0 .COJW ARCHIE CARR BULLETIN OF THE FLORIDA STATE MUSEUM BIOLOGICAL SCIENCES Volume 3 Number 2 FOSSIL TURTLES OF THE GENUS TERRAPENE IN FLORIDA Walter Auffenberg UNIVERSITY OF FLORIDA Gainesville 1958 HERPETOLOGY LIBRARYFLORIDA STATE MUSEul-,1 The numbers of THE BULLETIN OF THE FLORIDA STATE MUSEUM, BIOLOGICAL SCIENCES, will be published at irregular intervals. Volumes will contain about 800 pages and will not necessarily be completed in any one calendar year. WILLIAM J. RIEMER, Editor ROLAND F. HusSEY, Associate Editor All communications concerning purchase or exchange of the publication should be addressed to the Curator of Biological Sciences, Florida State Museum, Seagle Building, Gainesville, Florida. Manuscripts should be sent to the Editor of the BULLETIN, Flint Hall, University of Florida, Gainesville, Florida. Published 27 June 1958 Price for this issue $.60 FOSSIL TURTLES OF THE GENUS TERRAPENE IN FLORIDA WALTER AUFFENBERG 1 SYNOPSIS: Recent paleontological work in the Pleistocene of Florida has made available a larger series of fossil box turtles, Terrapene, than.has been studied pre- viously. A large number of modern specimens were' examined to determine the extent of variation existing in various populations and to evaluate characters found in the fossils. It is shown that two box turtles are represented in the known Pleistocene localities of the area. There is some evidence of intergradation be- tween them, and the two fossil forms are thus considered subspecies of the Recent form, Terrapene carolina. The smaller Pleistocene form represents the modern T. c. carolina Or T. c. bauri or both. The larger fossil form is termed T. c. put- nami. Terrapene innoxia and T. formosa are considered.synonyms of T. c. carolina or T. c. bauri or both. Terrapene canaticulata and T. antipex are synonyms of T. c. putnami. Terrapene singletoni seems to represent a specimen intermediate between T. c. carolina and T. c. putnami. Certain fossil populations are inter- mediate between these same two forms and are considered intergrades. A prob- able history of the genus Terrapehe in southeastern United States is presented. The Pleistocene localities of Florida are tentatively correlated on the· basis of the box turtles now available frbm them. Fossil box turtles are fairly common over much of eastern North America. Their remains are almost always composed of fragmentary elements of the 6arapace and plastron. Occasionally complete shells are found; rarely a skull. Most vertebrate paleontologists are only casually interested in fossil turtles, particularly Pleistocene forms. As a result, much remains to be learned concerning late Cenozoic turtles. No single comprehensive study has ever been attempted of the anatomy of Terrapene or. its evolutionary history, though several good regional studies have been published recently. Earlier papers tend to-obscure relationships of fossil and Recent forms under a pro- fusion of new names based, in many cases, on individuals so fragmen- tary as to make identification of new material almost impossible. This is particularly true in Florida, where six species have been described frpm Pleistocene deposits alone. The Recent species of Terrapene of the United States, ornata and carolina, are rather well understood, at least in broad outline. The Mexican species, T. mexicana, coahuila, klauberi, and nelsoni are little known. Some, if not all of them, are close to T. carolina. T. coahuila seems to be the most primitive member of the genus and m6st distantly ' i The author is an Associate in the Florida State Museum, Assistant Professor . of Biological Sciences, and curator in vertebrate paleontology of the University of Florida Collections. His most important contributions are concerned with the fossil history of small amphibians and reptiles. Manuscript submitted 28 May. 1958.-ED. 54 BULLETIN FLORIDA STATE MUSEUM Vol. 8 related to T. carolina. The latter includes four currently recognized geographic races: T. c. carolina, maior, bauri, and triunguis. The geographical and morphological limits of these forms are not clearly defined at present. Their relationships are not adequately understood, though mafor certainly seems the most primitive member of the spe- cies as now defined. Little is actually known of their subspecific characters other than ihat they care rather variable. This last fact is important when considering the described fossil forms, since it has been almost entirely ignored by past workers with the exception of Milstead (1956) and Barbour and Stetson (1981). The degree of vari- ability in Pleistocene populations has never been fully demonstrated. In the present studyit is assumed to be as great as in the Recent forms. ABBREVIATIONS AND ACKNOWLEDGMENTS The following abbfeviations are used when referring to collections: ACM-Amherst College Museum; AMNH-American Mu'seum of Nat- ural History; FGS-Florida Geological Survey; MCZ-Museum of , Comparative Zoology; UF-University of Florida Collections; USNM -United States National Museum; and VU-Vanderbilt University. I wish to thank the persons responsible for the collections in the various institutions mentioned above for the loan of specimens in their care. To A. Holman and R: Weigel, University of Flotida, I wish to express appreciation for allowing me to examine fossil box turtles collected by them while working on particular vertebrate deposits in the state and which have not as yet been deposited in a public institution. I also wish to thank W. Milstead and E. E. Williams for their valuable criticisms. DISCUSSION OF PREVIOUS STUDIES ON Tenapene The earliest fossil referred to this genus is Terrapene longinsulae Hay (1908a) from the lower Pliocene of Phillips County, Kansas. It has been thought to be somewhat closer to T. ornata than to T. care- lina, but the specimen deserves additional study. So far box turtle remains are unknown from the Florida Miocene or Pliocene. The Pleistocene forms from Florjda obviously belong to the carolina com- plex, so T. longinsulae has not been seriously considered as a possible - close relative in this study. Terrapene carolina appears first in the Pleistocene of Nosth Amer- ica. Remains have been found from Maryland to Texas and from Pennsylvania to Florida. A number of fossil species obviously be- longing to the carolina group have been described from Florida and Georgia, larkely by Hay. These are as follows: 1958 AUFFENBERG: TERRAPENE IN FLORIDA 55 TERRAPENE PUTNAMI Hay (1907).