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fill ^&y*t*\*ol\o^\*:^^\ auS^ s^eW-N. C. DOCUMENTS CLEARINGHOUSE AUG 10 1993 N.C. STATE LBRARY RALOGH number 18 June 1993 EDITORIAL STAFF Richard A. Lancia, Editor Suzanne A. Fischer, Assistant Editor Eloise F. Potter, Production Manager EDITORIAL BOARD James W. Hardin Rowland M. Shelley Professor ofBotany Curator ofInvertebrates North Carolina State University North Carolina State Museum ofNatural Sciences William M. Palmer Robert G. Wolk Director ofResearch and Collections Director ofPrograms North Carolina State Museum North Carolina State Museum ofNatural Sciences ofNatural Sciences Brimleyana, the Zoological Journal of the North Carolina State Museum of Natural Sciences, appears twice yearly in consecutively numbered issues. Subject matter focuses on systematics, evolution, zoogeography, ecology, behavior, and paleozoology in the southeastern United States. Papers stress the results of original empirical field studies, but synthesizing reviews and papers of significant historical interest to southeastern zoology are also included. Brief communications are accepted. All manuscripts are peer reviewed by specialists in the Southeast and elsewhere; final acceptability is determined by the Editor. Address manuscripts and related correspondence to Editor, Brimleyana, North Carolina State Museum of Natural Sciences, P.O. Box 27647, Raleigh, NC 27611. Information for contributors appears in the inside back cover. Address correspondence pertaining to subscriptions, back issues, and exchanges to Brimleyana Secretary, North Carolina State Museum of Natural Sciences, P.O. Box 27647, Raleigh, NC 27611. In citations please use the full name - Brimleyana. North Carolina State Museum of Natural Sciences Betsy Bennett, Director North Carolina Department of Agriculture James A. Graham, Commissioner CODN BRIMD 7 ISSN 0193-4406 The Myriapod Types of Oscar Harger (Arthropoda: Diplopoda, Chilopoda) Rowland M. Shelley North Carolina State Museum ofNatural Sciences, P.O. Box 27647, Raleigh, North Carolina 27611 ABSTRACT—The type specimens of all five milliped species — Trichopetalum lunatum, T. glomeratum, T. iuloides, lulus furcifer, and Polydesmus armatus—and one of the two centipedes, Lithobius pinetorum, authored by Oscar Harger in his only paper on myri-apods and previously thought to be lost, are housed at the Peabody Museum of Natural History, Yale University, New Haven, Connecti-cut. From our knowledge of the itinerary of the Yale paleontologi-cal expedition of 1871, we know the type locality of T. glomeratum, I. furcifer, P. armatus, and L. pinetorum, previously stated as the "John Day River Valley, Oregon" is restricted to the vicinity of Canyon City, Grant County, on the western slope of the Blue Mount-ains. The female holotype confirms that T. glomeratum is a repre-sentative of the chordeumatoid family Conotylidae, and the name is assigned provisionally to Taiyutyla pending collection of a male topotype. Unidentifiable female conotylids are also reported from another area in eastern Oregon and the Snake Mountains in eastern Nevada, which suggests that the family is widespread in montane forests at high elevations in the generally arid Columbia Plateau and Basin and Range Physiographic Provinces. To facilitate future studies, I provide gonopod drawings of male syntypes for /. furcifer and P. armatus. One of the more obscure authors of North American myriapods is Oscar Harger (1843-87), whose sole publication on these arthropods (Harger 1872) described the milliped genus Trichopetalum and seven species, two centipedes {Lithobius pinetorum and Geophilus gracilis) and five millipeds {Trichopetalum lunatum, T. glomeratum, T. iuloides, lulus furcifer, and Polydesmus armatus). Born at Oxford, Connecticut, Harger attended the Connecticut Literary Institute at Suffield and Yale College, graduating from the latter with honors in 1868 (Schuchert and LeVene 1940). After briefly studying zoology under Professor A. E. Verrill, Harger became the first assistant to the vertebrate paleontologist, O. C. Marsh, partici-pating on the latter's expeditions into the American West in 1871 and 1873. From July to September 1872, Harger dredged marine organisms on a Coast Survey steamer with Professors Verrill and Sydney I. Smith, Yale's first professor of Comparative Anatomy, who earlier had been naturalist to the U.S. Lake Survey and collected the types Brimleyana 18:1-13, June 1993 1 2 Rowland M. Shelley of T. iuloides. Harger was studious and an active reader, and Marsh valued his scientific opinions in paleontology. However, Marsh would not allow Harger to publish on vertebrate fossils, either alone or jointly with him, so Harger 's only papers are on invertebrates—that of 1872 on myriapods, two on isopods, and one on a fossil spider. From 1870 to 1873, Marsh led four vertebrate paleontological expeditions of Yale students and recent graduates into the West (Schuchert and LeVene 1940). The idea of such efforts arose from preliminary explorations he made on a trip to the end of the trans-continental railroad in Wyoming in August 1868 after attending a scientific meeting in Chicago. The 1871 expedition traveled to regions of Kansas, Wyoming, and Utah, where Harger collected 10 fossil species. The group then rested a few days in Salt Lake City with Brigham Young while Marsh prepared to explore a new area, the John Day River Basin in central Oregon. After traveling 12 days by rail and stage, the party crossed the Blue Mountains and arrived at Canyon City, Oregon, on the John Day River on 17 October 1871, where it waited several days for a military escort from Fort Harney, 75 mi (120 km) to the south. The group collected fossils from 31 October to 8 November in the John Day region before traveling down the Columbia River to Portland; it then traveled to San Francisco and returned east directly by rail or by boat via Panama. While the expedition was in the John Day River area, Harger, or Harger and Professor G. H. Collier, collected four Oregon myriapods that he described in 1872 — L. pinetorum, T. glomeratum, I. furcifer, and P. armatus. Both the publication and labels in the vials give the locality as just the "John Day River Valley," but knowledge of the group's activities enabled me to infer a more precise site. The John Day River arises on the western slope of the Blue Mountains in Grant and Umatilla counties, flows westward into Wheeler County, then heads northward to the Columbia River forming the boundaries between Wheeler/Wasco and Sherman/Gilliam counties. It is not to be confused with Days Creek, Douglas County, in the Umpqua River drainage of southwestern Oregon, the probable type locality for Zantona douglasia Chamberlin and Bollmanella oregona Chamberlin (Shear 1974, Gardner and Shelley 1989), which Chamberlin (1941a) misnamed as "John Day Creek." Because most millipeds require moist leaf litter and much of the John Day Basin is in the arid rain shadow of the Cascade Mountains, I (Shelley 1990) speculated that the site was probably near the confluence of the John Day and Columbia rivers in either Sherman or Gilliam county. However, as all the myriapods were collected in October 1871, and the expedition reached Canyon City on 17 October and only collected fossils from 31 October to 8 November Myriapod Types of Oscar Harger 3 after waiting for the military escort, it is evident that during most of the part of October that the group was in the John Day Valley, it was resting in Canyon City. Consequently, there was ample time for relaxed explorations in the vicinity of Canyon City, and I, therefore, believe that Harger's myriapods were collected near this town. Because Harger's paper specifies that L. pinetorum, I. furcifer, and P. armatus were collected by Professor Collier and himself, and T. glomeratum was taken by Harger alone, collecting probably occurred on at least two different dates, as one day Harger went out alone and the other he was accompanied by Collier. There could be as few as one site and as many as four, but further specification is not possible with what we know now. Consequently, the type locality for all of Harger's Oregon species is restricted to the vicinity of Canyon City, Grant County, on the western slope of the Blue Mountains. Harger's centipedes have received little attention since their description. They were included in the catalog of North American myriapods by Bollman (1893), who noted that G. gracilis Harger, 1872, was preoccupied by G. gracilis Meinert, 1870, proposed for a European geophilomorph. Cook and Collins (1891) remarked that Harger's description of G. gracilis conformed very closely to Schendyla nemorensis (C. L. Koch, 1837), and the former is now regarded as a junior synonym (Crabill 1953, 1961). Stuxberg (1875) included L. pinetorum in his list of North American lithobiids, but he had no personal knowledge of the species. Kevan (1983a) listed both species as potential inhabitants of Canada, recognizing the synonymy of G. gracilis under S. nemorensis. In contrast to the centipeds, Harger's millipeds have been cited in a number of publications, but the type specimens were thought to be lost. Chamberlin and Hoffman (1958) stated that their "present location [was] unknown" or that they were "not known to exist," and similarly, Shear (1971, 1972) said that the holotypes of T. glomeratum and T. iuloides were lost and that the whereabouts of that of T. lunatum was unknown. Causey (1967) guessed right when she stated that the holotype of T. lunatum was at the "Peabody Museum of Natural History, Yale University, if extant," but evidently she made no inquiries to confirm this supposition. While recently visiting the Peabody's Museum's collection, I unexpectedly discovered these types in the myriapod cabinet, where they have languished in obscurity for 120 years. A few vials were still capped with wax and had not been touched for decades. The types of P. armatus were in the general collection and not labeled as such, but those of the other millipeds were clearly marked as types and grouped in a clamp-top jar. A concerted search failed to reveal the types of G. gracilis, which ap- 4 Rowland M. Shelley parently are lost, but those of L. pinetorum were in an individual vial and clearly labeled. The sample consists of 12 nearly legless syntypes, seven males and five females, and is number 2175; according to the label it was collected by Harger alone, whereas the published account states that it was collected by him and Professor Collier. All the millipeds are listed in the continental checklist (Chamberlin and Hoffman 1958), and detailed accounts of those Harger assigned to Trichopetalum have recently appeared (Palmen 1952; Shear 1971, 1982; Shelley 1988, In Press). In the following accounts I update these reports by providing information on the type specimens, a brief historical review of each species, and pertinent anatomical observations. Complete synonymies are presented, and each species is placed in its proper order and family. Chordeumatida: Trichopetalidae Trichopetalum lunatum Harger Trichopetalum lunatum Harger, 1872:3, pi. II, figs. 1-4. Ryder, 1881:527. Packard, 1883:192. McNeill, 1888:8. Cook and Collins, 1895:63-64, pi. Ill, figs. 52-54. Williams and Hefner, 1928:115, fig. 12d. Causey, 1951:119, figs. 6-8; 1967:80, fig. 1. Palmen, 1952:8-11, figs. 10-17. Chamberlin and Hoffman, 1958:102-103. Shear, 1972:277, figs. 497-499. Kevan, 19836:2967. Shelley, 1988:1650. Trichopetalum album Cook and Collins, 1895:64-66, pis. II-III, figs. 22-29, 36-45. Chamberlin and Hoffman, 1958:102. Type Specimens—Five male and nine female syntypes (nos. 2208-2209) collected by O. Harger in May 1872 at New Haven, New Haven County, Connecticut; one male and one female syntype (no. 2125) taken by an unknown collector on an unknown date at Mt. Carmel, ca. 7 mi (11.2 km) north of New Haven, New Haven County. Remarks—Harger assigned three new species to his genus Trichopetalum but did not specify the type species, so Cook and Collins (1895) subsequently designated T lunatum. It is the only one of Harger's five milliped species to retain its original combination. The identity of T lunatum has been well established by Cook and Collins (1895), Palmen (1952), Causey (1967), and Shear (1972); a male syntype from New Haven that I dissected conformed to these Myriapod Types of Oscar Harger 5 accounts. For details of the genitalia, refer to the illustrations in Palmen (1952) and Shear (1972). Chordeumatida: Conotylidae Taiyutyla glomerata (Harger), new combination Trichopetalum glomeratum Harger, 1872:118, pi. II, fig. 5. Ryder, 1881:527 Packard, 1883:192. McNeill, 1888:8. Chamberlin and Hoffman, 1958:105. Shear, 1971:63. Craspedosoma glomeratum: Bollman, 1893:120. Conotyla glomerata: Cook and Collins, 1895:78. Cook, 1904:69. Type Specimen—Female holotype (No. 2173) collected by O. Harger in October 1871 from the vicinity of Canyon City, in the John Day River Valley, Grant County, Oregon. Remarks—The holotype is somewhat deformed, and its genitalia have been dissected and are lost. Cook and Collins (1895) stated that the original description was too brief to allow accurate generic placement but that the segment number, short fifth antennomere, and triangular eye patch resembled the condition in Conotyla. Shear (1971) agreed that accurate generic placement was impossible but perceived a similarity to Taiyutyla; he did not think the name could be referred to either Trichopetalum or Conotyla and considered it a nomen dubium. The holotype is about 8 mm long and has 30 post cephalic segments with obvious lateral tergal knobs that give rise to two prominent setae, so it is clearly a conotylid. Generic placement is impossible to determine with certainty until a male topotype is obtained, but the milliped is smaller and its lateral setae are much longer than those of comparative specimens of Conotyla atrolineata (Bollman), the western-most known representative of this genus, occurring in central British Columbia, northeastern Washington, and northern Idaho, over 200 mi (320 km) north northeast of Canyon City. These considerations tend to exclude Conotyla, but the type locality is also well removed from most of the known distribu-tions of the other northwestern conotylid genera Bollmanella and Taiyutyla, which are from southern coastal Oregon to Mason County, Washington, and in the Coast Ranges from San Francisco Bay to the Columbia River, respectively (Shear 1974, 1986). However, one species in each of these genera occurs east of the above ranges, B. bifurcata Shear, in the Wallowa Mountains, Wallowa County, Oregon, and T curvata Loomis and Schmitt, in Lincoln County, Montana, so either genus could occur in the Blue Mountains, which occupy an intermediate 6 Rowland M. Shelley geographical position between the Coast Range and both the Wallowa Mountains and Montana. Furthermore, Canyon City is only about 110 mi (176 km) southwest of the type locality of B. bifurcata. Therefore, I borrowed the types of both B. bifurcata and T. curvata for direct comparisons with that of glomerata. Few setae remain on the types of B. bifurcata, and those that do exist, on the caudal end of the male holotype, seem shorter and are not nearly as prominent as are those on glomerata. However, the setae on glomerata agree closely in length and prominence with those on the holotype of T. curvata. There is reasonable agreement in body dimensions between glomerata and both other conotylids, but because of the similarity in the setae, I provisionally assign glomerata to Taiyutyla, pending collection of a male topotype. This change, which formalizes Shear's (1971) perception of similarity to Taiyutyla, also necessitates the feminine suffix of the specific name. Fieldwork is needed in the Blue Mountains to collect a male conotylid to determine the identity and generic position of glomerata and to confirm or disprove this decision. Present evidence shows that the Conotylidae is much more wide-spread in the West than currently known. There is a female in the Florida State Collection of Arthropods from 12.5 mi (20 km) south of Baker City, Baker County, Oregon, that might be conspecific with glomerata, although this site is east of the Blue Mountains and presumably is drier than Canyon City. I also recently received two female conotylids that are superficially very similar to glomerata from the Snake Mountains, White Pine County, Nevada, in the eastern part of that state and hundreds of kilometers from any known site for the family. These two records plus glomerata suggest that conotylids could be scattered across the arid Columbia Plateau and Basin and Range Physiographic Provinces, where they are undoubtedly restricted to cool-er, forested regions at high elevations. The Ruby Mountains near Elko, Nevada, is another plausible area for conotylids, as are ranges in the central part of that state. Because only a few millipeds of any family have ever been collected from the "inselberg" mountains of these provinces, a concerted field effort is needed to both clarify the systema-tic positions of these conotylids and document the total diplopod fauna. Chordeumatida: Caseyidae Underwoodia iuloides (Harger) Trichopetalum iuliodes Harger, 1872:118. pi II, fig. 6. Trichopetalum juloides: Ryder, 1881:527. Trichopetalum iulioides: Packard, 1883:192. Trichopetalum iuloides: McNeill, 1888:8. Chordeuma iuloides: Bollman, 1893:121. Myriapod Types of Oscar Harger 7 Underwoodia polygama Cook and Collins, 1895:80-82, pi. X, figs. 180-190. Paleman, 1952:2-8, figs. l-9a. Chamberlin and Hoffman, 1958:107. Kevan, 19836:2968. Underwoodia iuloides: Cook and Collins, 1895:83-84, pi. X, figs. 177-178. Chamberlin and Hoffman, 1958:107. Kevan, 19836:2968. Shelley, 1988:1648-1649; In Press: Type Specimens—Eight female syntypes (No. 2207) collected by S. I. Smith in 1871 at Simon's Harbor (misspelled as Simmon's) on the north shore of Lake Superior, Ontario, Canada. This site is now in Pukaskwa National Park. Remarks—A review of Underwoodia with a description, discussion, and illustrations of U. iuloides is in press. For details on this species, see Shelley (1988). Fig. 1-3. Bollmaniulus furcifer, male syntype. 1, anterior gonopods, anterior view. 2, the same, posterior view. 3, posterior gonopods, anterior view. Scale line = 2.2 mm for figs. 1-2, 1.6 mm for fig. 3. Julida: Parajulidae Bollmaniulus furcifer (Harger) Figs. 1-3 lulus furcifer Harger, 1872:119, pi. II, fig. 7. Parajulus furcifer: Bollman, 1887:44. Cook, 1904:70-71, pi. V, figs. 5a-e. Chamberlin, 1920:35. 8 Rowland M. Shelley Paraiulus furcifer. Brolemann, 1895:69, pi. 7, figs. 21-23. Bollmaniulus furcifer: Verhoeff, 1926:65. Chamberlin and Hoffman, 1958:133. Buckett, 1964:18. Kevin, 19836:2964. Taijulus furcifer. Chamberlin, 1938:205. Caliulus furcifer. Chamberlin, 1940:15; 1944:80. Type Specimens—Three male and 13 female syntypes (No. 2172), most highly fragmented, collected by O. Harger and G. H. Collier in October 1871 from the vicinity of Canyon City, in the John Day River Valley, Grant County, Oregon. Remarks—Bollman (1887) transferred this species into Parajulus, misspelled as Paraiulus by Brolemann (1895), and Cook (1904) recorded it from Corvallis, Oregon. Chamberlin (1920) reported it from Clare-mont, Los Angeles County, California, surely a misidentification of another, possible congeneric parajulid. Verhoeff (1926) listed furcifer as the only component of his new genus Bollmaniulus, thereby mak-ing it the type species by monotypy as reported by Jeekel (1971). He did not specifically designate furcifer as the generotype, so this status does not result from original designation, as stated by Chamber-lin and Hoffman (1958). Chamberlin (1938, 1940) evidently was unaware of Verhoeffs action when he transferred furcifer into his new genera Taijulus and Caliulus, respectively, both of which have subsequently been placed in synonymy under Bollmaniulus (Chamberlin and Hoff-man 1958, Hoffman 1979). Chamberlin (1944) repeated the combination C. furcifer for a form from McCloud, Siskyou County, California, and added that the species was common over much of Oregon and California. Buckett (1964) recognized the combination Bollmaniulus furcifer and stated that it ranged from British Columbia into California. As noted by Hoffman (1979, 1992), the Parajulidae is one of the two most dominant Nearctic diplopod families in terms of com-ponent genera and species, the other being the Xystodesmidae (Polydesmida). It was studied from 1948 to about 1974 by Dr. Nell B. Causey, who amassed a large collection and examined most type specimens while conducting a detailed family revision. Unfortunately, she never completed the project and published only a few brief papers before her death in 1979. Consequently, knowledge of the Parajulidae is not nearly as advanced as those of the other major Nearctic diplopod families. Work on the taxon must essentially begin anew, a daunting task because of the diversity of the family and the enormous amount of preserved material in nearly every major and minor milliped repository on the continent. The types of /. furcifer will be crucial to an investigation of Pacific parajulids, because as the eighth oldest generic name in the family, Bollmaniulus has priority over such other nominal Pacific Myriapod Types of Oscar Harger Figs. 4-5 Chonaphe armata, male syntype. 4, telopodite of left gonopod, medial view. 5, the same, lateral view. Scale line = 1.14 mm for fig. 4, 1.0 mm for fig. 5. genera as Saiulus, Spathiulus, Sophiulus, Codiulus, and Simiulus, all authored by Chamberlin (1940), Tuniulus (Chamberlin 19415), and Mulaikiulus (Chamberlin 1941a), so additional generic synonymies could result from a study of these western forms. For the benefit of future students, I have included drawings of the gonopods of a male syntype (Figs. 1-3). Polydesmida: Xystodesmidae Chonaphe armata (Harger) Fig. 4-5 Polydesmus armatus Harger, 1872:119-120, pi. II, fig. 8. Leptodesmus armatus: Bollman, 1893:122. Chamberlin, 1911:264. Chonaphe armata: Cook, 1904:56-57, pi. Ill, figs. 2a-c. Attems, 1931:65- 67, figs. 100-101; 1938:156, fig. 177. Chamberlin, 1949:125. Chamberlin and Hoffman, 1958:27. Kevan, 19835:2968. Shelley, 1990:2314. Type Specimens—One male and two female syntypes, all highly fragmented, collected by O. Harger and G. H. Collier in October 10 Rowland M. Shelley 1871 from the vicinity of Canyon City, in the John Day River Valley, Grant County Oregon. This sample was discovered in the general milliped collection and is unnumbered. Remarks—Harger's single gonopod illustration enabled Cook (1904) to recognize that a male sent to him from an unknown locality in Washington was referrable to armatus. Bollman (1893) had earlier transferred armatus to Leptodesmus, a combination repeated by Chamberlin (1911), but Cook (1904) assigned it to his new genus, Chonaphe, a combination that subsequently has been recognized by Attems (1931, 1938), Chamberlin (1949), Chamberlin and Hoffman (1958), and Shelley (1990). Cook (1904) provided three additional genitalia drawings, and I include here medial and lateral views of the gonopod of a male syntype (Figs. 4—5). Five nominal species comprise Chonaphe, but Hoffman (1979) thought these might be subspecies. I (Shelley 1990) found few significant differences between these forms and concluded that the genus might be monotypic with C. armata being the oldest name. I am preparing a generic revision. ACKNOWLEDGMENTS—\ thank C. L. Remington and R. J. Pupedis for providing access to the Peabody Museum holdings and subsequently loaning Harger's types. The holotype of Taiyutyla curvata, housed at the National Museum of Natural History, Smithsonian Institution, Washington, D.C., was loaned by J. A. Coddington; the types of Bollmanella bifurcata, housed at the Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, were loaned by H. W. Levi. The conotylid from Baker County, Oregon, was discovered in material loaned by G. B. Edwards, Florida State Collection of Arthropods, Gainesville. Cathy Wood typed and retyped numerous drafts of the manuscript, and figures 1-5 were prepared by R. G. Kuhler. LITERATURE CITED Attems, C. G. 1931. Die familie Leptodesmidae und andere Poly-desmiden. Zoologica, Stuttgart, 30, Lief 3-4:1-149. Attems, C. G. 1938. Polydesmoidea II. Fam. Leptodesmidae, Platyrhacidae, Oxydesmidae, Gomphodesmidae. Das Tierreich, Lief 70:1-576. Bollman, C. H. 1887. Notes on North American Julidae. Annals of the New York Academy of Science 4:25-44. Bollman, C. H. 1893. The Myriapoda of North America. Bulletin No. 46, United States National Museum. Brolemann, H. W. 1895. Liste de myriapodes des Etats-Unis, et principale-ment de la Caroline du Nord, faisant partie des collections de M. Eugene Simon. Annales Societe Entomologique de France 65:43-70. Myriapod Types of Oscar Harger 11 Buckett, J. S. 1964. Annotated list of the Diplopoda of California. Simmons Publishing Company, Davis, California. Causey, N. B. 1951. New genera and species of chordeumoid millipeds in the United States, and notes on some established species. Proceedings of the Biological Society of Washington 64:117-124. Causey, N. B. 1967. Trichopetalum subterranewn, new species from Kentucky, new records and a key to the genus. Proceedings of the Biological Society of Washington 80:117-122. Chamberlin, R. V. 1911. Notes on myriopods from Alaska and Wash-ington. Canadian Entomologist 43:260-264. Chamberlin, R. V. 1920. Centipedes and millipedes from near Claremont. Pomona College Journal of Entomology and Zoology 12:35. Chamberlin, R. V. 1938. New diplopods. Proceedings of the Biological Society of Washington 51:205-208. Chamberlin, R. V. 1940. New genera and species of North American Paraiulidae. Bulletin of the University of Utah 30 [Biological Series, 5(7)]: 1-39. Chamberlin, R. V. 1941a. New western millipeds. Bulletin of the University of Utah 31[Biological Series, 6(5)]:l-23. Chamberlin, R. V. 1941b. New American millipeds. Bulletin of the University of Utah 31 [Biological Series, 6(4)]: 1-39. Chamberlin, R. V. 1944. Some records of myriopods collected by W. M. Pearce in California. Pan-Pacific Entomologist 20:79-80. Chamberlin, R. V. 1949. Some western millipeds of the family Chelodesmidae. Proceedings of the Biological Society of Washington 62:125-132. Chamberlin, R. V., and R. L. Hoffman. 