plate XCIX Fridericia Uriiw—cont. Fin. in. Pcnial bulb in a transverse section of the worm. Fir,, n. (Outline of spermatheca. 'Fridericia lenera Fin. 12. Outline of anterior end, lateral view. Fig. n. Superficial section of clitellar cells. Fig. 14. Seta. Fin. 1."). 0\itline of brain. Fig. Ifi. fdands in the hypodermis. Fig. Plate C Fridcriria ten era—cont. Fig. 17. Outline of nci)liri