-The hol6type is represented by a single hypoplastral element (AMNH 6097), dredged from the AlaGa River, Hillsborough County, Florida, about one mile above its entrance into Tampa Bay. Described on the basis of its large size and the thickness of the element, Hay estimated the carapace to have been about 265 mm. long. He also assignad two additional fragments of a carapace of a large box turtle in the collections of Vanderbilt Uni- versity to the same species. He e-stimates the carapace length of one of these specimens to have been about 820 mm. long. The holotype was taken from what was then termed the Peace Creek beds, orig- inally thought to represent the Pliocene. Subsequent work has sh6wn that the Peace Creek beds include both Pliocene and Pleistocene vertebrate remains (Wood, et al., 1941). Hay placed the species in the Pliocene with some reservation, however some subsequent authors have referred it to this period without question. New material indi- cates that the pieces are almost certainly Pleistocene. TERRAPENE CANALICULATA Hay (1907).-The holotype (USNM 5500) is composed of several fragmentary pieces of the carapace and plas- tron of a large bex turtle estimated to have had a carapace length of about 200 mm. The type was collected from Pleistocene deposits on either Whitemarsh or Skedaway Islands, below Savannah, Georgia. The most important diagnostic features are the- large size, a sharp longitudinal keel on the peripherals above the bridge, and a broad gutterlike groove above this keel. TERRAPENE ANTIPEX Hay (1916).-The holotype, originally in tbe collection of the Florida Geological Survey, is now in the United States National Museum (USNM 8820). It is represented by the posterior lobe of the plastron of a large box turtle estimated to have had a carapace length of about 220 mm. The specimen was presum- ably collected from stratum 8, Vero Beach, Indian River County, Florida. Stratum 2 has been shown to represent the Pleistocene bed. Stratum 3 is a much more recent deposit, containing Recent verte- brates as well as Pleistocene forms eroded from stratum 2. Terrapene antipex is here assumed to have originated from stratum 2, as will be discussed below. In addition to the type, another posterior lobe, part of an anterior lobe, and a few small pieces of carapace from the same locality are referred by Hay to this species. Still another posterior lobe from 28 miles south of St. Augustine, St. Johns County, Florida, and a fragment of a carapaee from the same locality are also referred to this species by Hay. Terrapene antipex is said to differ from T. 56 BULLETIN FLORIDA STATE MUSEUM Vol. 8 canaliculata in having a less conspicuous lateral keel, a thinner shell, and peripherals which are less flaring at the posterolateral margin of the shell. From putnami it is said to differ chiefly in size, being much smaller, and having a proportionately thinner hypoljlastron, TERRAPENE INNOXIA Hay (1916).-The holotype (USNM 8824) is a complete carapace, said to come from stratum 2, Vero Beach, Indian River County, Florida (fig. I). In the same collection a second cara- pace bearing the same locality data, and several fragments of the carapace and plastr6n of a similar-sized turtle from the same beds, are referred to this species by Hay. The major diagnostic features of this fossil species concern the size of the pieces, which represent a turtle much smaller than T. canaliculata, antipex, or putnami. The lateral keel so well developed in antipex and canaliculata is almost absent. The posterior peripherals are much less flared. From T. caro- lina (T. mafor of Hay, 1916) it differs in having the highest point of the carapace dver the middle of the shell rather than poiterior to the middle (Hay was using modern specimens of populations now termed T. c. bauri); in having a slightly less excavated nuchal bone at its anterior edge; in having a well-developed nuchal scale, which is said to be suppressed in Recent forms; and in having the rear. marginal scutes uniformly lower than in Recent shells he examined from Florida. TERRAPENE FORMOS'A Hay (1916).-The holotype is represented by the greater part of the posterior two thirds of a carapace (USNM 8825) taken from a Pleistocene Bssure in the Ocala limestone at Ocala, Mar- ion County, Florida (fig. 2) This is a poorly defined species that seems to have been described as new only because it came from ' Pleistocene bads and was slightly different than the type and referred material of T. innoxia from Vero Beach. TERRAPENE SINGLETONI Gilmore (1927).-The holotype (USNM 11181) consists of the greater part of a carapace from the Melbourne bone beds (presumably equivalent to stratum 2 of Vero Beach), two miles west of Meibourne, Brevard County, Florida. It is distinguished from all other fossil and Recent species of eastern United States, with the exception of T. canaliculata and putnami, in being much larger, the shell being 200 mm. long. From canaticulata it differs in being slightly smaller, in having a thinner shell, in lacking a lateral keel above the bridge, in being narrower, and in having peripherals which are less flared posteriorly. Gilmore placed Hay's T. anUpex in the synonymy of T. canaliculata Hay, largely on the basis of similar proportions of the posterior lobe of the plastron. Several complete carapaces from Melbourne in the 1958 AUFFENBERG: TERRAPENE IN FLORIDA 57 A B Figure 1.-Type of Terrapene innoxia Hay, USNM 8824, stratum 2 (?), Vero Beach, Indian River County, Florida, late Pleistocene (or Recent ?), referred to Terrapene carolina bauri. (A) dorsal view. (B) lateral view. St{ c I / f /1 1 /h ..\l / C ' - / C A B Figure 2.-Type of Terrapene formosa Hay, USNM 8825, from fissure fill in limestone quarry at Ocala, Maribn County, Florida, Pleistocene, ref6rred to Terrapene carolina (bauri or carolina). (A) dorsai view. (B) lateral view. (C) , posterior view, right half: . 