1958. Checklist of the millipeds of North America. Bulletin No. 212, United States National Museum. Cook, O. F. 1904. Myriapoda of northwestern North America. Harriman Alaska Expedition 8(Insects, part l):47-82. Cook, O. F., and G. N. Collins. 1891. Notes on North American Myriapoda of the family Geophilidae, with descriptions of three genera. Proceedings of the United States National Museum 13:383-396. Cook, O. F., and G. N. Collins. 1895. The Craspedosomatidae of North America. Annals of the New York Academy of Science 9:1-100. Crabill, R. E. 1953. The Schendylidae of northeastern North America (Chilopoda: Geophilomorpha: Schendylidae). Journal of the New York Entomological Society 61:93-98. Crabill, R. E. 1961. A catalogue of the Schendylinae of North America including Mexico, with a generic key and proposal of a new Simoporus (Chilopoda: Geophilomorpha: Schendylinae). Entomological News 72:67-80. Gardner, M. R., and R. M. Shelley. 1989. New records, species, and genera of caseyid millipeds from the Pacific Coast of North America. Pan-Pacific Entomologist 65:177-268. 12 Rowland M. Shelley Harger, O. 1872. New North American myriapods. American Journal of Science and Arts 4:116-121. Hoffman, R. L. 1979. Classification of the Diplopoda. Museum d'Histoire Naturelle, Geneva, Switzerland. Hoffman, R. L. 1992. On the taxonomy of the milliped genera Pseudojulus Bollman, 1887, and Georgiulus, gen. nov., of southeastern United States (Julida: Parajulidae). Jeffersoniana 1:1-19. Jeekel, C. A. W. 1971. Nomenclator generum et familiarum Diplopodorum: A list of the genus and family-group names in the class Diplopoda from the 10th edition of Linnaeus, 1758, to the end of 1957. Mono-grafieen van de Nederlandse Entomologische Vereniging, Number 5. Kevan, D. K. McE. 1983a. A preliminary survey of known and potentially Canadian and Alaskan centipedes (Chilopoda). Canadian Journal of Zoo-logy 61:2938-2955. Kevan, D. K. McE. 1983b. A preliminary survey of known and potentially Canadian millipedes (Diplopoda). Canadian Journal of Zoology 61:2956- 2975. McNeill, J. 1888. A list, with brief descriptions, of all the species, including one new to science, of Myriapoda of Franklin County, Indiana. Bulletin of the Brookville Society of Natural History, Number 3. Meinert, F. 1870. Myriopoda Musaei Hauniensis I. Geophili. Naturhistorisk Tidsskrift 3:241-268. Packard, A. S. 1883. A revision of the Lysiopetalidae, a family of Chilognath Myriopoda, with a notice of the genus Cambala. Proceedings of the American Philosophical Society 21:177-209. Palmen, E. 1952. Survey of the Diplopoda of Newfoundland. Annales Zoologici Societatis Zoologicae Botanicae Fennicae 'Vanamo' 15:1-31. Ryder, J. A. 1881. List of the North American species of myriapods belonging to the family of the Lysiopetalidae, with a description of a blind form from Luray Cave, Virginia. Proceedings of the United States National Museum 3:524-529. Schuchert, C, and C. M. Levene. 1940. O. C. Marsh, Pioneer in Paleontology. Yale University Press, New Haven, Connecticut. Shear, W. A. 1971. The milliped family Conotylidae in North America, with a description of the new family Adritylidae (Diplopoda: Chordeumida). Bulletin of the Museum of Comparative Zoology 141:55-98. Shear, W. A. 1972. Studies in the milliped order Chordeumida (Diplopoda): A revision of the family Cleidogonidae and a reclassification of the order Chordeumida in the New World. Bulletin of the Museum of Comparative Zoology 144:151-352. Shear, W. A. 1974. The milliped genus Bollmanella (Diplopoda, Chordeumida, Conotylidae). Psyche 81:134-146. Shear, W. A. 1976. The milliped family Conotylidae (Diplopoda: Chordeumida). Revision of the genus Taiyutyla with notes on recently proposed taxa. American Museum Novitates Number 2600. Myriapod Types of Oscar Harger 13 Shelley, R. M. 1988. The millipeds of eastern Canada (Arthropoda: Diplopoda). Canadian Journal of Zoology 66:1638-1663. Shelley, R. M. 1990. A new milliped of the genus Metaxycheir from the Pacific coast of Canada (Polydesmida: Xystodesmidae), with remarks on the tribe Chonaphini and the western Canadian and Alaskan diplopod fauna. Canadian Journal of Zoology 68:2310-2322. Shelley, R. M. The milliped genus Underwoodia (Chordeumatida: Caseyidae). Canadian Journal of Zoology, In Press. Stuxberg, A. 1875. Lithobioidae Americae Borealis. Ofversigt af Kongl. Vetenskaps-Akademiens Forhandlingar 22:23-32. Verhoeff, K. W. 1926. Chilognathen-Beitrage. Zoologisher Anzeiger 68:57- 71. Williams, S. R., and R. A. Hefner. 1928. The millipedes and centipedes of Ohio. Bulletin Number 18, Ohio Biological Survey, 4(3) [Ohio State University Bulletin, 33(7)]:93-146 Accepted 9 September 1992 14 THE SEASIDE SPARROW, ITS BIOLOGY AND MANAGEMENT Edited by Thomas L. Quay, John B. Funderburg, Jr., David S. Lee, Eloise F. Potter, and Chandler S. Robbins The proceedings of a symposium held at Raleigh, North Carolina, in October 1981, this book presents the keynote address of F. Eugene Hester, Deputy Director of the U.S. Fish and Wildlife Service, a bibliography of publications on the Seaside Sparrow, and 16 major papers on the species. Authors include Arthur W. Cooper, Oliver L. Austin, Jr., Herbert W. Kale, II, William Post, Harold W. Werner, Glen E. Woolfenden, Mary Victoria McDonald, Jon S. Greenlaw, Michael F. Delany, James A. Mosher, Thomas L. Merriam, James A. Kushlan, Oron L. Bass, Jr., Dale L. Taylor, Thomas A. Webber, and George F. Gee. A full-color frontispiece by John Henry Dick illustrates the nine races of the Seaside Sparrow, and a recording prepared by J. W. Hardy supplements two papers on vocalizations. "The Seaside Sparrow, with its extensive but exceedingly narrow breeding range in the coastal salt marshes, is a fascinating species. All the authors emphasize that the salt marsh habitat is at peril. . . . The collection is well worth reading." — George A. Hall, Wilson Bulletin. 1983 174 pages Softbound ISBN 0-917134-05-2 Price: $15 postpaid. North Carolina residents add 6% sales tax. Please make checks payable in U.S. currency to NCDA Museum Extension Fund. Send order to: SEASIDE SPARROW, N.C. State Museum of Natural Sciences, P.O. Box 27647, Raleigh, NC 27611. A Late Pleistocene Vertebrate Assemblage from the St. Marks River, Wakulla County, Florida Timothy S. Young1 and Joshua Laerm The University of Georgia Museum ofNatural History and Department of Zoology, University of Georgia, Athens, Georgia 30602 ABSTRACT—The St. Marks River in the central panhandle of Florida contains a well known, apparently late Pleistocene vertebrate as-semblage that has been only superficially examined and reported. Previous collections are reviewed, and we report on new fossil materials recently obtained. Included are 37 species of mammals, 26 birds, 13 reptiles, 2 amphibians, and 9 fish. Of these, 14 species of mammals and 2 reptiles are limited solely to the Pleis-tocene. The fauna is mixed and reflects heterochronous deposition over time beginning at least in the late Pleistocene (Wisconsinan) and extending through the Recent. The species present reflect mixed woodland and grassland terrestrial communities as well as mixed estuarine and freshwater aquatic communities. The St. Marks River assemblage compares well to other contemporaneous late Pleisto-cene Florida panhandle sites. One extralimital taxa is reported, Pylodictic cf. P. olivaris, the flathead catfish, whose natural range has not been reported east of the Mobile Bay drainage basin. Florida is characterized by a number of rich and well documented Pleistocene vertebrate assemblages (Webb 1974a, Lundelius et al. 1983, Webb and Wilkins 1984) that contain a mixture of extant and extinct South American immigrant and North American endemic species. The majority of these sites are distributed throughout the peninsular portion of the State (Webb 1974a, Webb and Wilkins 1984). However, with the exceptions of Wakulla Springs (Brodkorb 1963, Webb 1974a), Chipola River (Martin 1969, Webb 1974a), and Aucilla River (Olson 1972, Webb 1914a, Gillette 1976a) very little attention has been devoted to sites on the panhandle of Florida. The St. Marks River, located 32 km south of Tallahassee in Wakulla County, is a particularly rich, late Pleistocene panhandle site that has only been superficially investigated (Gillette 19766, Steadman 1980). Leidy (1870), who reported on the occurrence of Mammuthus columbi (now M. jeffersonii), provided the first record of vertebrate fossil remains recovered from the St. Marks River, though the exact locality was not given. Subsequently, the St. Marks River has attracted 'Present address: Department of Zoology, University of Florida, Gainesville, FL 32611. Brimleyana 18:15-57, June 1993 15 16 Timothy S. Young and Joshua Laerm numerous amateur collectors but a limited number of professionals. Storrs Olson (National Museum of Natural History) collected from a broad, shallow water area in Wakulla County near the Leon County line several times between 1968 and 1970. Tall Timbers Research Station sponsored collecting parties in the same area during 1974. Published accounts of the fauna are those of Gillette (1976b), who reported the mammals, and Steadman (1980), who discussed two specimens of Meleagris gallopavo. Storrs Olson (personal communication) examined the avian assemblage from the previous collections; however, he did not publish his findings. With the exception of Mammuthus sp. Mammut americanum, Synaptomys australis, and possibly Equus, the majority of species from Gillette's (1916b) report are extant. Gillette (19766) suggested that the assemblage was important because it represented a restricted temporal interval of the latest Pleistocene through the Holocene. Olsen (personal communication) felt the avian assemblage was very similar to that of today. Steadman (1980) characterized the site as a late Pleistocene deposit. The St. Marks River has also been reported by Lundelius et al. (1983) as being a naturally accumulating, fluvial Rancholabrean deposit. The purpose of our study is to review previous collections and to report on new fossil materials recently obtained from the St. Marks River. We provide information regarding the paleoenvironment of the depositional area and compare the St. Marks River fauna to other late Pleistocene faunas in the region. GEOLOGICAL AND GEOGRAPHIC SETTING Florida consists of five naturally occurring topographical divisions (Cooke 1939:14). The St. Marks River drainage basin is in the coastal lowlands division. Although the panhandle of Florida shows a topographical record of the relict shorelines, no ages have been securely assigned to these formations (Winker and Howard 1977a, 6). The coast line of the panhandle during the late Pleistocene is reported to be similar to that of today (Winker and Howard 19776). The St. Marks River is considered part of the Gulf Hammock region; it is underlain by the Upper Oligocene Suwannee Limestone (Harper 1914:302). The early Miocene St. Marks Formation overlies the Suwannee Formation in almost all of Wakulla County (Puri and Vernon 1964). The St. Marks Formation was revised to include the calcareous downdip facies of the Tampa Formation (Puri 1953). These formations can be found in many areas as outcroppings in springs and rivers (Spencer and Rupert 1987). The surface is mostly loamy St. Marks River Fauna 17 sand, probably Pleistocene in origin. The soil surrounding the river's edge is classified as Tooles-Nutall fine sand that is frequently flooded (Spencer and Rupert 1987). Topographically the region is nearly level, except for a few hilly areas (Harper 1914:302). The whole area east of the Apalachicola River in Wakulla County is called the Woodville Karst Plain (Hendry and Sproul 1966, Yon 1966), characterized by sand dunes overlying limestone (Hendry and Sproul 1966:154). The St. Marks River is fed by the St. Marks Spring located just inside Leon County (Fig. 1). Limestone lines the perimeter of the spring. The vent is located about 26 m below the water surface and has an average base flow of 14.7 m3/sec (Rosenau et al. 1977). This measurement was taken approximately 800 m down stream Leon County ) Basin ' Wakulla County • 3 •f f St. Marks River Wakulla River^— - /• // * 1 ( l_l \ I 1 mile N t Gulf of Mexico Fig. 1. St. Marks River in the central panhandle of Florida. Solid circles indicate 1987 collection sites. 18 Timothy S. Young and Joshua Laerm from the main vent. The pH and temperature as measured 16 July 1974 were 7.6 and 21.OC, respectively. Newport Spring also feeds the river about 800 m north of the U.S. Highway 98 bridge. The discharge of the spring as measured 2 March 1972 was 0.23 m3/sec with a pH of 7.8 and water temperature of 19C (Rosenau et al. 1977). Primary depositional site(s) were not located. The fossils are probably eroding out of the banks along much of the length of the river and washed down river by the current. Dense accumulations of fossils may be found in sand deposits, around submerged debris, and in deep holes along the entire length of the river. The St. Marks River with its shallow, relatively clear waters with abundant fossil and archaeological materials has been a popular recreational S.C.U.B.A. diving are for the past 30 years. Local divers report huge quantities of fossils have been recovered by amateur collectors. One of us (J.L.) observed an entire pick-up truck load of fossils being removed in 1978. Local divers report that have collected "tons of it." Although several large private collections of St. Marks material exist, unfortunately they have been mixed with fossils from other regional aquatic systems which makes their inclusion here inappropriate. METHODS We made extensive new collections and reviewed previously collected materials housed at the Florida Museum of Natural History, University of Florida (UF), and the National Museum of Natural History (USNM), Washington, D. C. Our collections are housed at the University of Georgia Museum of Natural History (UGAMNH). Collection efforts were concentrated in six separate locations approximately 3.2 km in either direction from the U.S. Highway 98 bridge that crosses the St. Marks River (Fig. 1). The fossils were collected from 16 to 19 July 1987 by a team of six people from the University of Georgia using S.C.U.B.A. gear. The majority of the fossil materials was collected by hand from these locations along the river. In addition, extensive sand samples were taken at each site for subsequent screening. To preliminarily identify recovered materials we used the Comparative Reference Skeletal Collection of the Zooarchaeology Laboratory, the University of Georgia Museum of Natural History. Reference sources were also used in preliminary identifications. All materials were subsequently taken to the Paleontology Laboratory, the Florida Museum of Natural History, University of Florida, to confirm identifications. Notes were made on the element identified, side, and fusion of bones where possible. St. Marks River Fauna 19 SYSTEMATIC PALEONTOLOGY Standardized common and current scientific names follow Robins et al. (1991) for fishes; Collins (1990) for amphibians and reptiles; American Ornithologists' Union (1983) for birds, and Kurten and Anderson (1980) and Jones et al. (1992) for mammals. Museum acronyms are indicated in the introduction. A complete faunal listing of the species recovered from the St. Marks site is provided in Table 1. CLASS MAMMALIA Order Didelphimorphia Family Didelphidae Didelphis virginiana Kerr Virginia Opossum Material—A single left dentary, UGAMNH1735. Remarks—The single element is identical to that of modern Didelphis virginiana. This was the only marsupial species present in North America during the Pleistocene. It is known from numerous fossil sites in Florida (Webb 1974a). Its stratigraphic range includes Middle Blancan to Recent (Kurten and Anderson 1980). It occurs in a variety of habitats, but it is usually found in forests and woodlands near water (Gardner 1973). We follow Marshall et al. (1990) in the use of the ordinal name Didelphimorphia as do Jones et al. (1992). Order Xenarthra Family Dasypodidae Holmesina septentrionalis (Leidy) Northern Pampathere Material—Right astragulus, UGAMNH2012; right calcaneus, UGAMNH2159; right metacarpus II, UGAMNH1981; two phalanges, UGAMNH1982-1983; numerous dermal plates, UGAMNH1954-1980, 1984- 2029, 2160, 2166. Remarks—The species is known from numerous sites throughout the South and Southeast. Its range is somewhat similar to that of its modern relative, Dasypus novemcinctus, and Holmsina had a similar preference for open woodlands (Kurten and Anderson 1980). Like its modern counterpart, Holmsina probably fed on insects and various invertebrates. Kurten and Anderson (1980) suggest this diet might have restricted them to relatively warm climates where food was available year round. Specimen UHAMNH2159, a right calcaneus, has rodent and carnivore gnaw marks that occurred prior to fossilization. Its stratigraphic range is early Irvingtonian to Wisconsinan (Kurten and Anderson 1980). 20 Timothy S. Young and Joshua Laerm Table 1. List of vertebrate species recovered from the St. Marks River. The figure f indicates extinct forms. Class Mammalia Order Didelphimorphia Family Didelphidae Didelphis virginiana Order Xenarthra Family Dasypodidae \Holmesina septentrionalis Family Megalonychidae ^Megalonyx jeffersonii Family Mylodontidae ^Glossotherium harlani Order Primates Family Hominidae Homo sapiens Order Lagomorpha Family Leporidae Sylvilagus sp. Order Rodentia Family Castoridae Castor canadensis Family Geomyidae Geomys pinetis Family Muridae Microtus sp. Microtus pinetorum Neofiber alleni Ondatra zibethicus Synaptomys australis Order Carnivora Family Mustelidae Mustela sp. Lutra canadensis Mephitis mephitis Family Canidae Canis sp. "\Canis dirus Urocyon cinereoargenteus Family Procyonidae Procyon lotor Family Ursidae Ursus cf. U. americanus Family Felidae Felis sp. ^Smilodon sp. Order Proboscidea Family Mammutidae \Mammuthus jeffersonii Family Elephantidae \Mammut americanum Order Perissodactyla Family Equidae Equus sp. Family Tapiridae ~\Tapirus sp. Order Artiodactyla Family Tayassuidae ^Platygonus compressus Family Suidae Sus scrofa Family Camelidae "\Hemiauchenia macrocephala ^Palaeolama mirifica Family Cervidae Odocoileus virginianus Family Bovidae Bison sp. Bison bison Bos taurus Class Aves Order Podicipediformes Family Podicipedidae Podiceps auritus Podilymbus podiceps Order Pelecaniformes Family Phalacrocoracidae Phalacrocorax auritus Order Ciconiformes Family Ardeidae Ardea herodias Butorides virescens Egretta caerulea Family Threskiornithidae Eudociums albus Order Anseriformes Family Anatidae Aix sponsa Anas acuta St. Marks River Fauna 21 Table 1. Continued. Anas americana Anas discors Anas platyrhynchos Anas sp. Aythya collaris Aythya sp. Branta canadensis Bucephala albeola Lophodytes cucullatus Mergus merganser Order Falconiformes Family Accipitridae Buteo jamaicensis Pandion haliaetus Order Galliformes Family Phasianidae Meleagris gallopavo Order Gruiiformes Family Rallidae Fulica americana Gallinula chloropus Family Aramidae Aramus guarauna Order Strigiformes Family Strigidae Strix varia Class Reptilia Order Testudines Family Chelydridae Chelydra serpentina Family Kinosternidae Gen. et spec, indet. Family Emydidae Pseudemys concinna Pseudemys floridanus Pseudemys nelsoni Trachemys scripta Terrapene Carolina Terrapene Carolina putnami Family Testudinidae Geochelone incisa Geochelone sp. Gopherus polyphemus Family Trionychidae Trionyx sp. Order Squamata Family Colubridae Elaphe obsoleta Gen et spec, indet. Order Crocodilia Family Alligatoridae Alligator mississippiensis Class Amphibia Order Caudata Family Sirenidae Siren sp. Order Anura Gen. et spec, indet. Class Osteichthyii Order Lepisosteiformes Family Lepisosteidae Lepisosteus sp. Order Amiiformes Family Amiidae Amia calva Order Siluriformes Family Ictaluridae PyIodictis cf. P. olivaris Family Ariidae Ariopsis felis Order Salmoniformes Esocidae Esox sp. Order Perciformes Family Percichthyidae Morone saxatilis Family Sparidae Archosargus probatocephalus Family Sciaenidae Sciaenops ocellatus Family Mugilidae Mugil sp. 22 Timothy S. Young and Joshua Laerm Family Megalonychidae Megalonyx jeffersonii (Desmarest) Jefferson's Ground Sloth Material—A single phalanx, UGAMNH2135, and tooth, UGAMNH2136. Remarks—Jefferson's ground sloth occurred in woodlands where it apparently fed on nuts, berries, leaves, and twigs (Stock 1925). It is known from a number of sites in the Southeast including Florida (Webb 1974a), Georgia (Ray 1967), South Carolina (Hay 1923, Roth and Laerm 1980), and Tennessee (Guilday et al. 1969). It could have tolerated a seasonally cool climate as evidenced by its Pleistocene occurrence in what is now Canada and Alaska (McNab 1985). It is reported from Irvingtonian to Rancholabrean sites with a terminal date of 13,890 years B.P., although Kurten and Anderson (1980) suggest it may have survived even longer in Florida. Family Mylodontidae Glossotherium harlani (Owen) Harlan's Ground Sloth Material—Two teeth, UGAMNH2137-2138. Remarks—This was an open plains and grassland species (Stock 1925). It is reported from Irvingtonian to Rancholabrean sites with a terminal date of 13,890 years B.P., although Kurten and Anderson (1980) suggest it may have survived even longer in Florida. Order Primates Family Hominidae Homo sapiens Linnaeus Human Material—Cranial fragment, UF21280. Remarks—This single specimen was recovered by Gillette (19766). Unfortunately, the cranial fragment was