58 BULLETIN FLORIDA STATE MUSEUM Vol. 8 collections of Amherst College and the United States National Museum (some of which are now in the Museum of Comparative Zoology) are referred by Gilmore to canaliculata (fig. 8). One specimen (USNM 11428) is provided with an associated complete plastron. The definite association of a plastron fitting, Hay's description of antipex with q carapace identical to his description of canaliculata leaves little doubt that Gilmore is correct in placing the former in the synonymy of the latter. ' iN A B C Figure 8.-Specimens typical of those referred to Terrapene carolina putnami (Hay), all from Golf Course locality, approximately three miles west of Melbourne, Brevard County, Florida, late Pleistocene. (A) dorsal view, AMC 25-144. (B) lateral view, AMC 25-145. (C) ventral view of carapace and plastron, showing axillary scale in contact with the third, fourth, and fifth marginal scutes, USNM 11428. Barbour and Stetson (1981), in reviewing the fossil b6x turtles then known fr6m Florida, placed T. singletoni, T. innoxia, and T, formosa in the synonymy of T. canaliculata, basing this action largely on a real- ization of the extreme variation within the shells of Recent T. carolina. The Pleistocene form, T.· canaliculata, was diagnosed mainly on .size alone. Terrapene putnami was retained as a much larger Pliocene species. Oelrich (1958), in describing a new Pleistocene species related to the carolina group (Uanensis from Kansas), suggests that formosa, in- noxia, and singletoni may be valid, but gives no evidence for the state- ment. Milstead (1956) agrees with Barbour and Stetson as to the synony- my of T. innoxia, f6rmosa, singletoni, and canaliculata. He also places 1958 . AUFFENBERG: TERRAPENE IN FLORIDA 59 ilanensis in'the synonymy of canaliculata and refers a series of shells from Friesenhahn Cave, Bexar County, Texas; to T. canaliculata. Two species described from the Pleistocene of Texas, Terrapene impensa and T. bulverda, are also placed in the synonymy of canaliculata. Al- though the western fossils of the genus are not considered in detail in the present study, it is signifcant to point out that Milstead re-empha- sizes the conclusions of Barbour and Stetson that the only distinctive difference between T. canaticulata and T. carolina lies in the larger size of the former. Milstead's work is certainly the most comprehensive so far in that it takes into account the variability of the Recent form, T. carolina, as well as illustrating for the first time the variability in a series of fossil shells. Even more important is the fact that in comparing the Friesenhahn Cave specimens with a small series from Ingleside, San Patricio County, Texas, and with published measurements of canalicu- lam from Florida, reference is made to intermediacy, a concept which, unfortunately, was not considered by earlier workers. The Ingleside specimens are said to be close to the Florida specimens of canalicu- lata, and the larger number of Friesenhahn Cave specimens intermedi- ate between these and the Recent carolina. ' In the present paper it is hoped that it can be shown that two types of turtles occur in the Florida Pleistocene, rather than only one as Barbour and Stetson[ indicated. Furthermore, there are intermediates betwean the two extremes, rep- resenting two races of the species carolina. Shell shape, sulcus patterns, and scute proportions are all highly variable within Terrapene carolina, as has been bbinted out by a num- ber of herpetologists. In some cases identification of Recent- speci- mens from several areas of the United States by means of these char- » acters beeomes almost impossible. However, regardless of the nomen- clatural problems confronting the herpetologist working with Recent forms, certain extremes can certainly be discerned. Thus, Terrapene c. bauri from central and southern peninsular Florida tends to be a rather small turtle with a high shell, with the highest point posterior to the middle of the carapace. Its shell is usually more triangular in + cross section than any of the other subspecies. The peripherals are not as greatly flared lis in T. c. major. In the latter subspecies the shell is not as gabled, but more flattened dorsoventrally and frequently ,with a depressed area on either side of the middorsal keel (fig. 4). The lateral keel above the bridge is much more obviously developed, and the posterior peripherals are much inore str6ngly flared. In ad- dition, the turtle attains considerably greater size. Though variable, the frst vertebral is usually straight-sided in bauri from extreme south- 60 BULLETIN FLORIDA STATE MUSEUM Vol. 8 ern Florida but is decidedly urn-shaped in specimens of maior. An - axillary scale is rarely absent in specimens of major and rarely present in bauri. Most of these differences appear to be clinal, though the degree of difference per unit distance may be different for each char- acter. The shells of T. c. triunguis and T. c. carolina are intermediate between those of bauri and malor. T. c. triunguis is certainly one of the most variable of the races as regards shell shape. C A B D Figure 4.-Recent specimen of Terrapene carolina major. St. Josephs Island, Gulf County, Florida. (A) dorsal view. (B) lateral view. (C) ventral view of anteriof peripherals, left side, showing axillary scale in contact with the third, fourth, and fifth marginal scutes. (D) posterior view, right half, showing flared posterior peripherals. Most important to the present study is the fact that at least two basic shell types are represented in tbe Pleistocene beds of Florida: (1) a small, high-domed, or even gabled-shelled turtle with peripherals not greatly flared and a well-developed axillary scale lacking, and (2) a much larger type in which the shell is usually depressed dorsoven- trally, peripherals widely Hared, and an axillary scale always well de- veloped. Both of these types are being termed canaliculata at the present time. As far as is presently known, except in deposits where admixture of faunas has deRnitely occurred, either one or the other is found. They are never found together in unmixed deposits. The smaller box turtle found so commonly in the Pleistocene of Florida certainly represents the species carolina. Furthermore, it is closer to T. c. bauri and T. c. carolina than to T. c. malor. It is indis- r p 1958 AUF.FENBERG: TERRAPENE IN FLORIDA 61 tinguishable from bauri and carolina in shape, proportions of the scutes, and in sulcus patterns of both the carapace and plastron. The larger turtle in the Pleistocene of Florida is certainly closer to the Recent subspecies major. The only character by which it can be separated from the modern subspecies is maximum size. The Pleis- tocene form grows .much larger. Terrapene c. carolina, maior, and bauri intergrade at their mutual borders at the present time (table 4). Fossil specimens intermediate between the smaller and larger Pleistocene forms occur at some locali- ties. Whether these are temporal or geographic intermediates is unknown at present. The smaller Pleistocene form, which lacks an axillary scale,, is here' considered synonymous with Terrapene carolina bauri or Terrapene carolina carolina or both, and the larger form in' which an axillary scale is always found is designated by the new nomenclatural combination Terrapene carolina putnami. The morpho- logical characters on which these designations are based are dis- cussed below. MATERIAL EXAMINED In order to evaluate variation in shell shape and sulcus pattern in Pleistocene specimens from Florida, 475 Recent specimens of Terra- pene carolina, including the cotypes of T. c. bauri and T. c. major, were examined. for particular characters. Specimens of T. carolina examined come from the following states: F16rida 165, Georgia 81, Tennessee 78, Arkansas 12, Louisiana 5, Texas 17, Illinois 6, Missouri 12, Alabama 22, South Carolina 4, Mississippi 8, North Carolina 8, Vir- ginia Il, Maryland 11, New Jersey 6, West Virginia 3, Ohio 13, New York 9, Pennsylvania 8, Massachusetts 8. Of the specimens from ex- treme southeastern United States, 54 can be termed "good" major, pos- sessing all of the diagnostic characters preseiitly used to separate this form from the others. Most of these specimens come from the general region of the lower reaches of the Apalachicola River in Florida. Specimens termed "good" bat*ri, in view of the fact that they possess ' all of the diagnostic characters of this race, total 59. These all come from peninsula Florida south of Marion County. The remaining 52 specimens from Florida are intermediate in at least some of their char- acters; intermediate between bauri and mator, bauri and carolina, or major and carolina. These specimens come from the northern tier of counties of Florida southward on the peninsula to include Marion County. The study of Recent Terrapene carolina is necessarily Florida- centered for two reasons. Specimens from this area are more com- 62 . BULLETIN FLORIDA STATE MUSEUM Vol. 8 mon in collections than from other southeastern states, and data from the fossil box turtles of Florida indicate that populations in and near Florida today are the most important from a comparative standpoint. In addition to Recent specimens of Terrapene carolina, 27 speci- mens of T. ornata from various parts of its range were also examined. One of the paratypes of T. coahuila (MCZ 53981), several individuals of T. mexicana (MCZ 4041, 4997, 9512, 56082, 56050), and one T. klauberi (MCZ 46855) were studied with regard to particular shell characters. The study of fossil box turtles fi~om Florida is based largely on 84 complete or nearly complete shells in which at least one entire half of the carapace is available. There are a few exceptions, namely, type specimens and Terrapene formosa in which the anterior quarter of the carapace is missing. In addition, a'large number of fragments, par- ticularly of the anterior and posterior peripherals were studied from several deposits. C A _ B D Figure 5.-Specimen referred to Terrapene carolina carolina, UF 9978, Arre- dondo I C, Alachua County, Florida, Pleistocene. (A) dorsal view. (B) lateral view. (C) ventral view of anterior peripherals, left side, showing absence of aullary scale. (D). Dosterior view, right half, showing n6nfiared posterior periph- erals. It is indeed fortunate that a Bne series of 16 complete or almost complete shells are available from one Pleistocene locality-a quarry of the Dixie Lime Products Company, approximately one-mile south of Reddick, Marion County, Florida. Most of these are in the Uni- versity of Florida Collections, although several are in the collections of the Florida Geological Survey. This series of shells constitutes the 1958 AUFFENBERG: TERRAPENE IN FLORIDA ,68 largest yet assembled of Terrapene from any single Pleistocene locality in North America. A specimen from Afredondo I, Locality C, Alachua County, Flor- ida (UP' 9978), is a particularly fortunate Bnd since it is provided with a perfect skull attached to the inner surface of the anterior 16be of the plastron (figs. 5 and 8) A small series of complete and partial shells is also available from the Melbourne area. These are deposited in the collections of Amherst College, the Museum of Comparative Zoology, and the United States National Museum. Single shells of less importance, which are com- plete or nearly complete, are available from several localities and are discussed below. In addition to the excellent specimens mentioned above, 106 non- associated anterior and posterior plastral lobes are available 'from the Pleistocene deposits of Florida, as are hundreds of fragmental pieces of carapace and some partial skulls. Some of the more noteworthy of these are discussed below. ANALYSIS AND DISCUSSION OF THE GENUS TERRAPENE The name canaliculata is now applied to all Pleistocene box turtle f remains from Florida. It has been applied to several populations out- side the state as well. This form in the restricted sense is best de= scribed by Gilmore (1927). He redefines the species on the basis of newer, more eomplete material .than was.previously available. He "outlines the main distinguishing features of this species as,... large size; shell thick and heavy; carapace with its free borders curved up- wards; posterior peripherals widely flaring; keel over the bridges con- necting free borders of the front and back