not available for examination. We are, therefore, unable to comment on the degree of mineralization. No other human remains were recovered in our efforts. The presence of considerable amounts of Native American cultural material (pottery shards) as well as 18-20th century European-American artifacts indicates the St. Marks River was a site of human occupation before and after European contact. St. Marks River Fauna 23 Order Lagomorpha Family Leporidae Sylvilagus sp. indet. Material—Tooth fragment, UF21301. Remarks—Rabbits are a common component of most Pleistocene sites in Florida. It is surprising no more than a single tooth fragment was encountered in the St. Marks River material. Two species of rabbit occur in the St. Marks region today, the eastern cottontail, Sylvilangus floridanus, and the more common swamp rabbit, S. aquaticus. The former prefers heavy brushy, forested areas with open areas nearby and edges of swamps. The latter is most common in marshes, swamps, and bottomlands (Golley 1962). Order Rodentia Family Castoridae Castor canadensis Kuhl Beaver Material—Left ulna, UGAMNH2126; right upper molar, UGAMNH2125; right M3 , UGAMNH2124; four molars, UF21294. Remarks—Two beaver species occurred in Florida in the late Pleistocene, Castoroides ohioensis and Castor canadensis. Both have even been found in the same deposits (Webb 1974a); however, only the latter is represented in the St. Marks River fauna. The beaver is found in any suitable water habitat including rivers, streams, lakes, and marshes (Lowery 1974). Its relative rarity in the St. Marks may be related to the presence of Alligator. The stratigraphic range is late Blancan to Recent (Kurten and Anderson 1980). Family Geomyidae Geomys pinetis Rafinesque Southeastern Pocket Gopher Material—A single lower fourth premolar, UF21291. Remarks—Geomys pinetis is the only species of pocket gopher in the Southeast. It is associated with the sandy soils of the Coastal Plain (Golley 1962) and is present today in the uplands adjacent the St. Marks River. It is known from late Irvingtonian to Recent (Kurten and Anderson 1980). Family Muridae Microtus sp. indet. Material—Left M3 , UGAMNH2127. Remarks—This fragment, while certainly Microtus, could not be 24 Timothy S. Young and Joshua Laerm referred to a species with confidence. We follow Jones et al. (1992) in their use of the familial name Muridae. Microtus pinetorum (LeConte) Pine Vole Material—Right M2 , UGAMNH2128. Remarks—This molar compares well to modern Microtus pinetorum. Regionally, the pine vole can be found in a wide range of habitats from hardwood and pine forests to overgrown fields (Golley 1962). The stratigraphic range is Sangamonian to Recent (Kurten and Anderson 1980). Neofiber alleni True Round-tailed Muskrat Material—right M2 , UGAMNH2121; right M3 , UGAMNH2123; right M3 , UGAMNH2122; maxilla, UF21293. Remarks—Neofiber alleni is a semi-aquatic mammal that prefers permanent bodies of water with emergent aquatic vegetation (Frazier 1977). Although it has a restricted range today, essentially extreme northern Florida and south Georgia, during the Pleistocene it ranged as far west as Kansas (Hibbard 1943). It is reported from late Irvingtonian to Recent (Kurten and Anderson 1980). The stratigraphic range is Illinoian to Recent (Kurten and Anderson 1980). Ondatra zibethicus (Linnaeus) Muskrat Material—Right dentary with M1 AND M2 , UGAMNH2120; dentary, UF21292. Remarks—The muskrat, like the round-tailed muskrat, is a semi-aquatic mammal that prefers permanent bodies of water (Nelson and Semken 1970). There is not overlap in the range of the two species today. However, Martin and Webb (1974) indicate they were sympatric in at least two late Pleistocene Florida faunas, Devils Den and Ichetucknee River. The occurrence of the two species in the St. Marks River fauna is not overly suggestive that they were sympatric here in the past because of the apparently heterochronous deposition at St. Marks. Furthermore, although the muskrat does not presently occur in the St. Marks River or Apalachicola River drainages, it is known from the extreme western panhandle and the Upper Coastal Plain of Georgia, a distance of 120 km. St. Marks River Fauna 25 Synaptomys australis Simpson Florida Bog Lemming Material—Left mandible with Mb UF21295. Remarks—The specimen referred to in Gillette's (1976b) review of the St. Marks River is the only record of this species at the site. In Florida it is known primarily from Sangamonian and Wisconsinan assemblages, although elsewhere it is known from the Illinoian through the Wisconsinan (Kurten and Anderson 1980). Its presence at Devils Den suggests it might have persisted until about 8,000 years B.P. (Martin and Webb 1974), although this radiocarbon date is considered suspect. The Florida bog lemming is similar morphologically to S. cooperi, the northern bog lemming, but differs considerably in size; it is about 35% larger than S. cooperi. Kurten and Anderson (1980) suggest it might represent a clinal variate of S. cooperi. It was an inhabitant of moist bogs and damp meadows (Burt 1928). Order Carnivora cf. Order Carnivora, gen. et sp. indet. Material—A left coronoid, UGAMNH1881. Remarks—This specimen, though carnivore-like, could not be identified to the familial level. Family Mustelidae Mustela sp. indet. cf. Weasel Material—A single left humerus, UGAMNH1738 and right P3 , UGAMNH1736. Remarks—Two species of weasel, Mustela frenata and M. vison, are common to the region today. Both are known from the Irvingtonian to Recent and are represented in regional fossil sites (Webb 1974a). However, fossil weasels have been reported from very few sites in Florida (Martin 1974, Webb 1974a). Lutra canadensis (Shreber) River Otter Material—Left humerus, UGAMNH1741. Remarks—This material compares well to modern Lutra canadensis. The stratigraphic range includes early Irvingtonian to Recent, and the species is represented in numerous regional sites (Kurten and Anderson 1980). The species occurs in woodlands near rivers and streams but is also known from tidal creeks and marshlands (Lowery 1974). 26 Timothy S. Young and Joshua Laerm Mephitis mephitis (Schreber) Striped Skunk Material—Right mandible, UGAMNH1746; left humerus, UGAMNH1737. Remarks—This material compares well to modern Mephitis mephitis, which can be found in mixed woodlands, brushlands, or prairies but generally in reasonable proximity to water (Lowery 1974). The stratigraphic range is mid Blancan to Recent (Kurten and Anderson 1980). Family Canidae Canis sp. indet. Material— Left ilium, UGAMNH1739; right dentary, UGAMNH1878, 1880; right scapula, UGAMNH1879. Remarks—None of these elements could be identified beyond the generic level. They are well mineralized, suggesting they are not modern C. familiaris contaminants. Several species of Canis are known from late Pleistocene sites in Florida. These include C. lupus, the gray wolf; C. rufus, the red wolf; C. latrans, the coyote; and C. dims, the dire wolf. Martin (1974) has concluded that only two species, C. rufus and C. dirus, are common to middle and late Pleistocene deposits of Florida. Canis lupis is typical of Irvingtonian deposits, whereas C. dirus is representative of the Rancholabrean. Canis dirus Leidy Dire Wolf Material—left radius, UGAMNH1877. Remarks—Canis dirus is known from a number of late Pleistocene sites in Florida (Webb 1914a) and is one of the more common species of mammals at numerous Rancholabrean sites throughout North America. It is thought to have inhabited a wide range of habitats because it was a hunter and scavenger (Kurten and Anderson 1980, Lundelius et al. 1983). The stratigraphic range is early Illinoian to Wisconsinan (Kurten and Anderson 1980). The most recent terminal date for extinction is given at about 8,000 years B.P. in Florida (Martin and Webb 1974), but somewhat earlier (approximately 9,000-10,000 year B.P.) elsewhere (Kurten and Anderson 1980). Urocyon cinereoargenteus (Shreber) Gray Fox Material—Right dentary, UGAMNH1743; left frontal, UGAMNH1744. St. Marks River Fauna 27 Remarks—This material is not well mineralized, which suggests that it is a modern contaminant. However, Urocyon cinereoargenteus would be expected in this fauna. It can be found in a wide range of habitats today, but brushy and woody areas probably best describe the preferred habitat in the South and Gulf Coast area (Lowery 1974). The stratigraphic range in Florida is Middle Rancholabrean to Recent (Martin and Webb 1974). Elsewhere it is known as early as the Blancan (Kurten and Anderson 1980). Family Procyonidae Procyon lotor (Linnaeus) Racoon Material—Three left dentaries, UGAMNH1742, 1747, 1750. A partial skeleton is represented by UF21296. Remarks—The University of Georgia material is not well mineralized, which suggests it could be a modern contaminant, since Procyon lotor is part of the modern fauna. In the Florida panhandle today, the racoon is an inhabitant of forested bottomland swamps. It fossil record in Florida extends from the Late Irvingtonian to Recent (Martin and Webb 1974). Family Ursidae Ursidae gen. et sp. indet. Material—Three phalanges, UGAMNH1745, 1749, 1752. Remarks—Generic identity of this material is uncertain. In addition to the modern black bear, Ursus americanus Pallas, several extinct species of bears are known from the Pleistocene of Florida. These include the cave bear, Tremarctos floridanus (Gidley), and the lesser short-faced bear, Arctodus pristinus Leidy, all of which persisted at least until the late Wisconsin (Kurten and Anderson 1980). Ursus cf. U. americanus Pallas cf. Black Bear Material—A single right dentary with M1? UGAMNH1751. Remarks—This specimen is well mineralized, but it is too worn for positive identification. The black bear can be found in forests and bottomland swamps throughout much of the Southeast (Golley 1962, Lowery 1974). It is represented in numerous late Pleistocene sites. The stratigraphic range is early Irvingtonian to Recent (Kurten and Anderson 1980). 28 Timothy S. Young and Joshua Laerm Family Felidae Felis sp. indet. Material—Left radius, UGAMNH1740. Remarks—This specimen is a large Felis, but it is too worn for positive identification. Webb (1974a) states that several species of Felis are known from the Late Pleistocene of Florida, and include F. atrox Leidy, F. concolor Linnaeus, F. onca (Linnaeus), F. pardalis Linnaeus, F. rufus Schreber, and F. yagouaroundi Geoffroy. Another possibility is Felis amnicola, a new, small cat described by Gillette (1976a). The description is based on several specimens from various localities in Florida and possibly Georgia. Smilodon sp. indet. Material—Left scapho-lunar, UGAMNH1748. Remarks—The sabertooth cats are reported from a dozen or more late Pleistocene sites in Florida (Webb 1974a, Kurten and Anderson 1980). Slaughter (1963) proposed a series of successional changes in Smilodon species throughout the North American Pleistocene. Webb (1974a) concurs that records of Smilodon in Florida support such a successional outline: Smilodon gracilis is a late Blancan and early Irvingtonian; S. fatalis is representative of late Irvingtonian and early Rancholabrean sites; and that S. floridanus is typical of the late Rancholabrean. The temporal span reflected by other faunal elements from the St. Marks would be more suggestive of the latter species; however, given the similarity of these species, more precise identification is impossible from the limited available material. Smilodon could probably have been found in habitats ranging from grassland to woodland (Merriam and Stock 1932, Lundelius et al. 1983). Order Proboscidea Proboscidea gen. et sp. indet. Material—Sesamoid, UGAMNH1098; tusk fragment, UF21255; skull fragment, UF21256; leg fragment, UF21257; vertebral fragment UF21258. Remarks—These specimens are very definitely proboscidean, but assignment to species is impossible. Family Mammutidae Mammut americanum (Kerr) American Mastodon Material—Axis fragment, UGAMNH1614; tooth fragments, UGAMNH1612, 1613, 1615, 1616, UF21267 and 21276; tusk fragments, UF21277-21278; proximal humerus, UF21279; calcaneus, UF21290. St. Marks River Fauna 29 Remarks—The morphology of the elements is consistent with its identification as Mammut americanum. Dreimanis (1968) suggested that M. americanum inhabited coniferous forests. The stratigraphic range is early Blancan to Wisconsinan (Kurten and Anderson 1980). Family Elephantidae Mammuthus jeffersonii (Osborn) Jefferson's Mammoth Material—Tooth fragments, UGAMNH1607-1611; tooth fragments, UF21259-21262, 21264-21266; mandibular symphysis, UF21263. Remarks—The morphology of the tooth fragments and mandibular symphysis is consistent with its identification as Mammuthus jeffersonii. Jefferson's mammoth probably inhabited open grasslands (Stock 1963, Harrington et al. 1974). The stratigraphic range is Illinoian to Wisconsinan (Kurten and Anderson 1980). Order Perissodactyla Family Equidae Equus sp. indet. Horse Material—left astragalus, UGAMNH1035; cervical vertebra, UGAMNH1170; left upper cheek tooth, UGAMNH1045; right upper cheek tooth, UGAMNH1031; right lower cheek tooth, UGAMNH1048; cheek tooth, UGAMNH1046, 1047, 1062; right deciduous P 2 , UGAMNH1042; left deciduous P2 , UGAMNH1061; right cuneiform, UGAMNH1049; left femoral head, UGAMNH1034; right distal humeral epiphysis, UGAMNH1054; left I 3 , UGAMNH1036; right I 2 , UGAMNH1041; lower incisor, UGAMNH1056; left I 3 , UGAMNH1059; left I 1 , UGAMNH1060; incisive fragment, UGAMNH1032; left upper molar, UGAMNH1038; right M2,UGAMNH1039; right M3 , UGAMNH1044; left M3 , UGAMNH1057; upper molar fragment UGAMNH1029; left navicular, UGAMNH1063; medial phalanges UGAMNH1030, 1050, 1053; distal phalanx, UGAMNH1051; proximal phalanx, UGAMNH1055; left P 2 , UGAMNH1037; right P 2 , UGAMNH1040; left upper premolar, UGAMNH1043; right lower premolar, UGAMNH1058; left scapula, UGAMNH1052; sesamoid, UGAMNH1033; medial phalanx, UF21228; teeth, UF21229-21238; teeth UF21240-UF21254; cheek tooth, axis, and pelvis, UF21297. Remarks—Equus is well represented in St. Marks River. A portion of the material is poorly mineralized and probably represent contaminants of the modern E. caballus. However, the majority of elements are well fossilized, and it is likely that most of the material 30 Timothy S. Young and Joshua Laerm is of late Pleistocene origin. Given the uncertain relationships of late Pleistocene horses in general and the likelihood of heterochronous deposition, we did not assign the material to a particular species. Pleistocene Equus was generally a grassland species (Kurten and Anderson 1980). Family Tapiridae Tapiridae gen. et spec, indet. Material—Left dentary, UGAMNH2068. Remarks—This edentulous specimen could not be assigned to Tapirus with confidence, although the morphology is similar. Tapirus sp. indet. Tapir Material—Right upper deciduous premolar, UGAMNH2070, left upper deciduous premolar, UGAMNH2071; right fibula, UGAMNH2069. Remarks—The available material, while certainly Tapirus, could not be referred to a species with confidence. Tapirs occur in wet woodlands (Simpson 1945, Gray and Crammer 1961). Order Artiodactyla Family Tayassuidae Platygonus compressus LeConte Flat-headed Peccary Material—Axis, UGAMNH2072. Remarks—The material has the diagnostic characters of Platygonus compressus which is thought to have wide environmental tolerances, but was probably most associated with open woodlands (Martin and Guilday 1967, Ray et al. 1970). The stratigraphic range is Sangamonian to Wisconsinan (Kurten and Anderson 1980). Family Suidae Sus scrofa Linnaeus Pig Material—Left maxilla with P3 and P4 , UGAMNH1159; right humeral fragment, UGAMNH1160; right femoral diaphysis, UGAMNH1162; right radial fragment, UGAMNH1161; left humeral fragments, UGAMNH1163, 1158; right ilial fragment, UGAMNH1157; distal humeral fragment, UGAMNH1178; left femur, UGAMNH1156. Remarks—None of the pig material showed evidence of significant mineralization. The pig was introduced during historic times and represents a domesticate. Specimen UGAMNH1178 shows marks of a saw. St. Marks River Fauna 31 Family Camelidae Camelidae gen. et sp. indet. Material—Proximal phalanges, UGAMNH1647, 1648; right scaphoid, UGAMNH1649; right astragalus, UGAMNH1650; right scapula, UGAMNH1651; left proximal femoral fragment, UGAMNH1652. Remarks—These specimens have distinctive camelid familial characters, but cannot be assigned to a particular species. Hemiauchenia macrocephala (Cope) Large-headed Llama Material—Proximal phalanges, UGAMNH2151, 2152. Remarks—The identification of these elements to Hemiauchenia macrocephala is based on the size of the phalanges. According to Webb (1974/?), H. macrocephala was a plains and grasslands inhabitant. The stratigraphic range is Wisconsinan to Recent (Kurten and Anderson 1980). Because this species has such a limited stratigraphic range, at least a portion of the deposit can be correlated to the Wisconsinan. Paleolama mirifica (Simpson) Stout-legged Llama Material—Left proximal metacarpal fragment, UGAMNH2146; right humerus, UGAMNH2145; left metatarsus, UGAMNH2158; left M3, UGAMNH2147; metapodial, UGAMNH2148; right distal humerus, UGAMNH2149; left pisiform, UGAMNH2150. Remarks—The stratigraphic range is late Irvingtonian to Wisconsinan (Kurten and Anderson 1980). Webb (1974b) reports them to be an inhabitant of grasslands and savannahs. Specimen UGAMNH2158, a left metatarsus, has longitudinal cracks indicative of weathering prior to fossilization. Family Cervidae Odocoileus virginianus (Zimmerman) White-tailed Deer Material—Antler fragments, UGAMNH1103, 1106, 1134, 1148; left astragalus, UGAMNH1101, 1124, 1210, 1757; right astragalus, UGAMNH1071, 1113, 1125, 1677, 1758, 2162; right calcaneus, UGAMNH1150, 1181, 1204, 1756; left calcaneus, UGAMNH1069, 1073, 1077, 1079, 1205, 1653, 2153, 2154, 1667, 1678; right cubonavicular, UGAMNH1074, 1111; right dentary with P1? P2 , P3 , M2 , M3 , UGAMNH1081; left dentary with Mlt UGAMNH1131; right dentary with Mh M2 , M3 , UGAMNH1108, 1126; right dentary with P2, P3 , M1? M2 , M3 , UGAMNH1191; right proximal femoral fragment, UGAMNH1184; femoral diaphysis, UGAMNH1064; left femoral head, 32 Timothy S. Young and Joshua Laerm UGAMNH1129; left femoral diaphysis, UGAMNH2163; right distal femur, UGAMNH11139; left distal femoral fragment, UGAMNH1085, 1088; left femoral lesser trochanter, UGAMNH1130; right femoral diaphysis, UGAMNH1139, 2164; right frontal with antler, UGAMNH1068, 1099; left frontal with antler, UGAMNH1012, 1082, 1118, 1121, 1666; frontal with antler pedicle, UGAMNH1196, 1203; right humeral fragments, UGAMNH1075, 1100, 1122, 1133, 1142, 1661, 1668; left humeral fragments, UGAMNH1070, 1093, 1136, 1137, 1143, 1185, 1186, 1235, 1679; right ilial fragments, UGAMNH1076, 1090, 1681; left ilial fragments, UGAMNH1079, 1086, 1147; left ischial fragments, UGAMNH1105, 1119, 1682; right lunate, UGAMNH1112; right maxilla with P4 , M1 , UGAMNH1206; left metacarpal fragments, UGAMNH1087, 1102, 1114, 1115, 1146, 1180, 1198, 1663; right metacarpal fragments, UGAMNH1072, 1084, 1092, 1097, 1116, 1665; metacarpal diaphysial fragments, UGAMNH1104, 1193-1195, 1670; right metatarsal fragments, UGAMNH1091, 1183, 1201, 1659, 1669, 1675, 1759; metatarsal diaphysial fragments, UGAMNH1117, 1199, 1200, 1208, 1212, 1656, 1672, 1673, 1676; left metatarsal fragments, UGAMNH1120, 1192, 1654, 1655, 1662, 1664, 1671; right M1 , UGAMNH1109, UGAMNH1151; left M3 , UGAMNH1289; right M3 , UGAMNH1190; left petrous, UGAMNH1110, 1128, 1202, 1214; medial phalanx, UGAMNH1080, 1141, 1209, 1213, 1753, 1754; proximal phalanx, UGAMNH1078, 1109, 1127, 1182, UGAMNH1211, 1755; left radial fragment, UGAMNH1065; right radial diaphysial fragment, UGAMNH1067; left radial fragments, UGAMNH1207, 2165; right radial fragments, UGAMNH1144; sacrum, UGAMNH1089; left scapular fragments, UGAMNH1140, 1188; right scapular fragments, UGAMNH1094, 1145; right distal tibial fragments, UGAMNH1095, 1123, 1658, 1680; left distal tibial fragments, UGAMNH1096, 1657; left proximal tibial fragment, UGAMNH1674; right proximal ulnar fragment, UGAMNH1132; left proximal ulnar fragment, UGAMNH1197; thoracic vertebral fragment, UGAMNH1760; lumbar vertebral fragment, UGAMNH1187; cervical vertebral fragment, UGAMNH1066; atlar fragments, UGAMNH1083, 1149; axial fragment, UGAMNH1660; antler, UF21289; five mandibles, UF21298. Remarks—The deer material shows a considerable range of mineralization. A significant portion is poorly mineralized and probably represents modern contaminants. The remaining material, however, is well mineralized, but mineralization alone is a poor indicator of possible Pleistocene age. The stratigraphic range of species is middle Blancan to Recent (Kurten and Anderson 1980). Odocoileus is a woodland and forest edge species (Golley 1962, Lowery 1974, Lundelius et al. 1983). St. Marks River Fauna 33 Family Bovidae Bovidae gen. et sp. indet. Material—Proximal phalanges, UGAMNH1621, 1627, 1631, 1632; left lunate, UGAMNH1633, 1634; left lunar, UGAMNH1625; left scapual spine, UGAMNH1635; rib head, UGAMNH1636; right scapula, UGAMNH1622; right P4, UGAMNH1623; left distal humerus, UGAMNH1624; left proximal femur, UGAMNH1626; right distal humeral epiphysis, UGAMNH1628; metatarsal diaphysial fragment, UGAMNH1629; left proximal tibial fragment, UGAMNH1630; tooth fragments, UF21239, 21281, 21282, 21285, 21288; distal humerus, UF21283; horn core tip, UF21284. Remarks—These elements are definitely bovid but the available material does not permit specific distinction. Bison bison (Linnaeus) Bison Material—-Right M2 , UGAMNH1620; left P3 , UGAMNH1619; left P2, UGAMNH1618; right M2 , UGAMNH1617; molar UF2299. Remarks—While Jones et al. (1992) have employed Bos bison for the American bison, we continue the traditional use of Bison bison. Two species of bison are known from Florida. The giant bison, B. latifrons, is known from Illinoian and Sangamonian and survived up until the late Wisconsinan. The American buffalo or bison, B. bison, was widespread throughout the Wisconsinan through the Recent (Kurten and Anderson 1980). Bison is typically associated with grasslands, though in the Southeast may well have ranged into woodlands (Golley 1962, Stock 1963). It became extinct in the southeastern United States early in the 19th Century. Bos taurus Linnaeus Cow Material—Right proximal humeral diaphysis, UGAMNH10863; left scapula, UGAMNH1155; left ilium, UGAMNH1152, 1154; right metatarsal diaphysis, UGAMNH117; right proximal humerus, UGAMNH1177; right distal humerus, UGAMNH1176; orbital portion of right maxilla, UGAMNH1 153; right proximal tibia, UGAMNH1171; right distal femoral epiphysis, UGAMNH1172; right astragalus, UGAMNH1173; distal phalanx, UGAMNH1174; metapodial, UF21300. Remarks—Bos taurus was introduced into North America sometime after 1492. All elements were poorly mineralized. The presence of cow indicates the site has modern contaminants. 