peripherals; gutter-like groove above this keel; first vertebral strongly urn-shaped with portion posterior to the middle as wide but usually wider than the anterior end; strongly elevated median longitudinal ridge, extending full length of ~rst vertebral; median... keel [not] interrupted by transverse sulci." All of the characters mentioned in the description above, as well as several others, have been examined to determine their variability in Recent specimens of cdrolina, as well as in the better and more abun- dant fossil material now available from Florida. One character of considerable interest, and never before mentioned by workers in this group, is the presence of a well-developed axillary scale in shells belonging to the large canaliculata-type turtles in Flor- ida , Recent specimens of Te~rapene c, major almost always possess a well-developed axillary scale in contact with the third and fourth; 64 ' BULLETIN FLORIDA STATE MUSEUM Vol. 3 or third, fourth, and fifth marginal scales. Color, size, and shell shape are the main characters used to diagnose this subspecies so far. Ex- amination of specimens of T. carolina from outside the range of major shows some variation in the development of the axillary scale. Rarely it is broken into a series of small scales. A similar abnormality occurs in other genera of turtles (Zangerl and J6hnson, 1957). When present as a single structure in Terrapene it varies from a large obvious scale to one of small size. In some it is totally absent. Fbr purposes of this study variation is simply tabulated'as a presence or absence of the scale. If present, the marginal scutes with ·which the axillary scale comes in contact is noted. No attempt is made to evaluate individual - if .-- \ 1 1 F *.to E/ , 15 ' too- 0 LEVY ALACHUA OUVAL 90· COUNTIES ,80 A PE R C EN T SP EC IM EN S W IT H AX IL LA R Y SC AL E 70· 3* 60, 50 40 D ' 30 B 10· D 0 GULF WAKULLA LAFAYETTE ALACHUA POLK DADE COUNTIES Figure 6.-Geographic variation in the percentage of specimens in various populations in, Florida that possess axillary scales. The representation is a gen- eralization and is meant to illustrate only major trends in this clinal type varia- tion. Variations among small localized inter-breeding populations in the same general . area are not of sufficient magnitude to alter the broad trends indicated here. luGo AUFFENBERG: TERRAPENE IN FLORIDA 65 variation in its size since the material at hand indicates that such a study would in no way alter the general conclusions attained by means 2 of the simpler method. t - ABCD Figure 7.-Variation in development of the axillary scale in Terrapene caro- - lina putnami. (A) UF 9928, Seminole Field, Pinellas County, Florida. (B) UF 2258, Bone Cave, Citrus County, Florida. (C) UF 2862, Slst St. locality, Braden. ton, Manatee County, Florida. ' (D) UF 2211, Haile VII A, Alachua County, Florida. Ali specimens Pleistocene. The axillary scale is absent in approximately two thirds of the spec- imens of Terrapene c. carolina from the eastern seaboard. It is absent in populations of bauri from southern Florida. It is variable in speci- mens of triunguis and c. carolina from the middle and western parts of the species range, but is never as frequent in c. carolina as in major. The specimens from northern and central Florida are intermediate as regards the number of individuals possessing the scale. Furthermore, a cline is suggested in Florida in which the scale becomes less frequent in populations from more southern areas of · the state. Populations from the western half of the peninsula have the scale more frequently than do those from the eastern half. Presumably this'is due to the in- fluence of maior existing in western Florida. Within the range of major in the pailhandle of the state, the populations which most fre- - quently have the scale present occur nearest the Gulf coast. Figure 6 illustrates variation in this character in Recent individuals of the species. Figure 7, illustrates variation in· the. size and shape of. this 66 BULLETIN FLORIDA STATE MUSEUM Vol. 8 scale in several shells of the large Pleistocene box turtle called canalicu- lata in the restricted sense of Hay (1907, and later) and Gilmore (1927).2 An axillary scale is present in the single Terrapene coahuila and T. klauberi examined. It is present in two of the six individuals of T. mexicana examined. No skull of T. canaliculata (sensu stricto) has yet been found. How- ever, a fossil 6f a carolina bauri-type animal (UF 9973) from the Pldisto- cene deposits at Arredondo I, Locality C, Alachua County, Florida, possesses a Ene skull. The associated shell is rather small (128 mm. in length), the peripherals are only slightly flared, the first vertebral is straight-sided, not um-shaped, and the area above the bridge lacks a keel. Unlike T. c. maior and most specimens· of bauri, the quadrato- jugular portion of the postorbital bar is completely lacking, though this element may have dropped out while the specimen was rotting. (It frequently does this in macerating specimens of bauri because- this element is somewhat loosely attached to its neighbors.) The presence or absence of the bar is apparently variable, at least in T. carolina. Unfortunately, the situation is much more complex than that indicated by Zangerl (1948a). In the fossil skull from Arredondo the postorbital is narrow and high as in T. c. tliunguis and carolina, and a more robust - element is found in T. c. bauri and inaior. The squamosal is relatively long and more like that in carolina and major than in triunguis and bauri. The posterior palatine foramina are small, and thus similar to those found in all of the North American subspecies with the exception of major, where they are usually rather large (fig. 8). As a unit the skull seems closest to that usually found in populations of T. c . CaTo- lina. The shell, because it is so high in proportion to its length, is more nearly like that typical of T. c. bauri (figs. 5 and 9), but is cer- tainly encountered in some carolina specimens as well. The fossil shell lacks the axillary scale found in canaliculata-type specimens and almost all Recent T. c. maior. Of the fossil forms described from Florida, it is obviously close to T. innoxia Hay and T. formosa Hay (figs. 1 and 2) The characters utilized by Gilmore in redefining Terrapene canalic- ulata are discussed in detail below, with particular emphasis being placed on variati6n in both Recent and fossil samples from Florida. ' A similar pattern of geographic variation has been shown.in the snake Coluber constrictor (Auffenberg, 1955). In the absence of any direct evidence bearing on the origip of such a geographic pattern it is quite reasonable to infer- that 'its basis rests in dispersal patterns of particular organisms during the Pleisto- cene. 1958 AUFFENBERG: TERRAPEN£ IN FLORIDA 67 Differences between the recent 'subspecies bauri and mafor are stressed in order to illustrate parallel differences between the two Pleistocene forms on the one hand represented by diinaliculata from Melbourbe, and on the other by the Arredondo specimen mentioned above as well as the type's of for,nosa and innoxia. A B C D E F 7«- - v G \ 1 Figure 8.-Variation in the development of the postorbital bar in Recent and Pleistocene specimens of Terrapene carolina. Dotted Iines represent reconstructed parts. (A, B; and C) Recent specimens of T. c. bauri, showing extremes of de- velopment; ali from Gainesville; Alachua County; Florida. Drawing C repre- sents the typical condition found in this part of the state. (D) Recent adult T. c. mator, St. Josephs Island, Gulf County, Florida. (E) Recent juvenile T. c. maior, 5 mi. W Apalachicola, Franklin County, Florida, ,(F) Recent T. c. caro- lina, Massachusetts. (G) UF 9978, Pleistocene, Arfedondo I C, Alachua County, Florida, referred to T. c. carolina. (H) UF 9988, Reddick I B, Marion County, Florida, Pleistocene, referred to T. c. Pcarolina XT. c. putnami. (I) Recent T. c. triunguis, (after Zangerl, 1948b). SIZE.-This has been one of the characters most persistently used not only in describing several fossil forms from Florida and Texas but also in separating canaliculata from carolina (Milstead, 1956; Bar- bour and Stetson, 1981). Size is a character of restricted diagnostic value when dealing with closely related reptiles. However, the several recognized sub- ' species of T. carolina show considerable differences in maximum size, as well as average size. Thus, T. c. maior definitely attains a greater size than does carolina or triunguis. T. c. bauri is the smallest sub- species of the group. Maximum and average sizes of the samples of Recent specimens of T. c. bauri, malor, and carolina are given in table 1. The available sample of triunguis is much too small to be signifi- 68 BULLETIN FLORIDA STATE MUSEUM Vol. 3 cant. Furthermore, the more western range of this form suggests that it need nbt be considered seriously in comparing Recent forms of carolina with the fossils from Florida. I C d l - A B D Figure 9.-Recent shall of T. c. bauri, Gainesville, Alachua County, Florida. (A) dorsal view showing dendritic scale abnormalities caused by fire, and an extra vertebral s€ute. (B) lateral view. (C) ventral view of anterior peripherals, left side, showing absence of axillary scale. (D) posterior view, right side, showing nonfiared peripherals. Similar measurements for the fossil shells from Florida are given iii table 2. The entire sample is divided into three categories; the fine series of shells from a single locality at Reddick, Marion County, Flor- ida, the small bauri-like shells from several localities, and the available ~ specimens clearly referable,to T. canaliculata (sensu stricto). The specimens of canaliculata (sensu stricto) are obviously much larger than any of the Recent forms. Their size is approached most closely by Terrapene c. major, less so by the Reddick sample. The latter is intermediate between the Recent mai'or and carolina. The bauri-type shells from the remaining fossil localities in Florida are essentially identical to T. c. bauri in size. The smallest living box turtles in Florida are found at the ex- treme southern end of the peninsula. Individuals from the Keys seem ' to be somewhat larger, approaching those of northern peninsular Florida in shell length. They also possess a flatter shell than those from the adjacent mainland, and the peripherals are more widely Bared. The largest living box turtles in North America are found along the Gulf coast from Bay County to Wakulla County, Florida. 1958 AUFFENBERG: TERRAPENE IN FLORIDA 69 Two specimens from St. Vincents Island are the largest box turtles I have measured (210 and 216 mm. respectively before the scutes were removed in preparing their skeletons). Northward from this coast the average carapace length· becomes progressively less, so that at the Georgia border the turtles are only slightly larger than typical speci- mens of the subspecies carolina over the remainder of its range. Mil- stead (1956) was unaware that' major attain so large a size. The largest Recent specimen of Terrapene that he measured is 162 mm. long. The largest fossil individual he measured is 208 mm., although fragments indicate that some specimens have attained lengths of from 240 to 260 mm. Living maior from a restricted area along.the Gulf coast attain a greater size than the majority of specimens from Friesen- hahn Cave referred to canaliculata., It is obvious that if size is the onlj' difference between Recent carolina and the Friesenhahn Cave ~ sample, the reference of that fossil series to an extinct species seems 180 160· OO LE NG TH O F PO ST ER IO R LO BE O F 140· cp» P LA S TR O N (I N M M .) 120- V A 08• 100. 19008 0 60 . 0 1%. * %0' Ip//40 -$ 60 100 150 200 250 300 350 CARAPACE LENGTH (IN MM.) Figure 10.-Correlation of carapace length against hypoplastral length in Recent and fossil Terrapen€ carolina. Solid circles represent actual measurements; hollow circles represent specimens in which only the hypoplastron is available and the carapace length is estimated. The size ranges for both fossil and Recent nomenelaturally distinct races are indicated. 