34 Timothy S. Young and Joshua Laerm CLASS AVES Order Podicipediformes Family Podicipedidae Podiceps auritus (Linnaeus) Horned Grebe Material—Distal portion of left ulna, USNM209968. Remarks—Today the species winters in coastal areas and infrequently occurs in freshwater (Sprunt 1954). Podilymbus podiceps (Linnaeus) Pied-billed Grebe Material—Right humerus, USNM210293, 210294, 210301, 210302; humerus, USNM210311; right proximal humerus, USNM210304; left humerus, USNM210307; left tibial fragments, USNM210292, 210308, 210309, 210315, 210322, 210325, 210327; right tibia, USNM210297, 210300, 210312, 210316, 210317; left ulna, USNM210296, 210303, 210305, 210306, 210310, 210328, 210329; right ulna USNM210313, 210321; radius, USNM210298; right coracoid, USNM210319; left coracoid, USNM210320; left femur, USNM210298; right carpometacarpus, USNM210321; left carpometacarpus, USNM210323; scapula, USNM210324; pedal phalanx, USNM210326; tarsometatarsus, USNM210314, 210318; left tibia, USNM210306. Remarks—The species inhabits freshwater marshes and ponds, but also is associated with saltwater in winter (Sprunt 1954). Order Pelecaniformes Family Phalacrocoracidae Phalacrocorax auritus (Lesson) Double-crested Cormorant Material—Left radius, USNM209845; left ulna, USNM209844; scapula, USNM209859; anterior sternum, USNM209843; left coracoid, USNM209858; phalanx 1 of digit II, USNM209861, 209852; phalanx 2 of digit II, USNM209856; left humerus, USNM209846; right humerus, USNM209851; proximal radius, USNM209860; right ulna, USNM209852; distal tibia, USNM209848; right femur, USNM209854; right mandible, USNM209855; left mandible, USNM209850; sternal fragment, USNM209857; left coracoid, USNM209849; right coracoid, USNM209847. Remarks—This species is distributed in large rivers and lakes as well as brackish and saltwater systems (Sprunt 1954). St. Marks River Fauna 35 Order Ciconiformes Family Ardeidae Ardea herodias Linnaeus Great Blue Heron Material—Cervical vertebrae, USNM210282, 210283, 210285, 210287; right mandible, USNM210281; mandible fragments, USNM210280, 210286, 210288; maxilla fragment, USNM210279; right coracoid, USNM210291; right proximal humerus, USNM210289; distal tarsometatarsus, USNM210290; right carpometacarpus, USNM210279. Remarks—The great blue heron has wide ecological tolerances, occurring in freshwater swamps and riparian habitats as well as saltwater marshes (Sprunt 1954). Butorides striatus (Linnaeus) Green-backed Heron Material—Right humerus, USNM209966. Remarks—Butorides striatus and B. virescens, sometimes regarded as separate species, are recognized as geographic races of B. striatus by the American Ornithologists Union (1983). It occurs along lake margins, streams, ponds, and freshwater and saltwater marshes (Sprunt 1954). Egretta caerulea (Linnaeus) Little Blue Heron Material—Mandibular tip with right ramus, USNM209862. Remarks—Freshwater swamps and saltwater marshes are the preferred habitats (Sprunt 1954). Family Threskiornithidae Eudocimus albus (Linnaeus) White Ibis Material—Right humerus, USNM209971; left proximal coracoid, USNM209972. Remarks—Eudocibus albus is associated with swampy forests, marshy sloughs, and saltwater marshes (Sprunt 1954). Order Anseriformes Family Anatidae Aix sponsa (Linnaeus) Wood Duck Material—Right carpometacarpus, USNM209931, 209938, 209939, 209944; left carpometacarpus, USNM209934; right ulna, 36 Timothy S. Young and Joshua Laerm USNM209927, 209946; left ulna, USNM209926, 209940, 209945; left humerus, USNM209928, 209930, 209932, 209933; right humerus, USNM209941; radius, USNM209929, 209936; scapula, USNM209942, 209943; proximal tibia, USNM209935; right coracoid, USNM20992; right femur, USNM209925; right tarsometatarsus, USNM209937. Remarks—The species is common today in freshwater woodland rivers, ponds, and marshes (Sprunt 1954). Anas sp. indet. Material—Right ulna, UGAMNH2078. Anas acuta Linnaeus North Pintail Material—Left coracoid, USNM209965. Remarks—The pintail is associated with freshwater marshes, ponds, and lakes (Sprunt 1954). Anas americana Gmelin American Wigeon Material—Left humerus, USNM210270; left ulna, USNM210267, 210273; right scapula, USNM210278; scapula USNM210272, 210274; right coracoid, USNM210268, 210275, 210276; right ulna, USNM210277; phalanx 1 of digit II, USNM210269; radius, USNM210271. Remarks—This species is an inhabitant of freshwater marshes, ponds, and shallow lakes (Sprunt 1954). Anas discors Linnaeus Blue-winged Teal Material—Left carpometacarpus, USNM209865, 209866, 209872- 209874; right carpometacarpus, USNM209870, 209871; right humerus, USNM209869; right coracoid, USNM209868; left ulna, USNM209864. Remarks—Sprunt (1954) reports the species from freshwater ponds and lakes. Anas platyrhynchos Linnaeus Mallard Material—Left humerus, USNM209910, 209911, 209914; right humerus, USNM209913, 209918; right scapula, USNM209916; left scapula, USNM209917; left coracoid, USNM209919; right coracoid, USNM209912; right carpometacarpus, USNM209920; furcula, USNM209915. St. Marks River Fauna 37 Remarks—The mallard prefers freshwater lakes and marshes (Sprunt 1954). Aythya sp. indet. Material—Right carpometacarpus, UGAMNH2073; left distal tibiotarsus, UGAMNH2077. Aythya collaris (Donovan) Ring-necked Duck Material—Humerus shaft, USNM209899; left humerus, USNM209884, 209878, 209886, 209890, 209894, USNM209910; right humerus, USNM209877, 209893, 209903; left ulna, 209885, 209888, 209897; right ulna, USNM209878, 209892, 209905-209907; left tibia, USNM209880, 209908; right tibia, USNM209909; left carpometacarpus, 209898, 209900; right tarsometatarsus, USNM209881, 209889; left tarsometatarsus, USNM209887; tarsometatarsus, USNM209902; right coracoid, 209895, 209896; proximal radius, USNM209882; distal radius, USNM209883; radius, USNM209904; right scapula, USNM209891; cervical vertebra, USNM209901. Remarks—This species is associated most commonly with wooded lakes, ponds, and rivers, but also is reported from saltwater systems (Sprunt 1954). Branta canadensis (Linnaeus) Canada Goose Material—Right coracoid, UGAMNH2074; right tarsometatarsus, USNM209875; right distal carpometacarpus, USNM209876. Remarks—Both USNM specimens from the 1970s are noted by Storrs Olson (personal communication) as small and possibly represent either a small subspecies or juveniles. The UGAMNH specimen from 1987 is large. Sprunt (1954:53) states the center of abundance in Florida for modern Branta canadensis is the St. Marks Refuge. This coracoid could possibly be assigned to Branta cf. B. dickeyi on the basis of size. Steven Emslie (Point Reyes Bird Observatory, personal communication) examined the St. Marks River specimen and thought it could be assigned to B. dickeyi. Measurements of the coracoid are larger than modern B. canadensis, but there is some overlap. Emslie (personal communication) reported a large B. dickeyi from the early Pleistocene of Florida. We refer the coracoid conservatively to B. canadensis. The species prefers freshwater lakes, rivers, and marshes (Sprunt 1954). 38 Timothy S. Young and Joshua Laerm Bucephala albeola (Linnaeus) Bufflehead Material—Right carpometacarpus, USNM209969. Remarks—The bufflehead is most common in saltwater bays and estuaries, and rarely in freshwater lakes and ponds (Sprunt 1954). Lophodytes cucullatus (Linnaeus) Hooded Merganser Material—Right proximal humerus USNM209975; right distal humerus USNM209976, 209980; right humerus, USNM209977; left humerus, USNM209978; left ulna, USNM209979. Remarks—The species occurs in freshwater wooded ponds, rivers, and lakes (Sprunt 1954). Mergus merganser Linnaeus Common Merganser Material—Left distal tarsometatarsus, USNM209863. Remarks—The common merganser inhabits wooded freshwater rivers and ponds but winters in saltwater bays (Sprunt 1954). Order Falconiformes Family Accipitridae Pandion haliaetus (Linnaeus) Osprey Material—Right distal tarsometatarsus, USNM209967. Remarks—The species prefers fresh and saltwater marshes, lakes, and bays (Sprunt 1954). Buteo jamaicensis (Gmelin) Red-tailed Hawk Material—Left distal humerus, USNM209970. Remarks—The red-tailed hawk is most common in deciduous forests adjacent to open grasslands (Sprunt 1954). Order Galliformes Family Phasianidae Meleagris gallopavo Linnaeus Wild Turkey Material—Left tarsometatarsus, UGAMNH2075; right proximal tibiotarsus, USNM209921; right proximal femur, USNM209922; tarsometatarsus shaft, USNM209923. Remarks—The species is known from drier swamps, open pine, and hardwoods as well as prairies (Sprunt 1954). St. Marks River Fauna 39 Order Gruiformes Family Rallidae Fulica americana (Gmelin) American Coot Material—Left distal tarsometatarsus, UGAMNH2076; left tibia, USNM209947, 209951; left distal tibiotarsus, USNM209956, 209958; right distal tibiotarsus, USNM20949, 209952, 209962; right tibiotarsus, USNM209961; tibiotarsus shaft, USNM209955; left ulna, USNM209954, 209966; right carpometacarpus, USNM209950, 209963; right distal femur, USNM209948; distal humerus, USNM209953; right scapula, USNM209959; left coracoid, USNM209960. Remarks—The American coot is primarily associated with open freshwater ponds and marshes (Sprunt 1954). Gallinula chloropus (Linnaeus) Common Moorhen Material—Right tarsometatarsus, USNM210259, 210260, 210255; right tibiotarsus, USNM210257; radius, USNM210256; left phalanx 1 of digit II, USNM210258. Remarks—This species prefers freshwater marshes and ponds with heavy aquatic vegetation (Sprunt 1954). Family Aramidae Aramas guarauna (Linnaeus) Limpkin Material—Left tarsometatarsus, USNM210262, 210266; right tarsometatarsus, USNM210261; right distal tarsometatarsus, USNM210265; left distal tibiotarsus, USNM210263; right distal tibiotarsus, USNM210264. Remarks—The limpkin is associated with open, freshwater swamps and marshes (Sprunt 1954). Order Strigiformes Family Strigidae Strix varia Barton Barred Owl Material—Right proximal femur, USNM209973; right tibiotarsus shaft, USNM209974. Remarks—The barred owl occurs in low, wet woodlands and swampy forests (Sprunt 1954). 40 Timothy S. Young and Joshua Laerm CLASS REPTILIA Order Testudines Family Kinosternidae Kinosternidae gen. et sp. indet. Material—Nuchal, UGAMNH2038, 2047; right peripheral 1, UGAMNH2041; left peripheral 2, UGAMNH2042; left peripheral 4, UGAMNH2052; right peripheral 4, UGAMNH2044; left peripheral 9, UGAMNH2053; right peripheral 10, UGAMNH2048; plastron fragment, UGAMNH2050, 2051; right pleural 1, UGAMNH2039; left pleural 1, UGAMNH2045; right pleural 2, UGAMNH2043; right pleural 6, UGAMNH2040; pleural fragments, UGAMNH2046, 2049. Remarks—None of the kinosternid material could be referred to genus or species. Family Chelydridae Chelydra serpentina (Linnaeus) Snapping Turtle Material—Right peripheral, UGAMNH2034, 2037; left peripheral 4, UGAMNH2035; peripheral UGAMNH2036. Remarks—This material compares well with modern Chelydra serpentina. The species prefers permanent freshwater systems (Conant 1975). Family Emydidae Emydidae gen. et spec, indet. Material—Right epiplastron, UGAMNH1350, 1351, 1355, 1356, 1366, 1368, 1370, 1402, 1403, 1405, 1444, 1462, 1482, 1510, 1530, 1535, 1538, 1546, 1548-1550, 1554, 1872; left epiplastron, UGAMNH1235, 1268, 1282, 1285, 1307, 1343, 1345, 1346, 1349, 1359, 1362, 1375, 1380, 1390, 1394, 1445, 1511, 1551; left humerus, UGAMNH1498; right hypoplastron at inguinal notch, UGAMNH1217, 1237, 1241, 1242, 1281, 1308, 1316, 1322, 1324, 1325, 1357, 1379, 1382, 1418, 1419, 1467, 1469, 1501, 1601, 1871; right hypoplastron at axillary notch, UGAMNH1238, 1240, 1301, 1352, 1358, 1404, 1452, 1474, 1503, 1547, 1553, 1555, 1565; right hypoplastron, UGAMNH1164, 1167, 1215, 1216, 1219, 1221, 1236, 1283, 1333, 1344, 1354, 1376, 1388, 1389, 1429, 1433, 1470, 1473, 1480, 1495, 1521, 1533, 1572, 1575, 1584, 1600, 1867, 1870; left hypoplastron at axial notch, UGAMNH1290, 1361, 1369, 1566, 1602; left hypoplastron at inguinal notch, UGAMNH1168, 1220, 1269, 1280, 1284, 1300, 1302, 1413, 1447, 1449, 1516, 1522, 1527, 1559, 1567, 1582; left hypoplastron, UGAMNH1218, 1222-1225, 1239, 1278, 1279, 1293, 1320, 1342, 1377, 1396, 1423, 1456, 1464, 1471, St. Marks River Fauna 41 1475, 1509, 1513, 1519, 1526, 1532, 1560, 1564, 1569, 1571, 1574, 1578, 1583; neural 1, UGAMNH1328, 1439, 1545, 1558; neural 2, UGAMNH1568, UGAMNH1588; neural 3, UGAMNH1573, 1581; neural 6, UGAMNH1461, 1577; neural 7, UGAMNH1260, 1341, 1579, 1874; neural 8, UGAMNH1271; neural 9, UGAMNH1410; neural, UGAMNH1233, 1258, 1259, 1277, 1291, 1309, 1310, 1312, 1363, 1364, 1372, 1384, 1392, 1393, 1406, 1409, 1414, 1440, 1441, 1494, 1524, 1563, 1570, 1576, 1580, 1589, 2139, 2141; nuchal, UGAMNH1261- 1264, 1313, 1411, 1417, 1427, 1457, 1486, 1504, 1508, 1518, 1595; right periphal 1, UGAMNH1165, 1321, 1399, 1398, 1451, 1489, 1525, 1528, 1592, 1866; left peripheral 1, UGAMNH1231, 1245, 1303, 1319, 1454, 1505, 1562; right peripheral 2, UGAMNH1381, 1397, 1407; left peripheral 2, UGAMNH1275, 1298, 1400; right peripheral 3, UGAMNH1169, 1294, 1442, 1531; left peripheral 3, UGAMNH1416, 1421, 1472; right peripheral 4, UGAMNH1540; left peripheral 4, UGAMNH2167; right peripheral 5, UGAMNH1395; left peripheral 5, UGAMNH1425, 1591; right peripheral 6, UGAMNH1244, 1517; left peripheral 6, UGAMNH1274, 1296, 1446; right peripheral 7, UGAMNH1246, 1428, 1594; left peripheral 7, UGAMNH1329, 1432, 1453, 1455; right peripheral 8, UGAMNH1232, 1552, 2143; left peripheral 8, 1424, 1542; right peripheral 9, UGAMNH1166, 1373, 1484, 1490; left peripheral 9, UGAMNH1249, 1299, 1492; right peripheral 10, H1273, 1276; left peripheral 10, UGAMNH1248; right peripheral 11, UGAMNH1326, 1332, 1429, 1587, 1597; left peripheral 11, UGAMNH1297, 1304, 1408, 1435; peripheral UGAMNH1000, 1243, 1247, 1311, 1420, 1426, 1434, 1442, 1449, 1450, 1536, 1593, 1604; right pleural 1, UGAMNH1234, 1334, 1336, 1374, 1385, 1437, 1485, 1493, 1502, 1554, 1875; left pleural 1, UGAMNH1365, 1371, 1378, 1391, 1438, 1468, 1487, 1507, 2142; left pleural 2, UGAMNH1292; right pleural 2, UGAMNH1465, UGAMNH1491; left pleural 3, UGAMNH1431; right pleural 3, UGAMNH1436, UGAMNH1430; left pleural 4, UGAMNH1460; right pleural 5, UGAMNH1596; left pleural 5, UGAMNH1492; right pleural 6, UGAMNH1488; right pleural 7, UGAMNH1340; Pleural, UGAMNH1265-1267, 1270, 1286-1289, 1295, 1305, 1306, 1314, 1318, 1330, 1339, 1348, 1352, 1367, 1383, 1387, 1401, 1483, 1537, 1543, 1585, 1586, 1590, 1598, 1599, 1603, 1868, 1873; pygal, UGAMNH1317, 1323, 1422, 1458, 1476, 1556, 1561; left scapula, UGAMNH1496, 1497; right scapula, UGAMNH1665; suprapygal, UGAMNH1499; right xiphiplastron, UGAMNH1226- 1230, 1250, 1251, 1253, 1255, 1257, 1331, 1347, 1360, 1415, 1463, 1472, 1479, 1481, 1869, 1876; left xiphiplastron, UGAMNH1252, 1254, 1256, 1315, 1335, 1337, 1338, 1386, 1512, 1514, 1515, 1523, 1539, 1544, 1459, 1478. 42 Timothy S. Young and Joshua Laerm Remarks—Most of the emydid material could only be identified to the familial level. Species level identification is difficult and requires nearly complete elements. Almost all the material was well mineralized. We are confident that the majority represents Pleistocene deposition as opposed to Recent. Pseudemys concinna (LeConte) River Cooter Material—Left peripheral 3, UGAMNH1882; left peripheral 4, UGAMNH1885; right peripheral 7, UGAMNH1884; right peripheral 11, UGAMNH1883. Remarks—Pseudemys concinna is distinguished by its distinctive carapace. It is most common in slow streams and rivers (Conant 1975). Pseudemys floridana (LeConte) Cooter Material—Left peripheral 3, UGAMNH2030; left pleural 3, UGAMNH2031; left pleural 4, UGAMNH2033; nuchal, UGAMNH2032. Remarks—The species is most commonly associated with permanent bodies of freshwater including swamps and rivers (Conant 1975). Pseudemys nelsoni Carr Florida Redbelly Turtle Material—Entoplastron, UGAMNH1904; right epiplastron, UGAMNH1920; right hypoplastron axial notch, UGAMNH1889, 1913, 1938; left hypoplastron axial notch UGAMNH1928, 1940; right hypoplastron inguinal notch, UGAMNH1908, 1943; left hypoplastron inguinal notch, UGAMNH1897, 1905; neural 7, UGAMNH1901; neural, UGAMNH1887; nuchal, UGAMNH1914, 1953; right peripheral 1, UGAMNH1899, 1906; right peripheral 2, UGAMNH1929; right peripheral 3, UGAMNH1937; left peripheral 3, UGAMNH1917; right peripheral 4, UGAMNH1930; left peripheral 5, UGAMNH1900; left peripheral 7, UGAMNH1890, 1898; right peripheral 8, UGAMNH1942, 1950; left peripheral 8, UGAMNH1945, 1948; right peripheral 9, UGAMNH1915; left peripheral 9, UGAMNH1886; left peripheral 10, UGAMNH1946, 1947; right peripheral 11, UGAMNH1506, 1918, 1919, 1941; left peripheral 11, UGAMNH1944; peripheral, UGAMNH1907, 1909, 1949; right pleural 1, UGAMNH1534, 1892, 1910; left pleural 1, UGAMNH1922, 1951, 2140; left pleural 2, UGAMNH1917; left pleural 3, UGAMNH1912; left pleural 4, UGAMNH1933; right pleural 5, UGAMNH1934; pleural, UGAMNH1891, 1893-1895, 1903, 1916, St. Marks River Fauna 43 1921, 1923-1927, 1931, UGAMNH1932, 1935, 1936, 1939; suprapygal, UGAMNH1888; right xiphiplastron, UGAMNH1896; left xiphiplastron UGAMNH1902, 1952. Remarks—This is a species associated with freshwater sloughs, marshes, streams, and ponds (Conant 1975). Trachemys scripta (Schoepff) Slider Material—Entoplastron, UGAMNH1763, 1792, 1801, 1831; right hypoplastron axial notch, UGAMNH1780; left hypoplastron axial notch, UGAMNH1828, 1833; right hypoplastron inguinal notch, UGAMNH1774; left hypoplastron inguinal notch, UGAMNH1819; neural 1, UGAMNH1790, 1834; neural 3, UGAMNH1789; neural 8, UGAMNH1846; neural, UGAMNH1766, 1769, 1819, 1837, 1842; nuchal, UGAMNH1520, 1764, 1773, 1776, 1778, 1782, 1784, 1787, 1791, 1823, 1841, 1844; right peripheral 1, UGAMNH1768, 1770, 1826, 1827; left peripheral 1, UGAMNH1783, 1840; right peripheral 2, UGAMNH1765, 1798, 1802; left peripheral 2, UGAMNH1776, 1806; right peripheral 3, UGAMNH1794; left peripheral 3, UGAMNH1796; left peripheral 5, UGAMNH1775; left peripheral 8, UGAMNH1835, 1836, right peripheral 9, UGAMNH1779, 1793; left peripheral 10, UGAMNH1767, 1820, 1839; right peripheral 11, UGAMNH1781, 1843; left peripheral 11, UGAMNH1762, 1803-1805, 1816, 1825, 1832, 1845; peripheral, UGAMNH1785, 1824, 1829; left pleural 1, UGAMNH1807; right pleural 2, UGAMNH1799; left pleural 2, UGAMNH1800; left pleural 4, UGAMNH1788; pleural, UGAMNH1771, 1772, 1786, 1808- 1815, 1821, 1822, 1838; pygal, UGAMNH1795, 1797, 1818, 1830. Remarks—This material has the distinctive sculpted appearance of Pleistocene Trachemys scripta. All the material is well mineralized. It occurs in freshwater ponds, streams, and rivers (Conant 1975). Terrapene Carolina (Linnaeus) Eastern Box Turtle Material—Right and left epiplastron, UGAMNH1703; left hypoplastron, UGAMNH2144; left and right hypoplastron and xiphiplastron, UGAMNH1697, 1698; right hypoplastron at hinge, UGAMNH1686, 1727; left hypoplastron at hinge, UGAMN1685, 1687, 1690, 1705, 1731; right hypoplastron at inguinal notch, UGAMNH1713; right hypoplastron, UGAMNH1714; left and right hypoplastron, UGAMNH1696, 1715; hypoplastron, UGAMNH1716; neural 1, pleural and peripheral 1 and 2, UGAMNH1699; neural 1 and left and right peripheral 1, UGAMNH1732; neural 5 and 6, UGAMNH1730; neural, UGAMNH1707; nuchal, UGAMNH1704, 1725; right peripheral 1 and 44 Timothy S. Young and Joshua Laerm 2, UGAMNH1728; left peripheral 1 and 2, UGAMNH1726; right peripheral 1, 2 and 3, UGAMNH1692; left peripheral 1, 2, 3 and pleural 1, UGAMNH1720; left peripheral 3, UGAMNH1722; left peripheral 3 and 4, UGAMNHA1721; right peripheral 3 and 4, UGAMNH1708; right peripheral 5, UGAMNH1688, 1706; left peripheral 5, UGAMNH1691; right peripheral 6 and 7 and pleural 4 and 5, UGAMNH1710; right peripheral 6, 7, and 8, UGAMNH1712; left peripheral 7, UGAMNH1684; left peripheral 8, UGAMNH1702; left peripheral 8, 9, and 10, UGAMNH1694; right peripheral 9, 10, and 11, UGAMNH1733; left peripheral 10, UGAMNH1689; right peripheral 10 and 11, UGAMNH1695, UGAMNH1718; left peripheral 10 and 11, UGAMNH1719, 1734; right peripheral 10 and 11 and pygal, UGAMNH1711; left peripheral 11, UGAMNH1701; left peripheral 11 and pygal, UGAMNH1709; left and right peripheral 11 and pygal, UGAMNH1717; right peripheral 11, UGAMNH1724; left pleural 2 and peripheral 4 and 5, UGAMNH1700; pygal, UGAMNH1683, UGAMNH1723; left xiphiplastron, UGAMNH1327, 1729; left and right xiphiplastron, UGAMNH1693. Remarks—Terrapene Carolina can be distinguished from its extinct relative T. Carolina putnami based on smaller size. It is a terrestrial woodland species (Conant 1975). Terrapene Carolina putnami Hay Giant Box Turtle Material—Right epiplastron, UGAMNH1860; left hypoplastron at inguinal notch, UGAMNH1855; left hypoplastron and epiplastron and entoplastron, UGAMNH1863; right hypoplastron and xiphiplastron, UGAMNH1864; neural 1 and pleural and peripheral 1, UGAMNH1856; nuchal, UGAMNH1865; left peripheral 3 and 4, UGAMNH1858; right peripheral 6 and 7, UGAMNH1859; right peripheral 10 and 11 and pygal, UGAMNH1862; right peripheral 1, UGAMNH1849; left peripheral 2, 3, and 4, UGAMNH1848; left peripheral 4, 5, and 6, UGAMNH1861; right peripheral 6, UGAMNH185; left peripheral 7 and 8, UGAMNH1852; left peripheral 8 and 9, UGAMNH1853; right peripheral 9, UGAMNH1847; left peripheral 10 and 11, UGAMNH1854; left pleural 2 and 3 and peripheral 4 and 5, UGAMNH1857; right pleural 2 and 3, UGAMNH1850. Remarks—This extinct giant subspecies is common in late Pleistocene deposits of Florida where it occurred in coastal marshes and lowland savannahs. (Auffenberg 1958, Kurten and Anderson 1980). It is readily distinguishable on the basis of its large size. St. Marks River Fauna 45 Family Testudinidae Testudinidae gen. et sp. indet. Material—Osteoderms, UGAMNH1645, UGAMNH1646. Remarks—These specimens represent a large tortoise, but the osteoderms are not diagnostic. Geochelone sp. indet. Material—Pleural, UGAMNH1638; left hypoplastron, UGAMNH1639; right pleural 2, UGAMNH1640; left pleural 4, UGAMNH1641. Remarks—The available material, while certainly Geochelone, could not be referred to a species with confidence. Geochelone incisa (Hay) Material—Right peripheral 7, UGAMNH1642; nuchal UGAMNH1643; right peripheral 5, UGAMNH1644. Remarks—This material compares well with the series of G. incisa in the collections of the Florida Museum of Natural History and corresponds to Auffenberg's (1963) description. The was apparently an open grassland inhabitant thought to require a frost free winter (Kurten and Anderson 1980); however, Martin and Guilday (1967) disagree. Gopherus polyphemus (Daudin) Gopher Tortoise Material—Nuchal, UGAMNH1637. Remarks—This material compares well with modern Gopherus polyphemus which ranges in dry sandy soils (Conant 1975). Family Trionychidae Trionyx sp. indet. Material—Carapacial fragment, UGAMNH1761. Remarks—The available material, while certainly Trionyx because of the distinctive pattern on the bone, could not be referred to a species with confidence. Order Squamata Family Colubridae Colubridae gen. et spec, indet. Material—Vertebrae, UGAMNH2054-2061. 46 Timothy S. Young and Joshua Laerm Elaphe obsoleta (Say) Rat Snake Material—Vertebra, UGAMNH2055. Remarks—This material compares well with modern Elaphe obsoleta which may be found in woodlands and grasslands (Conant 1975). Order Crocodilia Family Alligatoridae Alligator mississippiensis (Daudin) American Alligator Material—Left angular, UGAMNH1015; distal phalanx, UGAMNH1001, right dentary (without teeth), UGAMNH1012; dermal scutes, UGAMNH1003-1011 (1010 and 1011 exhibit crossmends), UGAMNH1023; right femur, UGAMNH1020, 1022; left humerus, UGAMNH1019; fused parietals, UGAMNH1025; left scapula, UGAMNH1014; right scapula, UGAMNH1016, 1018; teeth, UGAMNH1002, 1024, 1026, 1028, 1029; vertebra, UGAMNH1017; frontal, UGAMNH1013; left jugal, UGAMNH1021. Remarks—This material has the distinctive Alligator mississippiensis morphology and it compares well with modern examples. Alligators occur in both fresh and brackish waters (Conant 1975). CLASS AMPHIBIA Order Caudata Family Sirenidae Siren sp. indet. Material—Vertebrae, UGAMNH2129-2131, 2161. Remarks—The available material compares well with modern Siren. Order Anura Anura gen. et sp. indet. Material—Vertebrae, UGAMNH2132-2134-right humerii. Remarks—The available material, while certainly frog, could not be referred to a genus or species with confidence. CLASS OSTEICHTHYES Order Lepisosteiformes Family Lepisosteidae Lepisosteus sp. indet. Material—Scales, UGAMNH2109-21 11. St. Marks River Fauna 47 Remarks—The scales, while certainly Lepisosteus, could not be referred to a species with confidence. Lepisosteus occurs in freshwater and estuarine habitats (Hoese and Moore 1977, Lee et al. 1980). Order Amiiformes Family Amiidae Amia calva Linnaeus Bowfin Material—Left dentary, UGAMNH2088; left frontal, UGAMNH2089; cervical vertebra, UGAMNH2090. Remarks—This material compares well with modern specimens of Amia calva. The bowfin is a freshwater and estuarine species (Hoese and Moore 1977, Lee et al. 1980). Order Siluriformes Family Ictaluridae Ictaluridae gen. et sp. indet. Material—Spine, UGAMNH2112; vertebra, UGAMNH2113. Remarks—The available material, while certainly catfish, could not be referred to a genus or species with confidence. Pylodictis cf. P. olivaris (Rafinesque) Flathead Catfish Material—Left proximal coracoid, UGAMNH2119. Remarks—The morphology of the single element is very similar to modern specimens of P. olivaris and distinct from the other known regional ictalurids available for comparison. The specimen at hand shows some evidence of mineralization, but mineralization is not extensive. The species occurrence in the St. Marks River is outside its reported range which extends from northeastern Mexico east throughout Gulf of Mexico drainages to Mobile Bay (Lee et al. 1980 et seq.). However, in recent times the species has undergone introductions and populations are now known from at least the Appalachicola-Chatahoochee System (M. and B. J. Freeman, University of Georgia, personal communication). Uyeno and Miller (1962) reported some specimens of P. olivaris from the Trinity River Terrace, Texas. The deposit was dated to the Sangamon (late Pleistocene); however, that site is within the present range of the species. It is a freshwater species (Hoese and Moore 1977, Lee et al. 1980). 