70 BULLETIN FLORIDA STATE MUSEUM Vol. 8 unwarranted, especially in view of the considerable variation demon- strated in living material from Florida. However, the Florida speci- mens of canaliculata (sensu stricto) considerably exceed in size the largest Recent maioT measured so far . Partial plastra are by far much more abundant than complete carapaces. Of the plastral elements available from Florida, the poste- rior lobe is preserved complete most frequently. The posterior lobes of all Recent and fossil individuals were measured if a definite associ- ation with a particular carapace was possible. These measurements are plotted against carapace length (fig. 10). It then becomes possible to estimate carapace lengths of individuals represented only by the posterior lobe of the plastron, thus making it possible to obtain a bet- ter idea of size distribution in the various fossil populations than could be gained solely from the complete specimens now available. The results of these estimates are presented in table 3. It is obvious that two turtles can be distinguished in the Florida Pleistocene on size alone. Of particular interest is the fact that the type,of Terrapene putnami, described by Hay on the basis of its larger size, is not unduly larger than canaliculata (see fig. 10). A second fragment referred to putnami by Hay (1908b) is estimated to have originated from a specimen 820 mm. in length. There is every reason for believing that putnami bears the same relation to canaliculata (sensu stricto) as the latter does to T. c. maior, and as maior does to carolina, that is; a continuous trend toward increasing size from bauri on .the one hand, thr6ugh carolina. maior, canaliculata (sensu stricto), to putnumi. As members of a continuous series that can be separated by no other means, it seems unreasonable to assume that the relation- ship of all of these f6rms to one another is any more than a sub- specific one, if indeed one should go even that far on size alone. T. c. mator from the coastal areas of northwestern Florida, Alabama, Mississippi, and Louisiana is clearly differentiated from other nominal races of the same species on several characters other.than Size. This is also true of T. c. bauri. The fossil forms, canaliculata and putnami Hay 1907, are indistinguishable. T. putnami is considered to be sub- specifically related to T. carolina, and of all the forms in this latter species, it is closest to T. c. mator. The name T...carolina putnami as used below thus refers to the large, Rated-shelled, Pleistocene box tur- tles of Florida possessing an axillary scale. SHELL THICKNESS.-Shell thidkness is related to total size, and by itself a poor character with which to separate populations of Recent or fossil turtles. As a diagnostic tool it is valueless except as a cri- 1958 AUFFENBERG: TERRAPENE IN FLORIDA 71 TABLE 1 ~TANDARD CARAPACE LENGTH (IN MILLIMETERS) OF RECENT SPECIACENS 0* Terrapene carolina - Subspecies Mean Maximum Number major 178 216 46 bauri 133 158 44 carolina ' 156 168 101 TABLE 2 STANDARD CARAPACE LENGTH present, at 86 BULLETIN FLORIDA STATE MUSEUM Vol. 3 least in part, by a geographically somewhat restricted intermediate population, T. c. major. A tentative arrangement is presented in table 4 as a speculative correlation of Pleistocene vertebrate-bearing strata; this is based on taxonomic determinations of fossil Terrapene populations from Florida and the available stratigraphic data. Table 5 summarizes the taxo- nomic determinations made from complete or fragmentary shells of Terrapene for all of the deposits from which box turtles are now known in Florida. - T.C. CAROLINA P T, C, CAROLIN~ « T. C. PUTNAIAI A B \F~-0~T.C. BAURI F ..0...- T' 1 £ T. C. CAROLINA 4 -11- -Ze . 1 C ~~ *.~FT.C. BAURI D 4 T C BAURI, 1 l. ~4*4 Figure 15.-Major' distributional stages in the evolution and dispersal of Terrapene carolina in southeastern United States. (A) movement of a large, prim- itive stock (T. c. putnami) eastward along a circumferential Gulf coast marsh route sometime in the early Pleistocene. (B) hypothetical stage during some interglacial period when sea level was much higher than at present. (C) hypo- thetical stage during some later. glacial period when sea level was much lower than at present. (D) present distribution of the subspecies. T. c. mafor is assumed to represent a relict intergrade population between T. c. carolina to the north 'and Pleistocene T. c. putnami to the south. Stippled areas represent areas of intergradation. , 1958 AUFFENBERG: TERRAPENE IN FLORIDA 87 TABLE 4 TENTATIVE CORRELATION OF Fossm Terrapene LOCALITIES IN FLORIbA Time Bauri Bauri X Putnami Putnami - Vero Bea6h, stratum 8 Seminole (in part) RECENT . Wekiva Springs Itchatucknee River (in part) Bone Cave (in part) PLEISTOCENE - Seminole Field ' 8 mi: W Melbourne (in part) LaBelle Eichleberger Cave Sarasota A and B Venice Wisconsin "Uranium" Cave Vero Beach, stratum 8 maximum h . Mefford Cave I 8 mi. W Melbourne < Waccasassa River Kendrick I near St. Augustine Itchatucknee River 3 mi. W Melbourne (in part) 10 mi. W Melbourne Three Rivers Devel- Alafa River , opment Winter Beach Bradenton Field f Itchatucknee RiverLecanto Cave ~ (in part) Sangamon maximum · h « Bone Cave (in part) Mefford Cave II Reddick II Arredondo IA Paynes Prairie B Haile VII A (in part) Apopka Creek Reddick I B < ·Zuber I Williston Arrendondo I C; II Wall Company Pit Haile VII A (in part) ' Haile VII B Illinoian maximum h 88 BULLETIN FLORIDA STATE MUSEUM vol. 8, TABLE 5 DETERMINATIONS OF FOSSIL Terrapene FROM THE PLEISTOCENE BEDS OF GEORGIA AND FLORIDA Subspecies of Locality Terrapene carolina Remarks GEORGIA Skedaway or Whitemarsh putnami Types of T. canaliculata, Islands USNM 5500 FLORIDA Alachua County Arredondo I„ A putnami UF 2429 Arredondo I, B Terrapene sp'. UF 2045 Arredondo I, C Pcarolina A complete shell and skull, UF 9978 Arredondo II Pcarolina UF 2094 Wall Co. Pit carolina or bauri UF 2017, 5028-4, 5028 Paynes Prairie B putnami UF 6408 Haile VII, A putnami UF 2211 . carolina or bauri Complete shell and plastron. Private collection; not available for further study. Also FGS uncataloged and UF 1790, 9986. Haile VII, B ~ carolina or bauri FGS'V-5708 Brevard County 8 mi. W Melbourne ptitnami USNM 11428 (Gold Course loc.) AMC 25-144, 25-145 C MCZ 1024-1028 putnami X Pbauri Type of T. singletoni, USNM 11181 8 mi. W Melbourne bauri UF 1586 10 mi. W Melbourne putnami UF 1594 