48 Timothy S. Young and Joshua Laerm Family Ariidae Ariidae gen. et sp. indet. Material—Spine, UGAMNH2114-2116; cervical vertebrae, UGAMNH2117, UGAMNH2118. Remarks—These specimens show the characters of the marine catfishes, although species identification is not possible. Ariusfelis (Linnaeus) Hardhead Catfish Material—Spine, UGAMNH2091. Remarks—This spine compares well with the distinctive Arius felis morphology. This species is restricted to saltwater and estuaries (Hoese and Moore 1977, Lee et al. 1980). Order Salmoniformes Family Esocidae Esox sp. indet. Material—Right dentary, UGAMNH2101, left dentary UGAMNH2095, 2096, 2098-2100, 2102-2105; dentary, UGAMNH2097; parasphenoid, UGAMNH2106; pharyngeal grinding plates, UGAMNH2107, 2108. Remarks—These specimens closely resemble both E. americanus Gmelin and E. niger Lesueur. Both are considered freshwater species (Lee et al. 1980 et seq.) and occur in regional waters today. Order Perciformes Family Percichthyidae Morone saxatilis (Walbaum) Striped Bass Material—Right maxilla, UGAMNH2082; right premaxilla, UGAMNH2083; right quadrate, UGAMNH2084, 2085; left quadrate, UGAMNH2086; atlas, UGAMNH2087. Remarks—This material compares well with modern examples of Morone saxatilis which occurs in both coastal saltwater and estuaries (Hoese and Moore 1977, Lee et al. 1980). Family Sparidae Archosargus probatocephalus (Walbaum) Sheepshead Material—Right dentary, UGAMNH2079; left preoperculum, UGAMNH2080; tooth, UGAMNH2081. Remarks—This material compares well with modern examples of St. Marks River Fauna 49 Archosargus probatocephalus. The sheepshead is a coastal salt-water and estuary species (Hoese and Moore 1977, Lee et al. 1980). Family Sciaenidae Sciaenops ocellatus (Linnaeus) Red Drum Material—Quadrate, UGAMNH2092. Remarks—This material compares well with modern examples of Sciaenops ocellatus. It is a coastal saltwater species, but is also associated with estuaries (Hoese and Moore 1977, Lee et al. 1980). Family Mugilidae Mugil sp. indet. Material—Vertebrae, UGAMNH2093, UGAMNH2094. Remarks—The available material, while certainly Mugil, could not be referred to a species with confidence. Mugil is a coastal saltwater species (Hoese and Moore 1997, Lee et al. 1980). RESULTS AND DISCUSSION Chronology and Environment of Deposition Of several thousand separate skeletal elements recovered from the St. Marks River, 1,162 were referable to specific taxa. Included are 37 species of mammals, 3 birds, 13 reptiles, 2 amphibians, and 9 fish. An additional 23 species of birds were identified from the 1972 collection made by Storrs Olson. Of all species we reported, 14 mammals and 2 reptiles are restricted to the Pleistocene. The remaining are representative of the modern extant regional fauna. With the exception of modern contaminants, the latter are acceptable Pleistocene species; however, they more probably represent a mixture of Holocene and Pleistocene material. This is reflected in the range of mineralization observed in many species. In all cases those species known only from the Pleistocene are well mineralized. However, several species with both a Pleistocene and Recent occurrence such as horse and deer exhibit both well mineralized and, what appears to be, very recent unmineralized condition. Modern contaminants such as cow and pig are unmineralized. In general, mineralization is no criterion of Pleistocene deposition. The problem of apparent heterochronous deposition and separation of Pleistocene and Holocene materials is exacerbated by the apparent rapid mineralization that can occur in reducing environments. Neill (1957) noted that rapid mineralization of organic remains in Florida creates the illusion that Recent material is of older age. Nonetheless, the St. Marks River 50 Timothy S. Young and Joshua Laerm fauna is clearly mixed and reflects heterochronous deposition over time beginning no later than the late Pleistocene (Wisconsinan) and extending through the Recent. We compared the St. Marks River faunal list and a modern regional faunal list of the Apalachicola River system (Means 1976). Of the 344 species listed by Means, 29% of the mammals, 10% of the birds, 19% of the reptiles, 5% of the amphibians, and 2% of the fish are represented in the St. Marks River fauna. This bias toward mammals probably reflects taphonomic factors associated with the larger size of mammalian elements in a fluvial environment. Small, more fragile vertebrates (birds, reptiles, amphibians, and fish) are clearly under-represented in the St. Marks River fauna. This bias is reflected also in the mammalian fauna where chiropteran, insectivoran, and small rodent remains are conspicuously absent. While many of the species recovered from the St. Marks River are eurytopic and provide only limited information regarding the environment of deposition, a number are stenotopic and are considered good environmental indicators. Mammals—The mammalian fauna, in particular, is very useful in assessing the chronology and paleoenvironment of the St. Marks River. The reason for this is two-fold. First, mammals are the most numerous and have the largest component of extinct forms. Second, Florida has an extremely rich and well-documented late Pleistocene as well as modern mammalian fauna upon which comparisons to the St. Marks River fauna can be made. Thirteen (35%) of the mammalian fauna of the St. Marks River is represented by extinct forms. These include Holmsina septentrionalis, Megalonyx jeffersonii, Glossotherium harlani, Canis dims, Smilodon sp., Synaptomys australis, Tapirus, sp., Equus sp., Platygonus compressus, Hemiauchenia macrocephala, Paleolama mirifica, Mammut americanum, and Mammuthus jeffersonii. This closely approximates the relative percentage of extinct mammals from a number of Rancholabrean faunas from elsewhere in Florida (Martin and Webb 1974). The temporal span of the extinct forms ranges from Blancan through Recent. However, they all share a late Wisconsinan chronology. Those species representing extant forms, although individually some exhibit a longer stratigraphic history, also share a late Wisconsinan chronology. With few exceptions, all the extant species are represented in the local fauna today. Comparison of the known and inferred habitat preferences or requirements of the extant and extinct mammalian species suggests the depositional environment was heterogeneous. On one hand there are a number of essentially woodland species: Didelphis, Holmsina, St. Marks River Fauna 51 Megalonyx, Lutra, Mephitis, Urocyon, Ursus, Tapirus, Platygonus, Odocoileus, and Mammut. However, grassland species are well represented also: Glossotherium, Mephitis, Geomys, Equus, Hemiauchenia, Paleolama, Bison, and Mammuthus. From a simple listing it might appear that grassland species are about as common as woodland species. However, when compared by the number of identified specimens per taxon, woodland species are more prevalent. Despite criticism, this method is reliable for a comparison of relative abun-dances of species (Grayson 1984). In addition, a number of species indicate proximity of water: Didelphis, Lutra, Procyon, Ursus, Castor, Neofiber, Ondatra, Synaptomys, and Tapirus are all typically associated with moist, riparian, or standing water habitats. Birds—Storrs Olson's collection from the St. Marks River have never been published. He was kind enough to provide a list of the birds identified and has permitted us to include it in the present discussion. Olson (personal communication) felt that "there was very little of interest among the birds" mainly because the list of avian species recovered from the St. Marks River is essentially similar to the modern fauna (Means 1976). As a whole, birds are uninstructive concerning the dating of the St. Marks River fauna. They do, however, provide considerable information relating to the environment of deposition. The St. Marks River avian fauna is clearly biased toward large species with predominantly salt and freshwater marshland habitat preferences: Podiceps, Podilymbus, Phalacrocorax, Ardea, Butorides, Egretta, Eudocimbus, Aix, Anas, Aythya, Branta, Bucephala, Lophodytes, Padion, Fulica, Gallinula, and Aramus. In addition, a number of the species are typically associated with woodlands or woodland riparian habitats: Aix, Mergus, Buteo, and Strix. Conspicuously absent are the passeriforms. This probably represents the taphonomic bias referred to above. While a significant number of the birds are often present in saltwater marsh habitats, there are no shorebird (charadriform) species present. Reptiles and Amphibians—Many turtles, but few other reptiles, are reported from the St. Marks River. Emydid turtles, in particular, are well represented and make up approximately 90% of the recovered reptilian material. In fact, in numbers alone they make up well over one third the individual elements in the fauna. The emydid turtle species identified from the 1987 collection were Pseudemys concinna, P. floridanus, P. nelsoni, Trachemys scripta, and Terrapene Carolina, all of which are found in the area today. Pseudemys and Trachemys are indicative of a freshwater environment, while Terrapene is terrestrial. An extinct, large, late Pleistocene subspecies of Terrapene 52 Timothy S. Young and Joshua Laerm Carolina, T. c. putnami, is represented in the St. Marks River fauna by a number of elements. It was probably limited to the Coastal Plain and Savannah habitats (Auffenberg 1958) and is represented in many late Pleistocene sites in Florida. Other aquatic turtles recovered include one chelydrid, C. serpentina, and a number of unreferrable kinosternid fragments. Terrestrial testudinoid turtles present at the site are Geochelone incisa, Geochelone sp., and Gopherus polyphemus. Geochelone incisa represents a definite late Pleistocene species, as does Terrapene Carolina putnami. Gopherus polyphemus occurs in the area today. Only two snakes, Nerodia sp. and Elaphe obsoleta, were identified from the 1987 collection. Both snakes occur in the area today. No lizards were identified from any of the fossil collections. Two amphibians were recovered, one caudate and one anuran, neither of which could be identified to species. With the exception of the two late Pleistocene components, the herpetofauna is representative of the modern regional fauna and includes both lower Coastal Plain riverine and marshland species, as well as terrestrial forms. Fishes—The fish fauna described includes both freshwater and marine forms. Ariopsis felis, Morone saxatilis, Archosargus probatocephalus, Sciaenops ocellata, and Mugil sp. although typically marine are also estuarine tolerant. The freshwater fishes include Pylodictis cf. P. olivaris, Lepisosteus sp., Esox sp., and Amia calva. Of these, Lepisosteus sp., Esox sp., and A. calva tolerate estuarine, but not marine, conditions (Hoese and Moore 1977). In conclusion, the aquatic community suggests a mixed freshwater and marine, or more likely an estuarine environment, similar to the lower half of the St. Marks River drainage today. The terrestrial fauna indicates a wooded riparian environment also similar to that found in the St. Marks River drainage today. However, the presence of Hemiauchenia, Bison, Equus sp., and Mammumthus coupled with Geomys, Geochelone, and Gopherus suggests that more open, semi-forested savannah habitats were also represented. This is consistent with other late Pleistocene (Rancholabrean) faunas from the panhandle of Florida, some of which are considered below. Faunal Comparison The Chipola River sites (IA and HA)—This is a river deposit similar to the St. Marks River and contains similar species including Didelphis virginiana, Holmesina septentrionalis, Castor canadensis, Procyon lotor, Bison sp., Equus sp., Mammut americanum, Odocoileus virginianus, and Hemiauchenia macrocephala (Webb St. Marks River Fauna 53 1914a). Although no formal paleontological description of the site exists, the species present in that assemblage indicate a mixed woodland/grassland environment (Webb 1974a). The Aucilla River IA site—The site is also similar to the St. Marks River in depositional and temporal characters. No published paleontological description exists for this site either, but from the fauna a habitat of woodland and marsh can be assumed. It includes Didelphis virginiana, Holmesina septentrionalis, Glossotherium cf. G. harlani, Ondatra zibethicus, Castor canadensis, Neochoerus pinckneyi, Sylvilagus floridanus, Canis dims, and Tremarctos floridanus (Webb 1974a). Wakulla Springs—This, too, is similar to the St. Marks River in depositional and temporal characters. Included are Mammuthus sp., Mammut americanum, and Bison bison antiquus (Webb 1974a). No formal paleontological description of the site exists. Generally there are only slight differences between the St. Marks River and other Florida panhandle, riverine deposits. These differences can probably be attributed to a number of causes including collection by amateurs, undersampling, taphonomic events, or other collecting biases. Compared to the other Rancholabrean faunas from peninsular Florida (Martin and Webb 1974, Webb 1974a, Webb and Wilkins 1984), the St. Marks River assemblage probably is not representative of the full late Pleistocene fauna that existed in the area. For example, more than 50 species of mammals are known to have been present in Florida during the time of accumulation of the Ichetucknee River fauna, Columbia County, Florida (Martin and Webb 1974). As shown by Martin and Webb (1974) mammalian faunal diversity was considerably elevated in peninsular Florida during Rancholabrean time, and it is highly likely that is was the case along the rich fluvio-estuarine environment of the panhandle during the same period. ACKNOWLEDGMENTS—Thanks are due Chris McKensie, Locke Rogers, Brad Newsom, Tim Gaudin, and Luis Insignares for their efforts below and above the surface of the St. Marks River. Chris McKensie deserves added thanks for the many long hours he spent screening and sorting bones. The United States Department of Agriculture provided an advance copy of their publication on the soils of Wakulla County. Gary Morgan and Russ McArty at the Florida Museum of Natural History were more than just helpful. The gracious offer of Storrs Olson to make available his unpublished records of the avian material he collected and analyzed is greatly appreciated. Robert Martin, Robert Frey, and Elizabeth Reitz provided 54 Timothy S. Young and Joshua Laerm many helpful critical comments on earlier drafts. Funds for this study were provided through Department of Zoology and the Museum of Natural History, the University of Georgia. LITERATURE CITED American Ornithologists Union. 1983. Check-list of North American birds. Sixth edition. Allen Press, Lawrence, Kansas. Auffenberg, W. 1958. Fossil turtles of the genus Terrapene in Florida. Bulletin of the Florida State Museum 3:53-92. Auffenberg, W. 1963. Fossil Testudinine turtles of Florida, genera Geochelone and Floridemys. Bulletin of the Florida State Museum 7:54-97. Brodkorb, P. 1963. Catalogue of fossil birds, Part 1 (Archaeopterygiformes through Ardeiformes). Bulletin of the Florida State Museum 7:179- 293. Burt, W. H. 1928. Additional notes on the life history of the Goss lemming mouse. Journal of Mammalogy 9:212-216. Collins, J. T. 1990. Standard common and current scientific names for North American amphibians and reptiles. Third edition. Society for the Study of Amphibians and Reptiles Herpetological Circular 19:1-41. Conant, R. 1975. A field guide to the reptiles and amphibians of eastern/ central North America. Houghton Mifflin, Boston, Massachusetts. Cooke, C. W. 1939. Scenery of Florida interpreted by a geologist. Bulletin of the Florida Geological Survey 17:1-118. Dreimanis, A. 1968. Extinction of mastodons in eastern North America: testing a new climatic environmental hypothesis. Ohio Journal of Science 68:257-72. Frazier, M. F. 1977. New records of Neofiber leonardi (Rodentia, Cricetidae) and the paleoecology of the genus. Journal of Mammalogy 58:368-73. Gardner, A. L. 1973. The systematics of the Genus Didelphis (Marsupialia, Didelphidae) in North and Middle America. Special Publication of the Museum Texas Tech University 4:1-81. Gillette, D. D. 1976fl. A new species of small cat from the Late Quaternary of the southeastern United States. Journal of Mammalogy 57:664-676. Gillette, D. D. 19766. Late Quaternary mammals from the St. Marks River, Wakulla County, Florida. Florida Scientist 39:120-122. Golley, F. B. 1962. Mammals of Georgia. University of Georgia Press, Athens. Gray, S. W., and H. R. Crammer. 1961. A tapir mandible from a northwest Georgia cave. Bulletin of the Georgia Academy of Science 19:83-90. Grayson, D. K. 1984. Quantitative zooarchaeology. Academic Press, New York, New York. Guilday, J. E., A. D. McCrady, and H. W. Hamilton. 1969. The Pleistocene St. Marks River Fauna 55 vertebrate fauna of Robinson cave, Overton County, Tennessee. Paleovertebrata 1969:25-75. Harper, R. M. 1914. Geology and vegetation of northern Florida. Bulletin of the Florida Geological Survey 6:163-416. Harrington, C. R., H. W. Tipper, and R. J. Mott. 1974. Mammoth from Babine Lake, British Columbia. Canadian Journal of Earth Science 11:285-303. Hay, O. P. 1923. The Pleistocene of North America and its vertebrated animals from the states east of the Mississippi and from the Canadian provinces east of longitude 95. Carnegie Institute of Washington, Publication Number 32. Hendry, C. W., Jr., and C. R. Sproul. 1966. Geology and ground-water resources of Leon County. Bulletin of the Florida Geological Survey 47:1-178. Hibbard, C. W. 1943. The Rezabek Fauna, a new Pleistocene fauna from Lincoln County, Kansas. University of Kansas Scientific Bulletin 29:235-247. Hoese, H. D., and R. H. Moore. 1977. Fishes of the Gulf of Mexico, Texas, Louisiana, and adjacent waters. Texas A&M University Press, College Station. Jones, J. K., Jr., R. S. Hoffmann, D. W. Rice, C. Jones, R. J. Baker, and M. D. Engstrom. 1992. Revised checklist of North American mammals north of Mexico, 1991. Occasional Papers of the Museum Texas Tech. University 146:1-23. Kurten, B., and E. Anderson. 1980. Pleistocene mammals of North America. Columbia University Press, New York, New York. Lee, D. S., C. R. Gilbert, C. H. Hocutt, R. E. Jenkins, D. E. McAllister, and J. R. Stauffer, Jr. 1980. Atlas of North American freshwater fishes. North Carolina Biological Survey. Leidy, J. 1870. Narrative for a short visit to Boston and Cambridge. Proceedings of the Academy of Natural Sciences, Philadelphia 1870:96-99. Lowery, G. H. 1974. The Mammals of Louisiana and its adjacent waters. Louisiana State University Press, Baton Rogue. Lundelius, E. L., Jr., R. W. Graham, E. Anderson, J. Guilday, J. A. Holman, D. W. Steadman, and S. D. Webb. 1983. Terrestrial vertebrate faunas. Pages 311-353 in Late Quaternary environments of the United States (S. C. Porter, editor). University of Minnesota Press, Minneapolis. Martin, P. S., and J. E. Guilday. 1967. A bestiary for Pleistocene biologists. Pages 1-62 in Pleistocene extinctions: the search for a cause (P. S. Martin and H. E. Wright, Jr., editors). Yale University Press, New Haven, Connecticut. Martin, R. A. 1969. Taxonomy of giant Pleistocene beaver Castoroides from Florida. Journal of Paleontology 43:1033-1401. Martin, R. A. 1974. Fossil mammals from the Coleman IIA fauna, Sumpter County. Pages 35-99 in Pleistocene mammals of Florida (S. D. Webb, 56 Timothy S. Young and Joshua Laerm editor). University of Florida Press, Gainesville. Martin, R. A., and S. D. Webb. 1974. Late Pleistocene Mammals from the Devil's Den Fauna, Levy County. Pages 114-148 in Pleistocene mammals of Florida (S. D. Webb, editor). University of Florida Press, Gainesville. McNab, B. K. 1985. Energetics, population biology and distribution of xenarthrans, living and extinct. Pages 219-236 in The evolution and ecology of armadillos, sloths, and vermilinguas (G. G. Montgomery, editor). Smithsonian Institution Press, Washington, D. C. Means, D. B. 1976. Aspects of the significance to terrestrial vertebrates of the Apalachicola River Drainage Basin, Florida. Florida Marine Research Publication 26:37-67. Merriam, J. C, and C. Stock. 1932. The felidae of Rancho La Brea. Carnegie Institute of Washington Publication Number 422. Miller, W. E. 1971. Pleistocene vertebrates of the Los Angeles basin and vicinity (excusive of Rancho La Brea). Bulletin of the Los Angeles County Museum of Natural History 10:1-124. Neill, W. T. 1957. The rapid mineralization of organic remains in Florida and its bearing on supposed Pleistocene records. Quarterly Journal of the Florida Academy of Science 20:1-13. Nelson, R. S., and H. A. Semken. 1970. Paleoecology and stratigraphic significance of the muskrat in Pleistocene deposits. Bulletin of the Geological Society of America 81:3733-3738. Olson, S. L. 1972. A whooping crane from the Pleistocene of North Florida. Condor 74:341. Puri, H. S. 1953. Contributions to the study of the Miocene of the Florida Panhandle. Bulletin of the Florida Geological Survey 38:1- 345. Puri, H. S., and R. O. Vernon. 1964. Summary of the geology of Florida and a guidebook to the classic exposures. Florida Geological Survey Special Publication 5:1-311. Ray, C. E. 1967. Pleistocene mammals from Ladds, Bartow County, Georgia. Bulletin of the Georgia Academy of Science 25:120-150. Ray, C. E., C. S. Denny, and M. Rubin. 1970. A peccary, Platygonus compressus LeConte, from drift of Wisconsin age in northern Pennsylvania. American Journal of Science 268:78-94. Robins, C. R., R. M. Bailey, J. R. Brooker, E. A. Lachner, R. N. Lea, and W. B. Scott. 1991. Common and scientific names of fishes from the United States and Canada. American Fisheries Society Special Publication 20. Rosenau, J. C, G. L. Faulkner, C. W. Hendrey, Jr., and R. W. Hall. 1977. Springs of Florida. Bulletin of the Florida Geological Survey (Revised) 31:1-461. Roth, J. A., and J. Laerm. 1980. A Late Pleistocene vertebrate assemblage from Edisto Island, South Carolina. Brimleyana 3:1-29. Semken, H. A., Jr. 1966. Stratigraphy and Paleontology of the McPherson Equus beds (Sandahl local fauna) McPherson County, Kansas. St. Marks River Fauna 57 Contributions of the Museum of Paleontology, University of Michigan 20:6:121-178. Simpson, G. G. 1945. Notes on Pleistocene and Recent tapirs. Bulletin of the American Museum of Natural History 86:2:37-81. Slaughter, B. H. 1963. Some observations concerning the genus Smilodon, with special reference to Smilodon fatalis. Texas Journal of Science 15:68-81. Spencer, S. M., and F. R. Rupert. 1987. The geology and ground water , of Wakulla County, Florida. United States Department of Agriculture Soil Conservation Service (unpublished advanced copy). Sprunt, A., Jr. 1954. Florida bird life. Coward-McCann Inc. New York, New York. Steadman, D. W. 1980. A review of the osteology and paleontology of turkeys (Aves:Meleagridinae). Contributions to Science, Los Angeles County Museum 330:131-207. Stevens, M. S. 1965. A new species of Urocyon from the Upper Pliocene of Kansas. Journal of Mammalogy 46:265-269. Stock, C. 1925. Cenozoic gravigrade edentates of western North American with special reference to the Pleistocene Megalonychidae and Mylodontidae of Rancho La Brea. Carnegie Institute of Washington Publication Number 331. Stock, C. 1963. Rancho La Brea. Los Angeles County Museum Scientific Series 20:1-83. Uyeno, T., and R. R. Miller. 1962. Late Pleistocene fishes from Trinity River Terrace, Texas. Copeia 25:338-345. Webb, S. D. 1974a. Pleistocene Mammals of Florida. University of Florida Press, Gainesville. Webb, S. D. 1914b. Pleistocene llamas of Florida, with a brief review of Lamini. Pages 170-213 in Pleistocene Mammals of Florida (S. D. Webb, editor). University of Florida Press, Gainesville. Webb, S. D., and K. T. Wilkins. 1984. Historical biogeography of Florida Pleistocene mammals. Pages 370-383 in Contributions in Quaternary Vertebrate Paleontology: a volume in memorial to John E. Guilday (H. H. Geonways and M. R. Dawson, editors.). Carnegie Museum of Natural History, Pittsburgh, Pennsylvania. Winker, C. D., and J. D. Howard. 1977a. Correlation of technically deformed shorelines of the southern Atlantic Coastal Plains. Geology 5:123-127. Winker, C. D., and J. D. Howard. 1971b. Plio-Pleistocene paleogeography of the Florida Gulf Coast interpreted from relict shorelines. Transactions of the Gulfcoast Association of Geologists Society 27:409- 420. Yon, J. W. 1966. Geology of Jefferson County, Florida. Bulletin of the Florida Geological Survey 48:1-119. Accepted 29 July 1992 58 AUTUMN LAND-BIRD MIGRATION ON THE BARRIER ISLANDS OF NORTHEASTERN NORTH CAROLINA by Paul W. Sykes, Jr. For three consecutive years Sykes investigated the autumn migration of land birds in the Bodie Island and Pea Island area of coastal North Carolina. During a 102-day period in 1965, he recorded 110,482 individual birds of 148 species. He was able to correlate major influxes of migratory species with specific weather patterns. His data show seasonal peaks of southward movement for the land-bird species that pass along the North Carolina coast in large numbers. In addition, Sykes recorded five species native to the western United States. Three of these vagrants provided the first reports of Swainson's Hawk, Sage Thrasher, and Western Meadowlark for North Carolina. 1986 49 pages Softbound ISBN 0-917134-12-5 Price: $5 postpaid. North Carolina residents add 6% sales tax. Please make checks payable in U.S. currency to NCDA Museum Extension Fund. Send order to: LAND-BIRD MIGRATION, N.C. State Museum of Natural Sciences, P.O. Box 27647, Raleigh, NC 27611. No Decline in Salamander (Amphibia: Caudata) Populations: A Twenty-Year Study in the Southern Appalachians Nelson G. Hairston, Sr., and R. Haven Wiley Department of Biology, University ofNorth Carolina Chapel Hill, North Carolina 27599-3280 ABSTRACT—Identical observations, conducted 1-4 times per year for 15-20 years at two locations in the southern Appalachians, have yielded quantitative data on populations of six species of salamanders. Although the numbers have fluctuated for various reasons, there has been no trend in the numbers of any of the species. The "world-wide decline of amphibian populations" has not occurred in the two localities studied. Recently, much attention has been given to a decline in many populations of amphibians (Barringer 1990, Blaustein and Wake 1990, Phillips 1990). There is a suggestion by some authors that there is a general cause for a supposed "world-wide" decline. We do not deny that many amphibian species have decreased in abundance. Among the causes that have been suggested are acid precipitation (Harte and Hoffman 1989, Beebee et al. 1990) and ultraviolet increase due to ozone depletion (Barringer 1990, Blaustein and Wake 1990, Phillips 1990). The same authors have considered overcollecting and rejected it as a general cause. Habitat destruction is also widely mentioned. The last cause is common to all species, except for some pioneering ones, and would not apply only to amphibians. The situation is regard-ed by many herpetologists as very serious, so much so that the World Conservation Union (IUCN), Species Survival Commission, has activated a Declining Amphibian Populations Task Force. This group has estab-lished local subgroups throughout the United States and elsewhere in the Americas to promote research on the problem. If there has been a general cause for the decline in amphibian populations, all amphibian populations should be involved; if they are not, the original claim of a "world-wide decline" must be modified, either by eliminating some taxonomic groups, some ecolog
Object Description
Description
Title | Brimleyana |
Contributor |
North Carolina State Museum of Natural Sciences. |
Date | 1993 |
Subjects |
Zoology--Southern States--Periodicals Ecology--Southern States--Periodicals Natural history--Southern States--Periodicals |
Place | North Carolina, United States |
Time Period | (1990-current) Contemporary |
Description | Vol. 18;"The journal of the North Carolina State Museum of Natural History." |
Publisher | [Raleigh, NC : North Carolina State Museum of Natural History] |
Agency-Current |
North Carolina Department of Environmental Quality |
Rights | State Document see http://digital.ncdcr.gov/u?/p249901coll22,63754 |
Physical Characteristics | v. :ill. ;23 cm. |
Collection | North Carolina State Documents Collection. State Library of North Carolina |
Type | text |
Language | English |
Format | Periodicals |
Digital Characteristics-A | 8390 KB; 150 p. |
Digital Collection | North Carolina Digital State Documents Collection |
Digital Format | application/pdf |
Audience | All |
Pres File Name-M | pubs_serial_brimleyana1993v18.pdf |
Pres Local File Path-M | Preservation_content\StatePubs\pubs_serial_brimleyana\images_master |
Full Text | fill ^&y*t*\*ol\o^\*:^^\ auS^ s^eW-N. C. DOCUMENTS CLEARINGHOUSE AUG 10 1993 N.C. STATE LBRARY RALOGH number 18 June 1993 EDITORIAL STAFF Richard A. Lancia, Editor Suzanne A. Fischer, Assistant Editor Eloise F. Potter, Production Manager EDITORIAL BOARD James W. Hardin Rowland M. Shelley Professor ofBotany Curator ofInvertebrates North Carolina State University North Carolina State Museum ofNatural Sciences William M. Palmer Robert G. Wolk Director ofResearch and Collections Director ofPrograms North Carolina State Museum North Carolina State Museum ofNatural Sciences ofNatural Sciences Brimleyana, the Zoological Journal of the North Carolina State Museum of Natural Sciences, appears twice yearly in consecutively numbered issues. Subject matter focuses on systematics, evolution, zoogeography, ecology, behavior, and paleozoology in the southeastern United States. Papers stress the results of original empirical field studies, but synthesizing reviews and papers of significant historical interest to southeastern zoology are also included. Brief communications are accepted. All manuscripts are peer reviewed by specialists in the Southeast and elsewhere; final acceptability is determined by the Editor. Address manuscripts and related correspondence to Editor, Brimleyana, North Carolina State Museum of Natural Sciences, P.O. Box 27647, Raleigh, NC 27611. Information for contributors appears in the inside back cover. Address correspondence pertaining to subscriptions, back issues, and exchanges to Brimleyana Secretary, North Carolina State Museum of Natural Sciences, P.O. Box 27647, Raleigh, NC 27611. In citations please use the full name - Brimleyana. North Carolina State Museum of Natural Sciences Betsy Bennett, Director North Carolina Department of Agriculture James A. Graham, Commissioner CODN BRIMD 7 ISSN 0193-4406 The Myriapod Types of Oscar Harger (Arthropoda: Diplopoda, Chilopoda) Rowland M. Shelley North Carolina State Museum ofNatural Sciences, P.O. Box 27647, Raleigh, North Carolina 27611 ABSTRACT—The type specimens of all five milliped species — Trichopetalum lunatum, T. glomeratum, T. iuloides, lulus furcifer, and Polydesmus armatus—and one of the two centipedes, Lithobius pinetorum, authored by Oscar Harger in his only paper on myri-apods and previously thought to be lost, are housed at the Peabody Museum of Natural History, Yale University, New Haven, Connecti-cut. From our knowledge of the itinerary of the Yale paleontologi-cal expedition of 1871, we know the type locality of T. glomeratum, I. furcifer, P. armatus, and L. pinetorum, previously stated as the "John Day River Valley, Oregon" is restricted to the vicinity of Canyon City, Grant County, on the western slope of the Blue Mount-ains. The female holotype confirms that T. glomeratum is a repre-sentative of the chordeumatoid family Conotylidae, and the name is assigned provisionally to Taiyutyla pending collection of a male topotype. Unidentifiable female conotylids are also reported from another area in eastern Oregon and the Snake Mountains in eastern Nevada, which suggests that the family is widespread in montane forests at high elevations in the generally arid Columbia Plateau and Basin and Range Physiographic Provinces. To facilitate future studies, I provide gonopod drawings of male syntypes for /. furcifer and P. armatus. One of the more obscure authors of North American myriapods is Oscar Harger (1843-87), whose sole publication on these arthropods (Harger 1872) described the milliped genus Trichopetalum and seven species, two centipedes {Lithobius pinetorum and Geophilus gracilis) and five millipeds {Trichopetalum lunatum, T. glomeratum, T. iuloides, lulus furcifer, and Polydesmus armatus). Born at Oxford, Connecticut, Harger attended the Connecticut Literary Institute at Suffield and Yale College, graduating from the latter with honors in 1868 (Schuchert and LeVene 1940). After briefly studying zoology under Professor A. E. Verrill, Harger became the first assistant to the vertebrate paleontologist, O. C. Marsh, partici-pating on the latter's expeditions into the American West in 1871 and 1873. From July to September 1872, Harger dredged marine organisms on a Coast Survey steamer with Professors Verrill and Sydney I. Smith, Yale's first professor of Comparative Anatomy, who earlier had been naturalist to the U.S. Lake Survey and collected the types Brimleyana 18:1-13, June 1993 1 2 Rowland M. Shelley of T. iuloides. Harger was studious and an active reader, and Marsh valued his scientific opinions in paleontology. However, Marsh would not allow Harger to publish on vertebrate fossils, either alone or jointly with him, so Harger 's only papers are on invertebrates—that of 1872 on myriapods, two on isopods, and one on a fossil spider. From 1870 to 1873, Marsh led four vertebrate paleontological expeditions of Yale students and recent graduates into the West (Schuchert and LeVene 1940). The idea of such efforts arose from preliminary explorations he made on a trip to the end of the trans-continental railroad in Wyoming in August 1868 after attending a scientific meeting in Chicago. The 1871 expedition traveled to regions of Kansas, Wyoming, and Utah, where Harger collected 10 fossil species. The group then rested a few days in Salt Lake City with Brigham Young while Marsh prepared to explore a new area, the John Day River Basin in central Oregon. After traveling 12 days by rail and stage, the party crossed the Blue Mountains and arrived at Canyon City, Oregon, on the John Day River on 17 October 1871, where it waited several days for a military escort from Fort Harney, 75 mi (120 km) to the south. The group collected fossils from 31 October to 8 November in the John Day region before traveling down the Columbia River to Portland; it then traveled to San Francisco and returned east directly by rail or by boat via Panama. While the expedition was in the John Day River area, Harger, or Harger and Professor G. H. Collier, collected four Oregon myriapods that he described in 1872 — L. pinetorum, T. glomeratum, I. furcifer, and P. armatus. Both the publication and labels in the vials give the locality as just the "John Day River Valley," but knowledge of the group's activities enabled me to infer a more precise site. The John Day River arises on the western slope of the Blue Mountains in Grant and Umatilla counties, flows westward into Wheeler County, then heads northward to the Columbia River forming the boundaries between Wheeler/Wasco and Sherman/Gilliam counties. It is not to be confused with Days Creek, Douglas County, in the Umpqua River drainage of southwestern Oregon, the probable type locality for Zantona douglasia Chamberlin and Bollmanella oregona Chamberlin (Shear 1974, Gardner and Shelley 1989), which Chamberlin (1941a) misnamed as "John Day Creek." Because most millipeds require moist leaf litter and much of the John Day Basin is in the arid rain shadow of the Cascade Mountains, I (Shelley 1990) speculated that the site was probably near the confluence of the John Day and Columbia rivers in either Sherman or Gilliam county. However, as all the myriapods were collected in October 1871, and the expedition reached Canyon City on 17 October and only collected fossils from 31 October to 8 November Myriapod Types of Oscar Harger 3 after waiting for the military escort, it is evident that during most of the part of October that the group was in the John Day Valley, it was resting in Canyon City. Consequently, there was ample time for relaxed explorations in the vicinity of Canyon City, and I, therefore, believe that Harger's myriapods were collected near this town. Because Harger's paper specifies that L. pinetorum, I. furcifer, and P. armatus were collected by Professor Collier and himself, and T. glomeratum was taken by Harger alone, collecting probably occurred on at least two different dates, as one day Harger went out alone and the other he was accompanied by Collier. There could be as few as one site and as many as four, but further specification is not possible with what we know now. Consequently, the type locality for all of Harger's Oregon species is restricted to the vicinity of Canyon City, Grant County, on the western slope of the Blue Mountains. Harger's centipedes have received little attention since their description. They were included in the catalog of North American myriapods by Bollman (1893), who noted that G. gracilis Harger, 1872, was preoccupied by G. gracilis Meinert, 1870, proposed for a European geophilomorph. Cook and Collins (1891) remarked that Harger's description of G. gracilis conformed very closely to Schendyla nemorensis (C. L. Koch, 1837), and the former is now regarded as a junior synonym (Crabill 1953, 1961). Stuxberg (1875) included L. pinetorum in his list of North American lithobiids, but he had no personal knowledge of the species. Kevan (1983a) listed both species as potential inhabitants of Canada, recognizing the synonymy of G. gracilis under S. nemorensis. In contrast to the centipeds, Harger's millipeds have been cited in a number of publications, but the type specimens were thought to be lost. Chamberlin and Hoffman (1958) stated that their "present location [was] unknown" or that they were "not known to exist," and similarly, Shear (1971, 1972) said that the holotypes of T. glomeratum and T. iuloides were lost and that the whereabouts of that of T. lunatum was unknown. Causey (1967) guessed right when she stated that the holotype of T. lunatum was at the "Peabody Museum of Natural History, Yale University, if extant," but evidently she made no inquiries to confirm this supposition. While recently visiting the Peabody's Museum's collection, I unexpectedly discovered these types in the myriapod cabinet, where they have languished in obscurity for 120 years. A few vials were still capped with wax and had not been touched for decades. The types of P. armatus were in the general collection and not labeled as such, but those of the other millipeds were clearly marked as types and grouped in a clamp-top jar. A concerted search failed to reveal the types of G. gracilis, which ap- 4 Rowland M. Shelley parently are lost, but those of L. pinetorum were in an individual vial and clearly labeled. The sample consists of 12 nearly legless syntypes, seven males and five females, and is number 2175; according to the label it was collected by Harger alone, whereas the published account states that it was collected by him and Professor Collier. All the millipeds are listed in the continental checklist (Chamberlin and Hoffman 1958), and detailed accounts of those Harger assigned to Trichopetalum have recently appeared (Palmen 1952; Shear 1971, 1982; Shelley 1988, In Press). In the following accounts I update these reports by providing information on the type specimens, a brief historical review of each species, and pertinent anatomical observations. Complete synonymies are presented, and each species is placed in its proper order and family. Chordeumatida: Trichopetalidae Trichopetalum lunatum Harger Trichopetalum lunatum Harger, 1872:3, pi. II, figs. 1-4. Ryder, 1881:527. Packard, 1883:192. McNeill, 1888:8. Cook and Collins, 1895:63-64, pi. Ill, figs. 52-54. Williams and Hefner, 1928:115, fig. 12d. Causey, 1951:119, figs. 6-8; 1967:80, fig. 1. Palmen, 1952:8-11, figs. 10-17. Chamberlin and Hoffman, 1958:102-103. Shear, 1972:277, figs. 497-499. Kevan, 19836:2967. Shelley, 1988:1650. Trichopetalum album Cook and Collins, 1895:64-66, pis. II-III, figs. 22-29, 36-45. Chamberlin and Hoffman, 1958:102. Type Specimens—Five male and nine female syntypes (nos. 2208-2209) collected by O. Harger in May 1872 at New Haven, New Haven County, Connecticut; one male and one female syntype (no. 2125) taken by an unknown collector on an unknown date at Mt. Carmel, ca. 7 mi (11.2 km) north of New Haven, New Haven County. Remarks—Harger assigned three new species to his genus Trichopetalum but did not specify the type species, so Cook and Collins (1895) subsequently designated T lunatum. It is the only one of Harger's five milliped species to retain its original combination. The identity of T lunatum has been well established by Cook and Collins (1895), Palmen (1952), Causey (1967), and Shear (1972); a male syntype from New Haven that I dissected conformed to these Myriapod Types of Oscar Harger 5 accounts. For details of the genitalia, refer to the illustrations in Palmen (1952) and Shear (1972). Chordeumatida: Conotylidae Taiyutyla glomerata (Harger), new combination Trichopetalum glomeratum Harger, 1872:118, pi. II, fig. 5. Ryder, 1881:527 Packard, 1883:192. McNeill, 1888:8. Chamberlin and Hoffman, 1958:105. Shear, 1971:63. Craspedosoma glomeratum: Bollman, 1893:120. Conotyla glomerata: Cook and Collins, 1895:78. Cook, 1904:69. Type Specimen—Female holotype (No. 2173) collected by O. Harger in October 1871 from the vicinity of Canyon City, in the John Day River Valley, Grant County, Oregon. Remarks—The holotype is somewhat deformed, and its genitalia have been dissected and are lost. Cook and Collins (1895) stated that the original description was too brief to allow accurate generic placement but that the segment number, short fifth antennomere, and triangular eye patch resembled the condition in Conotyla. Shear (1971) agreed that accurate generic placement was impossible but perceived a similarity to Taiyutyla; he did not think the name could be referred to either Trichopetalum or Conotyla and considered it a nomen dubium. The holotype is about 8 mm long and has 30 post cephalic segments with obvious lateral tergal knobs that give rise to two prominent setae, so it is clearly a conotylid. Generic placement is impossible to determine with certainty until a male topotype is obtained, but the milliped is smaller and its lateral setae are much longer than those of comparative specimens of Conotyla atrolineata (Bollman), the western-most known representative of this genus, occurring in central British Columbia, northeastern Washington, and northern Idaho, over 200 mi (320 km) north northeast of Canyon City. These considerations tend to exclude Conotyla, but the type locality is also well removed from most of the known distribu-tions of the other northwestern conotylid genera Bollmanella and Taiyutyla, which are from southern coastal Oregon to Mason County, Washington, and in the Coast Ranges from San Francisco Bay to the Columbia River, respectively (Shear 1974, 1986). However, one species in each of these genera occurs east of the above ranges, B. bifurcata Shear, in the Wallowa Mountains, Wallowa County, Oregon, and T curvata Loomis and Schmitt, in Lincoln County, Montana, so either genus could occur in the Blue Mountains, which occupy an intermediate 6 Rowland M. Shelley geographical position between the Coast Range and both the Wallowa Mountains and Montana. Furthermore, Canyon City is only about 110 mi (176 km) southwest of the type locality of B. bifurcata. Therefore, I borrowed the types of both B. bifurcata and T. curvata for direct comparisons with that of glomerata. Few setae remain on the types of B. bifurcata, and those that do exist, on the caudal end of the male holotype, seem shorter and are not nearly as prominent as are those on glomerata. However, the setae on glomerata agree closely in length and prominence with those on the holotype of T. curvata. There is reasonable agreement in body dimensions between glomerata and both other conotylids, but because of the similarity in the setae, I provisionally assign glomerata to Taiyutyla, pending collection of a male topotype. This change, which formalizes Shear's (1971) perception of similarity to Taiyutyla, also necessitates the feminine suffix of the specific name. Fieldwork is needed in the Blue Mountains to collect a male conotylid to determine the identity and generic position of glomerata and to confirm or disprove this decision. Present evidence shows that the Conotylidae is much more wide-spread in the West than currently known. There is a female in the Florida State Collection of Arthropods from 12.5 mi (20 km) south of Baker City, Baker County, Oregon, that might be conspecific with glomerata, although this site is east of the Blue Mountains and presumably is drier than Canyon City. I also recently received two female conotylids that are superficially very similar to glomerata from the Snake Mountains, White Pine County, Nevada, in the eastern part of that state and hundreds of kilometers from any known site for the family. These two records plus glomerata suggest that conotylids could be scattered across the arid Columbia Plateau and Basin and Range Physiographic Provinces, where they are undoubtedly restricted to cool-er, forested regions at high elevations. The Ruby Mountains near Elko, Nevada, is another plausible area for conotylids, as are ranges in the central part of that state. Because only a few millipeds of any family have ever been collected from the "inselberg" mountains of these provinces, a concerted field effort is needed to both clarify the systema-tic positions of these conotylids and document the total diplopod fauna. Chordeumatida: Caseyidae Underwoodia iuloides (Harger) Trichopetalum iuliodes Harger, 1872:118. pi II, fig. 6. Trichopetalum juloides: Ryder, 1881:527. Trichopetalum iulioides: Packard, 1883:192. Trichopetalum iuloides: McNeill, 1888:8. Chordeuma iuloides: Bollman, 1893:121. Myriapod Types of Oscar Harger 7 Underwoodia polygama Cook and Collins, 1895:80-82, pi. X, figs. 180-190. Paleman, 1952:2-8, figs. l-9a. Chamberlin and Hoffman, 1958:107. Kevan, 19836:2968. Underwoodia iuloides: Cook and Collins, 1895:83-84, pi. X, figs. 177-178. Chamberlin and Hoffman, 1958:107. Kevan, 19836:2968. Shelley, 1988:1648-1649; In Press: Type Specimens—Eight female syntypes (No. 2207) collected by S. I. Smith in 1871 at Simon's Harbor (misspelled as Simmon's) on the north shore of Lake Superior, Ontario, Canada. This site is now in Pukaskwa National Park. Remarks—A review of Underwoodia with a description, discussion, and illustrations of U. iuloides is in press. For details on this species, see Shelley (1988). Fig. 1-3. Bollmaniulus furcifer, male syntype. 