Citrus County Bone Cave putnami Large part of shell, UF 2258 Large pieces of shells, plas- tra, etc., UF 2257, 2616, 2625, 6520 bauri - Anterior parts of carapaces, numerous plastra, UF 2622, 2623 Lecanto Cave putnami X Pbauri FGS uncataloged. (Saber Tooth) [continued] 1958 AWFFENBERG: TERRAPENE IN FLORIDA 89 TABLE 5-(continued) DETERMINATIONS OF FOSSIL Terrapene FROM THE PLEISTOCENE BEDS OF GEORGIA AND FLORIDA Subspecies of Locality Terrapene carolina Remarks Columbia County Itchatucknee River carolina or baud UF 1986 putnemi UF 1616 Three Rivers Develop- carolina or bauri UF 16I1 ment DeSoto County Prairia Creek Terrapene sp. UF 6618 Hardy County Near LaBelle putnami UF 5817 Hillsborough County Alafa River putnami ' Type of T. putnami, AMNH 6097 Indian River County Winter Beach Locality .Pputnami UF 1865 Vero Beach, stratum 2 bauri Type of T. innoxia, USNM 8824 Also FGS V-5471, 7085 Vero Beach, stratum 8 putnami Type of T. antipex, USNM 8820 Also FGS V-4485, 5478, 5480,5255 Levy County Wekiva Springs bauri UF 2588 Waccasassa River putnami UF 4267 Williston (Holman carolina X putnami FGS uncataloged locality) Manatee County Bradenton, 51st Street putnami Large parts of earapaces and locality plastra, UF 2809,2362, 2663, 9862 Marion County "Ocala" carolina or bauri Type of T. formosa, USNM 8825 Zuber I ?carolina X putriami Entire rear of shell, UF 5019 Reddick I, B Pcarolina X putnami Entire, or large parts of shells, UF 1479, 2059, 2179,2388, 2552, 4747-a, -b, -c, 5697, 5699, 5700, 6137, 6415, 6422, 6612 FCS V-5709 [continued] , 90 BULLETIN FLORIDA STATE MUSEUM Vol. 3 TABLE 5-(continued) DETERMINATIONS OF FOSSIL Terrapene FROM THE PLEISTOCENE BEDS OF ~EORGIA AND FLORIDA Subspecies of Locality Terrapene carolina Remarks Plus many plastra, skull pieces, etc. Reddick II putnathi Almost complete shell, UF 9972, and large ~ piece of shell, UF 9918 Eichleberger Cave A Pbauri UF 1476-9 Eichleberger Cave B Pbauri UF 1658 Pbauri UF 1458Uranium Cave, 2 mi. north Bellview Mefford Cave I carolina or bauri UF 2115 Mefford Cave II putnami UF 6521-2 ~ Kendrick I carolina or bauri UF 2650 Orange County Apopka Creek putnami FGS V-1694 Pinellas County ~ "Seminole Field" putnami UF 9928 Sarasota County' Near Sarasota Fitnami UF »1054 St. Johns Couhty 28 mi. S St. Augustine putnami Uncataloged, Allen collec- tion (Hay 1916) 20 mi. N St: Augustine Putnami FGS V-4485 LITERATURE CITED AufFenberg, W. 1955. A reconsideration of the raeer, Coluber constrictor, in eastern United States. Tulane Studies Zo61., vol. 2, no. 6, pp 89-155. 1956. Additional records of Pleistocene lizards from Florida. Quart, Jour. Florida Adad. Sci., vol. 19, nos. 2-5, pp 157-167. [MS.] The fossil snakes of Florida. Thesis, submitted · to University of Florida, 1956. 128 pp. Bader, R. 1957. Two Pleistocene mammaIian faunas from Alacliua Couhty, Florida. Bull. Florida State Mus., vol. 2, no. 5, pp. 58-75. Barbour, T., and H. C. Stetson 1931. A revision of the Pleistocene species of Terrapene of Florida. Bull. Mus. Comp, Zo61., vol. 72, no. 8, pp 295-299. 1958 AUFFENBERG: TERRAPENE IN FLORIDA 91 Brodkorb, P. 1957. New passerine birds from the Pleistocene of Reddick, Florida. Jour. Paleontol., vol. 81, no. 1, pp. 129-188. Carr, A. F., Jr. 1952. Handbook of turtles. Ithaca, New York: Cornell Univ. Press; pp. i-xviii, 1-542. Cooke, C. W. 1945. Geology of Florida. Florida State Geol. Surv., Geol. Bull; no. 29, pp. 1-839. Cupe, E. D. 1869: Synopsis of extinct Batrachia, Reptilia and Aves of North America. Trans, Amen Phil, Soc., vol. 14, pp. 1-124, Gilmore, C. W. 1927. On fossil turtles from the Pleistocene of Florida. Proc. U.S. Natl. Mus., vol. 71, art. 15, pp. 1-10. Hay, 0. P. 1907. Descriptions of seven new species of . turtles from the Tertiary of the . United States. BulL Amer. Mus. Nat. Hist:, vol. 28, art. 84, pp. 847-863. 1908a. Descriptions of five species of North American fossil turtles, four of which are new. Proc. U.S. Natl. Mus., vol. 35, no. 1640, pp. 161-169. 1908b. The fossil turtles of, North America. Carnegie Inst. Washington, publ. no. 75, pp. 1-568. 1916. Descriptions of some Floridian fossil vertebrates belonging mostly to the Pleistocene. Florida State Geol. Surv., 8th Ann. Rept., pp. 89-76. 1928. The Pleistocene of North America and its vertebrate animals from the states east of the Mississippi River, and from the Canadian Provinces east of longitude 95". Carnegie In5t-. Washington, pubI. no. 822, pp. 1-405. ' McConkey, E. H. 1957. The subspecies of Eumeces egregius, a lizard of the southeastern United States. Bull. Florida State Mus., vol. 2, no. 2, pp. 18-28. Milstead, W. W. 1956. Fossil turtles of Friesenhahn Cave, Texas, with the description of a new species of Testudo. Copeia, 1956, no. 8, pp 162-171. Neill, W. T. 1957. Historical biogeography of present-day Florida. Bull. Florida State Mus., vol. 2, no. 7, pp. 175-220. Oelrich, T. M. 1958. A new box turtle from the Pleistocene of southwestern Kansas, Copeia, 1958, no. 1, pp. 83-88. Rouse, I. 1951. A survey of Indian River archeology. Yale University Publ., Anthropol., no. 44, pA 1-296. Wood, H. E., et al. 1941. Nomenclature.,and correlation of the North American continental Tei- tiary. Bull. Geol. Soc. Amen, vol. 52, no. 1, pp. 1-48. 92 BULLETIN FLORIDA STATE MUSEUM Vol. 8 Zangerl, R. 1948a. The vertebrate fauna of the Selma formation of Alabama. iI. The pleurodiran turtles. Fieldiana, Geology Memoirs, vol. 3, pp. 28-56. 1948b. The methods of comparative anatomy and its contribution to the study of evolution, Evolution, vol. 2, no, 4, pp. 851-374. Zangerl, R., and R. G. Johnson 1957. The nature of shell abnormalities in the turtle shell. Fieldiana, Geology, vol. 10, no. 29, pp. 841-862. Contributions to the· BULLETIN OF THE FLORIDA STATE MUSEUM may be in any Reid of biology. Manuscripts dealing with natural history or systematic problems involving the. southeastefn United States or the Caribbean area are solicited especially. Manuscripts,should be of medium length-12 to 200 printed pages. Examination for suitability is made by an Editorial Board. The BULLETIN is distributed worldwide thr6ugh institutional subscriptions and exchanges Only. 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