1, anterior gonopods, anterior view. 2, the same, posterior view. 3, posterior gonopods, anterior view. Scale line = 2.2 mm for figs. 1-2, 1.6 mm for fig. 3. Julida: Parajulidae Bollmaniulus furcifer (Harger) Figs. 1-3 lulus furcifer Harger, 1872:119, pi. II, fig. 7. Parajulus furcifer: Bollman, 1887:44. Cook, 1904:70-71, pi. V, figs. 5a-e. Chamberlin, 1920:35. 8 Rowland M. Shelley Paraiulus furcifer. Brolemann, 1895:69, pi. 7, figs. 21-23. Bollmaniulus furcifer: Verhoeff, 1926:65. Chamberlin and Hoffman, 1958:133. Buckett, 1964:18. Kevin, 19836:2964. Taijulus furcifer. Chamberlin, 1938:205. Caliulus furcifer. Chamberlin, 1940:15; 1944:80. Type Specimens—Three male and 13 female syntypes (No. 2172), most highly fragmented, collected by O. Harger and G. H. Collier in October 1871 from the vicinity of Canyon City, in the John Day River Valley, Grant County, Oregon. Remarks—Bollman (1887) transferred this species into Parajulus, misspelled as Paraiulus by Brolemann (1895), and Cook (1904) recorded it from Corvallis, Oregon. Chamberlin (1920) reported it from Clare-mont, Los Angeles County, California, surely a misidentification of another, possible congeneric parajulid. Verhoeff (1926) listed furcifer as the only component of his new genus Bollmaniulus, thereby mak-ing it the type species by monotypy as reported by Jeekel (1971). He did not specifically designate furcifer as the generotype, so this status does not result from original designation, as stated by Chamber-lin and Hoffman (1958). Chamberlin (1938, 1940) evidently was unaware of Verhoeffs action when he transferred furcifer into his new genera Taijulus and Caliulus, respectively, both of which have subsequently been placed in synonymy under Bollmaniulus (Chamberlin and Hoff-man 1958, Hoffman 1979). Chamberlin (1944) repeated the combination C. furcifer for a form from McCloud, Siskyou County, California, and added that the species was common over much of Oregon and California. Buckett (1964) recognized the combination Bollmaniulus furcifer and stated that it ranged from British Columbia into California. As noted by Hoffman (1979, 1992), the Parajulidae is one of the two most dominant Nearctic diplopod families in terms of com-ponent genera and species, the other being the Xystodesmidae (Polydesmida). It was studied from 1948 to about 1974 by Dr. Nell B. Causey, who amassed a large collection and examined most type specimens while conducting a detailed family revision. Unfortunately, she never completed the project and published only a few brief papers before her death in 1979. Consequently, knowledge of the Parajulidae is not nearly as advanced as those of the other major Nearctic diplopod families. Work on the taxon must essentially begin anew, a daunting task because of the diversity of the family and the enormous amount of preserved material in nearly every major and minor milliped repository on the continent. The types of /. furcifer will be crucial to an investigation of Pacific parajulids, because as the eighth oldest generic name in the family, Bollmaniulus has priority over such other nominal Pacific Myriapod Types of Oscar Harger Figs. 4-5 Chonaphe armata, male syntype. 4, telopodite of left gonopod, medial view. 5, the same, lateral view. Scale line = 1.14 mm for fig. 4, 1.0 mm for fig. 5. genera as Saiulus, Spathiulus, Sophiulus, Codiulus, and Simiulus, all authored by Chamberlin (1940), Tuniulus (Chamberlin 19415), and Mulaikiulus (Chamberlin 1941a), so additional generic synonymies could result from a study of these western forms. For the benefit of future students, I have included drawings of the gonopods of a male syntype (Figs. 1-3). Polydesmida: Xystodesmidae Chonaphe armata (Harger) Fig. 4-5 Polydesmus armatus Harger, 1872:119-120, pi. II, fig. 8. Leptodesmus armatus: Bollman, 1893:122. Chamberlin, 1911:264. Chonaphe armata: Cook, 1904:56-57, pi. Ill, figs. 2a-c. Attems, 1931:65- 67, figs. 100-101; 1938:156, fig. 177. Chamberlin, 1949:125. Chamberlin and Hoffman, 1958:27. Kevan, 19835:2968. Shelley, 1990:2314. Type Specimens—One male and two female syntypes, all highly fragmented, collected by O. Harger and G. H. Collier in October 10 Rowland M. Shelley 1871 from the vicinity of Canyon City, in the John Day River Valley, Grant County Oregon. This sample was discovered in the general milliped collection and is unnumbered. Remarks—Harger's single gonopod illustration enabled Cook (1904) to recognize that a male sent to him from an unknown locality in Washington was referrable to armatus. Bollman (1893) had earlier transferred armatus to Leptodesmus, a combination repeated by Chamberlin (1911), but Cook (1904) assigned it to his new genus, Chonaphe, a combination that subsequently has been recognized by Attems (1931, 1938), Chamberlin (1949), Chamberlin and Hoffman (1958), and Shelley (1990). Cook (1904) provided three additional genitalia drawings, and I include here medial and lateral views of the gonopod of a male syntype (Figs. 4—5). Five nominal species comprise Chonaphe, but Hoffman (1979) thought these might be subspecies. I (Shelley 1990) found few significant differences between these forms and concluded that the genus might be monotypic with C. armata being the oldest name. I am preparing a generic revision. ACKNOWLEDGMENTS—\ thank C. L. Remington and R. J. Pupedis for providing access to the Peabody Museum holdings and subsequently loaning Harger's types. The holotype of Taiyutyla curvata, housed at the National Museum of Natural History, Smithsonian Institution, Washington, D.C., was loaned by J. A. Coddington; the types of Bollmanella bifurcata, housed at the Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, were loaned by H. W. Levi. The conotylid from Baker County, Oregon, was discovered in material loaned by G. B. Edwards, Florida State Collection of Arthropods, Gainesville. Cathy Wood typed and retyped numerous drafts of the manuscript, and figures 1-5 were prepared by R. G. Kuhler. LITERATURE CITED Attems, C. G. 1931. Die familie Leptodesmidae und andere Poly-desmiden. Zoologica, Stuttgart, 30, Lief 3-4:1-149. Attems, C. G. 1938. Polydesmoidea II. Fam. Leptodesmidae, Platyrhacidae, Oxydesmidae, Gomphodesmidae. Das Tierreich, Lief 70:1-576. Bollman, C. H. 1887. Notes on North American Julidae. Annals of the New York Academy of Science 4:25-44. Bollman, C. H. 1893. The Myriapoda of North America. Bulletin No. 46, United States National Museum. Brolemann, H. W. 1895. Liste de myriapodes des Etats-Unis, et principale-ment de la Caroline du Nord, faisant partie des collections de M. Eugene Simon. Annales Societe Entomologique de France 65:43-70. Myriapod Types of Oscar Harger 11 Buckett, J. S. 1964. Annotated list of the Diplopoda of California. Simmons Publishing Company, Davis, California. Causey, N. B. 1951. New genera and species of chordeumoid millipeds in the United States, and notes on some established species. Proceedings of the Biological Society of Washington 64:117-124. Causey, N. B. 1967. Trichopetalum subterranewn, new species from Kentucky, new records and a key to the genus. Proceedings of the Biological Society of Washington 80:117-122. Chamberlin, R. V. 1911. Notes on myriopods from Alaska and Wash-ington. Canadian Entomologist 43:260-264. Chamberlin, R. V. 1920. Centipedes and millipedes from near Claremont. Pomona College Journal of Entomology and Zoology 12:35. Chamberlin, R. V. 1938. New diplopods. Proceedings of the Biological Society of Washington 51:205-208. Chamberlin, R. V. 1940. New genera and species of North American Paraiulidae. Bulletin of the University of Utah 30 [Biological Series, 5(7)]: 1-39. Chamberlin, R. V. 1941a. New western millipeds. Bulletin of the University of Utah 31[Biological Series, 6(5)]:l-23. Chamberlin, R. V. 1941b. New American millipeds. Bulletin of the University of Utah 31 [Biological Series, 6(4)]: 1-39. Chamberlin, R. V. 1944. Some records of myriopods collected by W. M. Pearce in California. Pan-Pacific Entomologist 20:79-80. Chamberlin, R. V. 1949. Some western millipeds of the family Chelodesmidae. Proceedings of the Biological Society of Washington 62:125-132. Chamberlin, R. V., and R. L. Hoffman. 1958. Checklist of the millipeds of North America. Bulletin No. 212, United States National Museum. Cook, O. F. 1904. Myriapoda of northwestern North America. Harriman Alaska Expedition 8(Insects, part l):47-82. Cook, O. F., and G. N. Collins. 1891. Notes on North American Myriapoda of the family Geophilidae, with descriptions of three genera. Proceedings of the United States National Museum 13:383-396. Cook, O. F., and G. N. Collins. 1895. The Craspedosomatidae of North America. Annals of the New York Academy of Science 9:1-100. Crabill, R. E. 1953. The Schendylidae of northeastern North America (Chilopoda: Geophilomorpha: Schendylidae). Journal of the New York Entomological Society 61:93-98. Crabill, R. E. 1961. A catalogue of the Schendylinae of North America including Mexico, with a generic key and proposal of a new Simoporus (Chilopoda: Geophilomorpha: Schendylinae). Entomological News 72:67-80. Gardner, M. R., and R. M. Shelley. 1989. New records, species, and genera of caseyid millipeds from the Pacific Coast of North America. Pan-Pacific Entomologist 65:177-268. 12 Rowland M. Shelley Harger, O. 1872. New North American myriapods. American Journal of Science and Arts 4:116-121. Hoffman, R. L. 1979. Classification of the Diplopoda. Museum d'Histoire Naturelle, Geneva, Switzerland. Hoffman, R. L. 1992. On the taxonomy of the milliped genera Pseudojulus Bollman, 1887, and Georgiulus, gen. nov., of southeastern United States (Julida: Parajulidae). Jeffersoniana 1:1-19. Jeekel, C. A. W. 1971. Nomenclator generum et familiarum Diplopodorum: A list of the genus and family-group names in the class Diplopoda from the 10th edition of Linnaeus, 1758, to the end of 1957. Mono-grafieen van de Nederlandse Entomologische Vereniging, Number 5. Kevan, D. K. McE. 1983a. A preliminary survey of known and potentially Canadian and Alaskan centipedes (Chilopoda). Canadian Journal of Zoo-logy 61:2938-2955. Kevan, D. K. McE. 1983b. A preliminary survey of known and potentially Canadian millipedes (Diplopoda). Canadian Journal of Zoology 61:2956- 2975. McNeill, J. 1888. A list, with brief descriptions, of all the species, including one new to science, of Myriapoda of Franklin County, Indiana. Bulletin of the Brookville Society of Natural History, Number 3. Meinert, F. 1870. Myriopoda Musaei Hauniensis I. Geophili. Naturhistorisk Tidsskrift 3:241-268. Packard, A. S. 1883. A revision of the Lysiopetalidae, a family of Chilognath Myriopoda, with a notice of the genus Cambala. Proceedings of the American Philosophical Society 21:177-209. Palmen, E. 1952. Survey of the Diplopoda of Newfoundland. Annales Zoologici Societatis Zoologicae Botanicae Fennicae 'Vanamo' 15:1-31. Ryder, J. A. 1881. List of the North American species of myriapods belonging to the family of the Lysiopetalidae, with a description of a blind form from Luray Cave, Virginia. Proceedings of the United States National Museum 3:524-529. Schuchert, C, and C. M. Levene. 1940. O. C. Marsh, Pioneer in Paleontology. Yale University Press, New Haven, Connecticut. Shear, W. A. 1971. The milliped family Conotylidae in North America, with a description of the new family Adritylidae (Diplopoda: Chordeumida). Bulletin of the Museum of Comparative Zoology 141:55-98. Shear, W. A. 1972. Studies in the milliped order Chordeumida (Diplopoda): A revision of the family Cleidogonidae and a reclassification of the order Chordeumida in the New World. Bulletin of the Museum of Comparative Zoology 144:151-352. Shear, W. A. 1974. The milliped genus Bollmanella (Diplopoda, Chordeumida, Conotylidae). Psyche 81:134-146. Shear, W. A. 1976. The milliped family Conotylidae (Diplopoda: Chordeumida). Revision of the genus Taiyutyla with notes on recently proposed taxa. American Museum Novitates Number 2600. Myriapod Types of Oscar Harger 13 Shelley, R. M. 1988. The millipeds of eastern Canada (Arthropoda: Diplopoda). Canadian Journal of Zoology 66:1638-1663. Shelley, R. M. 1990. A new milliped of the genus Metaxycheir from the Pacific coast of Canada (Polydesmida: Xystodesmidae), with remarks on the tribe Chonaphini and the western Canadian and Alaskan diplopod fauna. Canadian Journal of Zoology 68:2310-2322. Shelley, R. M. The milliped genus Underwoodia (Chordeumatida: Caseyidae). Canadian Journal of Zoology, In Press. Stuxberg, A. 1875. Lithobioidae Americae Borealis. Ofversigt af Kongl. Vetenskaps-Akademiens Forhandlingar 22:23-32. Verhoeff, K. W. 1926. Chilognathen-Beitrage. Zoologisher Anzeiger 68:57- 71. Williams, S. R., and R. A. Hefner. 1928. The millipedes and centipedes of Ohio. Bulletin Number 18, Ohio Biological Survey, 4(3) [Ohio State University Bulletin, 33(7)]:93-146 Accepted 9 September 1992 14 THE SEASIDE SPARROW, ITS BIOLOGY AND MANAGEMENT Edited by Thomas L. Quay, John B. Funderburg, Jr., David S. Lee, Eloise F. Potter, and Chandler S. Robbins The proceedings of a symposium held at Raleigh, North Carolina, in October 1981, this book presents the keynote address of F. Eugene Hester, Deputy Director of the U.S. Fish and Wildlife Service, a bibliography of publications on the Seaside Sparrow, and 16 major papers on the species. Authors include Arthur W. Cooper, Oliver L. Austin, Jr., Herbert W. Kale, II, William Post, Harold W. Werner, Glen E. Woolfenden, Mary Victoria McDonald, Jon S. Greenlaw, Michael F. Delany, James A. Mosher, Thomas L. Merriam, James A. Kushlan, Oron L. Bass, Jr., Dale L. Taylor, Thomas A. Webber, and George F. Gee. A full-color frontispiece by John Henry Dick illustrates the nine races of the Seaside Sparrow, and a recording prepared by J. W. Hardy supplements two papers on vocalizations. "The Seaside Sparrow, with its extensive but exceedingly narrow breeding range in the coastal salt marshes, is a fascinating species. All the authors emphasize that the salt marsh habitat is at peril. . . . The collection is well worth reading." — George A. Hall, Wilson Bulletin. 1983 174 pages Softbound ISBN 0-917134-05-2 Price: $15 postpaid. North Carolina residents add 6% sales tax. Please make checks payable in U.S. currency to NCDA Museum Extension Fund. Send order to: SEASIDE SPARROW, N.C. State Museum of Natural Sciences, P.O. Box 27647, Raleigh, NC 27611. A Late Pleistocene Vertebrate Assemblage from the St. Marks River, Wakulla County, Florida Timothy S. Young1 and Joshua Laerm The University of Georgia Museum ofNatural History and Department of Zoology, University of Georgia, Athens, Georgia 30602 ABSTRACT—The St. Marks River in the central panhandle of Florida contains a well known, apparently late Pleistocene vertebrate as-semblage that has been only superficially examined and reported. Previous collections are reviewed, and we report on new fossil materials recently obtained. Included are 37 species of mammals, 26 birds, 13 reptiles, 2 amphibians, and 9 fish. Of these, 14 species of mammals and 2 reptiles are limited solely to the Pleis-tocene. The fauna is mixed and reflects heterochronous deposition over time beginning at least in the late Pleistocene (Wisconsinan) and extending through the Recent. The species present reflect mixed woodland and grassland terrestrial communities as well as mixed estuarine and freshwater aquatic communities. The St. Marks River assemblage compares well to other contemporaneous late Pleisto-cene Florida panhandle sites. One extralimital taxa is reported, Pylodictic cf. P. olivaris, the flathead catfish, whose natural range has not been reported east of the Mobile Bay drainage basin. Florida is characterized by a number of rich and well documented Pleistocene vertebrate assemblages (Webb 1974a, Lundelius et al. 1983, Webb and Wilkins 1984) that contain a mixture of extant and extinct South American immigrant and North American endemic species. The majority of these sites are distributed throughout the peninsular portion of the State (Webb 1974a, Webb and Wilkins 1984). However, with the exceptions of Wakulla Springs (Brodkorb 1963, Webb 1974a), Chipola River (Martin 1969, Webb 1974a), and Aucilla River (Olson 1972, Webb 1914a, Gillette 1976a) very little attention has been devoted to sites on the panhandle of Florida. The St. Marks River, located 32 km south of Tallahassee in Wakulla County, is a particularly rich, late Pleistocene panhandle site that has only been superficially investigated (Gillette 19766, Steadman 1980). Leidy (1870), who reported on the occurrence of Mammuthus columbi (now M. jeffersonii), provided the first record of vertebrate fossil remains recovered from the St. Marks River, though the exact locality was not given. Subsequently, the St. Marks River has attracted 'Present address: Department of Zoology, University of Florida, Gainesville, FL 32611. Brimleyana 18:15-57, June 1993 15 16 Timothy S. Young and Joshua Laerm numerous amateur collectors but a limited number of professionals. Storrs Olson (National Museum of Natural History) collected from a broad, shallow water area in Wakulla County near the Leon County line several times between 1968 and 1970. Tall Timbers Research Station sponsored collecting parties in the same area during 1974. Published accounts of the fauna are those of Gillette (1976b), who reported the mammals, and Steadman (1980), who discussed two specimens of Meleagris gallopavo. Storrs Olson (personal communication) examined the avian assemblage from the previous collections; however, he did not publish his findings. With the exception of Mammuthus sp. Mammut americanum, Synaptomys australis, and possibly Equus, the majority of species from Gillette's (1916b) report are extant. Gillette (19766) suggested that the assemblage was important because it represented a restricted temporal interval of the latest Pleistocene through the Holocene. Olsen (personal communication) felt the avian assemblage was very similar to that of today. Steadman (1980) characterized the site as a late Pleistocene deposit. The St. Marks River has also been reported by Lundelius et al. (1983) as being a naturally accumulating, fluvial Rancholabrean deposit. The purpose of our study is to review previous collections and to report on new fossil materials recently obtained from the St. Marks River. We provide information regarding the paleoenvironment of the depositional area and compare the St. Marks River fauna to other late Pleistocene faunas in the region. GEOLOGICAL AND GEOGRAPHIC SETTING Florida consists of five naturally occurring topographical divisions (Cooke 1939:14). The St. Marks River drainage basin is in the coastal lowlands division. Although the panhandle of Florida shows a topographical record of the relict shorelines, no ages have been securely assigned to these formations (Winker and Howard 1977a, 6). The coast line of the panhandle during the late Pleistocene is reported to be similar to that of today (Winker and Howard 19776). The St. Marks River is considered part of the Gulf Hammock region; it is underlain by the Upper Oligocene Suwannee Limestone (Harper 1914:302). The early Miocene St. Marks Formation overlies the Suwannee Formation in almost all of Wakulla County (Puri and Vernon 1964). The St. Marks Formation was revised to include the calcareous downdip facies of the Tampa Formation (Puri 1953). These formations can be found in many areas as outcroppings in springs and rivers (Spencer and Rupert 1987). The surface is mostly loamy St. Marks River Fauna 17 sand, probably Pleistocene in origin. The soil surrounding the river's edge is classified as Tooles-Nutall fine sand that is frequently flooded (Spencer and Rupert 1987). Topographically the region is nearly level, except for a few hilly areas (Harper 1914:302). The whole area east of the Apalachicola River in Wakulla County is called the Woodville Karst Plain (Hendry and Sproul 1966, Yon 1966), characterized by sand dunes overlying limestone (Hendry and Sproul 1966:154). The St. Marks River is fed by the St. Marks Spring located just inside Leon County (Fig. 1). Limestone lines the perimeter of the spring. The vent is located about 26 m below the water surface and has an average base flow of 14.7 m3/sec (Rosenau et al. 1977). This measurement was taken approximately 800 m down stream Leon County ) Basin ' Wakulla County • 3 •f f St. Marks River Wakulla River^— - /• // * 1 ( l_l \ I 1 mile N t Gulf of Mexico Fig. 1. St. Marks River in the central panhandle of Florida. Solid circles indicate 1987 collection sites. 18 Timothy S. Young and Joshua Laerm from the main vent. The pH and temperature as measured 16 July 1974 were 7.6 and 21.OC, respectively. Newport Spring also feeds the river about 800 m north of the U.S. Highway 98 bridge. The discharge of the spring as measured 2 March 1972 was 0.23 m3/sec with a pH of 7.8 and water temperature of 19C (Rosenau et al. 1977). Primary depositional site(s) were not located. The fossils are probably eroding out of the banks along much of the length of the river and washed down river by the current. Dense accumulations of fossils may be found in sand deposits, around submerged debris, and in deep holes along the entire length of the river. The St. Marks River with its shallow, relatively clear waters with abundant fossil and archaeological materials has been a popular recreational S.C.U.B.A. diving are for the past 30 years. Local divers report huge quantities of fossils have been recovered by amateur collectors. One of us (J.L.) observed an entire pick-up truck load of fossils being removed in 1978. Local divers report that have collected "tons of it." Although several large private collections of St. Marks material exist, unfortunately they have been mixed with fossils from other regional aquatic systems which makes their inclusion here inappropriate. METHODS We made extensive new collections and reviewed previously collected materials housed at the Florida Museum of Natural History, University of Florida (UF), and the National Museum of Natural History (USNM), Washington, D. C. Our collections are housed at the University of Georgia Museum of Natural History (UGAMNH). Collection efforts were concentrated in six separate locations approximately 3.2 km in either direction from the U.S. Highway 98 bridge that crosses the St. Marks River (Fig. 1). The fossils were collected from 16 to 19 July 1987 by a team of six people from the University of Georgia using S.C.U.B.A. gear. The majority of the fossil materials was collected by hand from these locations along the river. In addition, extensive sand samples were taken at each site for subsequent screening. To preliminarily identify recovered materials we used the Comparative Reference Skeletal Collection of the Zooarchaeology Laboratory, the University of Georgia Museum of Natural History. Reference sources were also used in preliminary identifications. All materials were subsequently taken to the Paleontology Laboratory, the Florida Museum of Natural History, University of Florida, to confirm identifications. Notes were made on the element identified, side, and fusion of bones where possible. St. Marks River Fauna 19 SYSTEMATIC PALEONTOLOGY Standardized common and current scientific names follow Robins et al. (1991) for fishes; Collins (1990) for amphibians and reptiles; American Ornithologists' Union (1983) for birds, and Kurten and Anderson (1980) and Jones et al. (1992) for mammals. Museum acronyms are indicated in the introduction. A complete faunal listing of the species recovered from the St. Marks site is provided in Table 1. CLASS MAMMALIA Order Didelphimorphia Family Didelphidae Didelphis virginiana Kerr Virginia Opossum Material—A single left dentary, UGAMNH1735. Remarks—The single element is identical to that of modern Didelphis virginiana. This was the only marsupial species present in North America during the Pleistocene. It is known from numerous fossil sites in Florida (Webb 1974a). Its stratigraphic range includes Middle Blancan to Recent (Kurten and Anderson 1980). It occurs in a variety of habitats, but it is usually found in forests and woodlands near water (Gardner 1973). We follow Marshall et al. (1990) in the use of the ordinal name Didelphimorphia as do Jones et al. (1992). Order Xenarthra Family Dasypodidae Holmesina septentrionalis (Leidy) Northern Pampathere Material—Right astragulus, UGAMNH2012; right calcaneus, UGAMNH2159; right metacarpus II, UGAMNH1981; two phalanges, UGAMNH1982-1983; numerous dermal plates, UGAMNH1954-1980, 1984- 2029, 2160, 2166. Remarks—The species is known from numerous sites throughout the South and Southeast. Its range is somewhat similar to that of its modern relative, Dasypus novemcinctus, and Holmsina had a similar preference for open woodlands (Kurten and Anderson 1980). Like its modern counterpart, Holmsina probably fed on insects and various invertebrates. Kurten and Anderson (1980) suggest this diet might have restricted them to relatively warm climates where food was available year round. Specimen UHAMNH2159, a right calcaneus, has rodent and carnivore gnaw marks that occurred prior to fossilization. Its stratigraphic range is early Irvingtonian to Wisconsinan (Kurten and Anderson 1980). 20 Timothy S. Young and Joshua Laerm Table 1. List of vertebrate species recovered from the St. Marks River. The figure f indicates extinct forms. Class Mammalia Order Didelphimorphia Family Didelphidae Didelphis virginiana Order Xenarthra Family Dasypodidae \Holmesina septentrionalis Family Megalonychidae ^Megalonyx jeffersonii Family Mylodontidae ^Glossotherium harlani Order Primates Family Hominidae Homo sapiens Order Lagomorpha Family Leporidae Sylvilagus sp. Order Rodentia Family Castoridae Castor canadensis Family Geomyidae Geomys pinetis Family Muridae Microtus sp. Microtus pinetorum Neofiber alleni Ondatra zibethicus Synaptomys australis Order Carnivora Family Mustelidae Mustela sp. Lutra canadensis Mephitis mephitis Family Canidae Canis sp. "\Canis dirus Urocyon cinereoargenteus Family Procyonidae Procyon lotor Family Ursidae Ursus cf. U. americanus Family Felidae Felis sp. ^Smilodon sp. Order Proboscidea Family Mammutidae \Mammuthus jeffersonii Family Elephantidae \Mammut americanum Order Perissodactyla Family Equidae Equus sp. Family Tapiridae ~\Tapirus sp. Order Artiodactyla Family Tayassuidae ^Platygonus compressus Family Suidae Sus scrofa Family Camelidae "\Hemiauchenia macrocephala ^Palaeolama mirifica Family Cervidae Odocoileus virginianus Family Bovidae Bison sp. Bison bison Bos taurus Class Aves Order Podicipediformes Family Podicipedidae Podiceps auritus Podilymbus podiceps Order Pelecaniformes Family Phalacrocoracidae Phalacrocorax auritus Order Ciconiformes Family Ardeidae Ardea herodias Butorides virescens Egretta caerulea Family Threskiornithidae Eudociums albus Order Anseriformes Family Anatidae Aix sponsa Anas acuta St. Marks River Fauna 21 Table 1. Continued. Anas americana Anas discors Anas platyrhynchos Anas sp. Aythya collaris Aythya sp. Branta canadensis Bucephala albeola Lophodytes cucullatus Mergus merganser Order Falconiformes Family Accipitridae Buteo jamaicensis Pandion haliaetus Order Galliformes Family Phasianidae Meleagris gallopavo Order Gruiiformes Family Rallidae Fulica americana Gallinula chloropus Family Aramidae Aramus guarauna Order Strigiformes Family Strigidae Strix varia Class Reptilia Order Testudines Family Chelydridae Chelydra serpentina Family Kinosternidae Gen. et spec, indet. Family Emydidae Pseudemys concinna Pseudemys floridanus Pseudemys nelsoni Trachemys scripta Terrapene Carolina Terrapene Carolina putnami Family Testudinidae Geochelone incisa Geochelone sp. Gopherus polyphemus Family Trionychidae Trionyx sp. Order Squamata Family Colubridae Elaphe obsoleta Gen et spec, indet. Order Crocodilia Family Alligatoridae Alligator mississippiensis Class Amphibia Order Caudata Family Sirenidae Siren sp. Order Anura Gen. et spec, indet. Class Osteichthyii Order Lepisosteiformes Family Lepisosteidae Lepisosteus sp. Order Amiiformes Family Amiidae Amia calva Order Siluriformes Family Ictaluridae PyIodictis cf. P. olivaris Family Ariidae Ariopsis felis Order Salmoniformes Esocidae Esox sp. Order Perciformes Family Percichthyidae Morone saxatilis Family Sparidae Archosargus probatocephalus Family Sciaenidae Sciaenops ocellatus Family Mugilidae Mugil sp. 22 Timothy S. Young and Joshua Laerm Family Megalonychidae Megalonyx jeffersonii (Desmarest) Jefferson's Ground Sloth Material—A single phalanx, UGAMNH2135, and tooth, UGAMNH2136. Remarks—Jefferson's ground sloth occurred in woodlands where it apparently fed on nuts, berries, leaves, and twigs (Stock 1925). It is known from a number of sites in the Southeast including Florida (Webb 1974a), Georgia (Ray 1967), South Carolina (Hay 1923, Roth and Laerm 1980), and Tennessee (Guilday et al. 1969). It could have tolerated a seasonally cool climate as evidenced by its Pleistocene occurrence in what is now Canada and Alaska (McNab 1985). It is reported from Irvingtonian to Rancholabrean sites with a terminal date of 13,890 years B.P., although Kurten and Anderson (1980) suggest it may have survived even longer in Florida. Family Mylodontidae Glossotherium harlani (Owen) Harlan's Ground Sloth Material—Two teeth, UGAMNH2137-2138. Remarks—This was an open plains and grassland species (Stock 1925). It is reported from Irvingtonian to Rancholabrean sites with a terminal date of 13,890 years B.P., although Kurten and Anderson (1980) suggest it may have survived even longer in Florida. Order Primates Family Hominidae Homo sapiens Linnaeus Human Material—Cranial fragment, UF21280. Remarks—This single specimen was recovered by Gillette (19766). Unfortunately, the cranial fragment was not available for examination. We are, therefore, unable to comment on the degree of mineralization. No other human remains were recovered in our efforts. The presence of considerable amounts of Native American cultural material (pottery shards) as well as 18-20th century European-American artifacts indicates the St. Marks River was a site of human occupation before and after European contact. St. Marks River Fauna 23 Order Lagomorpha Family Leporidae Sylvilagus sp. indet. Material—Tooth fragment, UF21301. Remarks—Rabbits are a common component of most Pleistocene sites in Florida. It is surprising no more than a single tooth fragment was encountered in the St. Marks River material. Two species of rabbit occur in the St. Marks region today, the eastern cottontail, Sylvilangus floridanus, and the more common swamp rabbit, S. aquaticus. The former prefers heavy brushy, forested areas with open areas nearby and edges of swamps. The latter is most common in marshes, swamps, and bottomlands (Golley 1962). Order Rodentia Family Castoridae Castor canadensis Kuhl Beaver Material—Left ulna, UGAMNH2126; right upper molar, UGAMNH2125; right M3 , UGAMNH2124; four molars, UF21294. Remarks—Two beaver species occurred in Florida in the late Pleistocene, Castoroides ohioensis and Castor canadensis. Both have even been found in the same deposits (Webb 1974a); however, only the latter is represented in the St. Marks River fauna. The beaver is found in any suitable water habitat including rivers, streams, lakes, and marshes (Lowery 1974). Its relative rarity in the St. Marks may be related to the presence of Alligator. The stratigraphic range is late Blancan to Recent (Kurten and Anderson 1980). Family Geomyidae Geomys pinetis Rafinesque Southeastern Pocket Gopher Material—A single lower fourth premolar, UF21291. Remarks—Geomys pinetis is the only species of pocket gopher in the Southeast. It is associated with the sandy soils of the Coastal Plain (Golley 1962) and is present today in the uplands adjacent the St. Marks River. It is known from late Irvingtonian to Recent (Kurten and Anderson 1980). Family Muridae Microtus sp. indet. Material—Left M3 , UGAMNH2127. Remarks—This fragment, while certainly Microtus, could not be 24 Timothy S. Young and Joshua Laerm referred to a species with confidence. We follow Jones et al. (1992) in their use of the familial name Muridae. Microtus pinetorum (LeConte) Pine Vole Material—Right M2 , UGAMNH2128. Remarks—This molar compares well to modern Microtus pinetorum. Regionally, the pine vole can be found in a wide range of habitats from hardwood and pine forests to overgrown fields (Golley 1962). The stratigraphic range is Sangamonian to Recent (Kurten and Anderson 1980). Neofiber alleni True Round-tailed Muskrat Material—right M2 , UGAMNH2121; right M3 , UGAMNH2123; right M3 , UGAMNH2122; maxilla, UF21293. Remarks—Neofiber alleni is a semi-aquatic mammal that prefers permanent bodies of water with emergent aquatic vegetation (Frazier 1977). Although it has a restricted range today, essentially extreme northern Florida and south Georgia, during the Pleistocene it ranged as far west as Kansas (Hibbard 1943). It is reported from late Irvingtonian to Recent (Kurten and Anderson 1980). The stratigraphic range is Illinoian to Recent (Kurten and Anderson 1980). Ondatra zibethicus (Linnaeus) Muskrat Material—Right dentary with M1 AND M2 , UGAMNH2120; dentary, UF21292. Remarks—The muskrat, like the round-tailed muskrat, is a semi-aquatic mammal that prefers permanent bodies of water (Nelson and Semken 1970). There is not overlap in the range of the two species today. However, Martin and Webb (1974) indicate they were sympatric in at least two late Pleistocene Florida faunas, Devils Den and Ichetucknee River. The occurrence of the two species in the St. Marks River fauna is not overly suggestive that they were sympatric here in the past because of the apparently heterochronous deposition at St. Marks. Furthermore, although the muskrat does not presently occur in the St. Marks River or Apalachicola River drainages, it is known from the extreme western panhandle and the Upper Coastal Plain of Georgia, a distance of 120 km. St. Marks River Fauna 25 Synaptomys australis Simpson Florida Bog Lemming Material—Left mandible with Mb UF21295. Remarks—The specimen referred to in Gillette's (1976b) review of the St. Marks River is the only record of this species at the site. In Florida it is known primarily from Sangamonian and Wisconsinan assemblages, although elsewhere it is known from the Illinoian through the Wisconsinan (Kurten and Anderson 1980). Its presence at Devils Den suggests it might have persisted until about 8,000 years B.P. (Martin and Webb 1974), although this radiocarbon date is considered suspect. The Florida bog lemming is similar morphologically to S. cooperi, the northern bog lemming, but differs considerably in size; it is about 35% larger than S. cooperi. Kurten and Anderson (1980) suggest it might represent a clinal variate of S. cooperi. It was an inhabitant of moist bogs and damp meadows (Burt 1928). Order Carnivora cf. Order Carnivora, gen. et sp. indet. Material—A left coronoid, UGAMNH1881. Remarks—This specimen, though carnivore-like, could not be identified to the familial level. Family Mustelidae Mustela sp. indet. cf. Weasel Material—A single left humerus, UGAMNH1738 and right P3 , UGAMNH1736. Remarks—Two species of weasel, Mustela frenata and M. vison, are common to the region today. Both are known from the Irvingtonian to Recent and are represented in regional fossil sites (Webb 1974a). However, fossil weasels have been reported from very few sites in Florida (Martin 1974, Webb 1974a). Lutra canadensis (Shreber) River Otter Material—Left humerus, UGAMNH1741. Remarks—This material compares well to modern Lutra canadensis. The stratigraphic range includes early Irvingtonian to Recent, and the species is represented in numerous regional sites (Kurten and Anderson 1980). The species occurs in woodlands near rivers and streams but is also known from tidal creeks and marshlands (Lowery 1974). 26 Timothy S. Young and Joshua Laerm Mephitis mephitis (Schreber) Striped Skunk Material—Right mandible, UGAMNH1746; left humerus, UGAMNH1737. Remarks—This material compares well to modern Mephitis mephitis, which can be found in mixed woodlands, brushlands, or prairies but generally in reasonable proximity to water (Lowery 1974). The stratigraphic range is mid Blancan to Recent (Kurten and Anderson 1980). Family Canidae Canis sp. indet. Material— Left ilium, UGAMNH1739; right dentary, UGAMNH1878, 1880; right scapula, UGAMNH1879. Remarks—None of these elements could be identified beyond the generic level. They are well mineralized, suggesting they are not modern C. familiaris contaminants. Several species of Canis are known from late Pleistocene sites in Florida. These include C. lupus, the gray wolf; C. rufus, the red wolf; C. latrans, the coyote; and C. dims, the dire wolf. Martin (1974) has concluded that only two species, C. rufus and C. dirus, are common to middle and late Pleistocene deposits of Florida. Canis lupis is typical of Irvingtonian deposits, whereas C. dirus is representative of the Rancholabrean. Canis dirus Leidy Dire Wolf Material—left radius, UGAMNH1877. Remarks—Canis dirus is known from a number of late Pleistocene sites in Florida (Webb 1914a) and is one of the more common species of mammals at numerous Rancholabrean sites throughout North America. It is thought to have inhabited a wide range of habitats because it was a hunter and scavenger (Kurten and Anderson 1980, Lundelius et al. 1983). The stratigraphic range is early Illinoian to Wisconsinan (Kurten and Anderson 1980). The most recent terminal date for extinction is given at about 8,000 years B.P. in Florida (Martin and Webb 1974), but somewhat earlier (approximately 9,000-10,000 year B.P.) elsewhere (Kurten and Anderson 1980). Urocyon cinereoargenteus (Shreber) Gray Fox Material—Right dentary, UGAMNH1743; left frontal, UGAMNH1744. St. Marks River Fauna 27 Remarks—This material is not well mineralized, which suggests that it is a modern contaminant. However, Urocyon cinereoargenteus would be expected in this fauna. It can be found in a wide range of habitats today, but brushy and woody areas probably best describe the preferred habitat in the South and Gulf Coast area (Lowery 1974). The stratigraphic range in Florida is Middle Rancholabrean to Recent (Martin and Webb 1974). Elsewhere it is known as early as the Blancan (Kurten and Anderson 1980). Family Procyonidae Procyon lotor (Linnaeus) Racoon Material—Three left dentaries, UGAMNH1742, 1747, 1750. A partial skeleton is represented by UF21296. Remarks—The University of Georgia material is not well mineralized, which suggests it could be a modern contaminant, since Procyon lotor is part of the modern fauna. In the Florida panhandle today, the racoon is an inhabitant of forested bottomland swamps. It fossil record in Florida extends from the Late Irvingtonian to Recent (Martin and Webb 1974). Family Ursidae Ursidae gen. et sp. indet. Material—Three phalanges, UGAMNH1745, 1749, 1752. Remarks—Generic identity of this material is uncertain. In addition to the modern black bear, Ursus americanus Pallas, several extinct species of bears are known from the Pleistocene of Florida. These include the cave bear, Tremarctos floridanus (Gidley), and the lesser short-faced bear, Arctodus pristinus Leidy, all of which persisted at least until the late Wisconsin (Kurten and Anderson 1980). Ursus cf. U. americanus Pallas cf. Black Bear Material—A single right dentary with M1? UGAMNH1751. Remarks—This specimen is well mineralized, but it is too worn for positive identification. The black bear can be found in forests and bottomland swamps throughout much of the Southeast (Golley 1962, Lowery 1974). It is represented in numerous late Pleistocene sites. The stratigraphic range is early Irvingtonian to Recent (Kurten and Anderson 1980). 28 Timothy S. Young and Joshua Laerm Family Felidae Felis sp. indet. Material—Left radius, UGAMNH1740. Remarks—This specimen is a large Felis, but it is too worn for positive identification. Webb (1974a) states that several species of Felis are known from the Late Pleistocene of Florida, and include F. atrox Leidy, F. concolor Linnaeus, F. onca (Linnaeus), F. pardalis Linnaeus, F. rufus Schreber, and F. yagouaroundi Geoffroy. Another possibility is Felis amnicola, a new, small cat described by Gillette (1976a). The description is based on several specimens from various localities in Florida and possibly Georgia. Smilodon sp. indet. Material—Left scapho-lunar, UGAMNH1748. Remarks—The sabertooth cats are reported from a dozen or more late Pleistocene sites in Florida (Webb 1974a, Kurten and Anderson 1980). Slaughter (1963) proposed a series of successional changes in Smilodon species throughout the North American Pleistocene. Webb (1974a) concurs that records of Smilodon in Florida support such a successional outline: Smilodon gracilis is a late Blancan and early Irvingtonian; S. fatalis is representative of late Irvingtonian and early Rancholabrean sites; and that S. floridanus is typical of the late Rancholabrean. The temporal span reflected by other faunal elements from the St. Marks would be more suggestive of the latter species; however, given the similarity of these species, more precise identification is impossible from the limited available material. Smilodon could probably have been found in habitats ranging from grassland to woodland (Merriam and Stock 1932, Lundelius et al. 1983). Order Proboscidea Proboscidea gen. et sp. indet. Material—Sesamoid, UGAMNH1098; tusk fragment, UF21255; skull fragment, UF21256; leg fragment, UF21257; vertebral fragment UF21258. Remarks—These specimens are very definitely proboscidean, but assignment to species is impossible. Family Mammutidae Mammut americanum (Kerr) American Mastodon Material—Axis fragment, UGAMNH1614; tooth fragments, UGAMNH1612, 1613, 1615, 1616, UF21267 and 21276; tusk fragments, UF21277-21278; proximal humerus, UF21279; calcaneus, UF21290. St. Marks River Fauna 29 Remarks—The morphology of the elements is consistent with its identification as Mammut americanum. Dreimanis (1968) suggested that M. americanum inhabited coniferous forests. The stratigraphic range is early Blancan to Wisconsinan (Kurten and Anderson 1980). Family Elephantidae Mammuthus jeffersonii (Osborn) Jefferson's Mammoth Material—Tooth fragments, UGAMNH1607-1611; tooth fragments, UF21259-21262, 21264-21266; mandibular symphysis, UF21263. Remarks—The morphology of the tooth fragments and mandibular symphysis is consistent with its identification as Mammuthus jeffersonii. Jefferson's mammoth probably inhabited open grasslands (Stock 1963, Harrington et al. 1974). The stratigraphic range is Illinoian to Wisconsinan (Kurten and Anderson 1980). Order Perissodactyla Family Equidae Equus sp. indet. Horse Material—left astragalus, UGAMNH1035; cervical vertebra, UGAMNH1170; left upper cheek tooth, UGAMNH1045; right upper cheek tooth, UGAMNH1031; right lower cheek tooth, UGAMNH1048; cheek tooth, UGAMNH1046, 1047, 1062; right deciduous P 2 , UGAMNH1042; left deciduous P2 , UGAMNH1061; right cuneiform, UGAMNH1049; left femoral head, UGAMNH1034; right distal humeral epiphysis, UGAMNH1054; left I 3 , UGAMNH1036; right I 2 , UGAMNH1041; lower incisor, UGAMNH1056; left I 3 , UGAMNH1059; left I 1 , UGAMNH1060; incisive fragment, UGAMNH1032; left upper molar, UGAMNH1038; right M2,UGAMNH1039; right M3 , UGAMNH1044; left M3 , UGAMNH1057; upper molar fragment UGAMNH1029; left navicular, UGAMNH1063; medial phalanges UGAMNH1030, 1050, 1053; distal phalanx, UGAMNH1051; proximal phalanx, UGAMNH1055; left P 2 , UGAMNH1037; right P 2 , UGAMNH1040; left upper premolar, UGAMNH1043; right lower premolar, UGAMNH1058; left scapula, UGAMNH1052; sesamoid, UGAMNH1033; medial phalanx, UF21228; teeth, UF21229-21238; teeth UF21240-UF21254; cheek tooth, axis, and pelvis, UF21297. Remarks—Equus is well represented in St. Marks River. A portion of the material is poorly mineralized and probably represent contaminants of the modern E. caballus. However, the majority of elements are well fossilized, and it is likely that most of the material 30 Timothy S. Young and Joshua Laerm is of late Pleistocene origin. Given the uncertain relationships of late Pleistocene horses in general and the likelihood of heterochronous deposition, we did not assign the material to a particular species. Pleistocene Equus was generally a grassland species (Kurten and Anderson 1980). Family Tapiridae Tapiridae gen. et spec, indet. Material—Left dentary, UGAMNH2068. Remarks—This edentulous specimen could not be assigned to Tapirus with confidence, although the morphology is similar. Tapirus sp. indet. Tapir Material—Right upper deciduous premolar, UGAMNH2070, left upper deciduous premolar, UGAMNH2071; right fibula, UGAMNH2069. Remarks—The available material, while certainly Tapirus, could not be referred to a species with confidence. Tapirs occur in wet woodlands (Simpson 1945, Gray and Crammer 1961). Order Artiodactyla Family Tayassuidae Platygonus compressus LeConte Flat-headed Peccary Material—Axis, UGAMNH2072. Remarks—The material has the diagnostic characters of Platygonus compressus which is thought to have wide environmental tolerances, but was probably most associated with open woodlands (Martin and Guilday 1967, Ray et al. 1970). The stratigraphic range is Sangamonian to Wisconsinan (Kurten and Anderson 1980). Family Suidae Sus scrofa Linnaeus Pig Material—Left maxilla with P3 and P4 , UGAMNH1159; right humeral fragment, UGAMNH1160; right femoral diaphysis, UGAMNH1162; right radial fragment, UGAMNH1161; left humeral fragments, UGAMNH1163, 1158; right ilial fragment, UGAMNH1157; distal humeral fragment, UGAMNH1178; left femur, UGAMNH1156. Remarks—None of the pig material showed evidence of significant mineralization. The pig was introduced during historic times and represents a domesticate. Specimen UGAMNH1178 shows marks of a saw. St. Marks River Fauna 31 Family Camelidae Camelidae gen. et sp. indet. Material—Proximal phalanges, UGAMNH1647, 1648; right scaphoid, UGAMNH1649; right astragalus, UGAMNH1650; right scapula, UGAMNH1651; left proximal femoral fragment, UGAMNH1652. Remarks—These specimens have distinctive camelid familial characters, but cannot be assigned to a particular species. Hemiauchenia macrocephala (Cope) Large-headed Llama Material—Proximal phalanges, UGAMNH2151, 2152. Remarks—The identification of these elements to Hemiauchenia macrocephala is based on the size of the phalanges. According to Webb (1974/?), H. macrocephala was a plains and grasslands inhabitant. The stratigraphic range is Wisconsinan to Recent (Kurten and Anderson 1980). Because this species has such a limited stratigraphic range, at least a portion of the deposit can be correlated to the Wisconsinan. Paleolama mirifica (Simpson) Stout-legged Llama Material—Left proximal metacarpal fragment, UGAMNH2146; right humerus, UGAMNH2145; left metatarsus, UGAMNH2158; left M3, UGAMNH2147; metapodial, UGAMNH2148; right distal humerus, UGAMNH2149; left pisiform, UGAMNH2150. Remarks—The stratigraphic range is late Irvingtonian to Wisconsinan (Kurten and Anderson 1980). Webb (1974b) reports them to be an inhabitant of grasslands and savannahs. Specimen UGAMNH2158, a left metatarsus, has longitudinal cracks indicative of weathering prior to fossilization. Family Cervidae Odocoileus virginianus (Zimmerman) White-tailed Deer Material—Antler fragments, UGAMNH1103, 1106, 1134, 1148; left astragalus, UGAMNH1101, 1124, 1210, 1757; right astragalus, UGAMNH1071, 1113, 1125, 1677, 1758, 2162; right calcaneus, UGAMNH1150, 1181, 1204, 1756; left calcaneus, UGAMNH1069, 1073, 1077, 1079, 1205, 1653, 2153, 2154, 1667, 1678; right cubonavicular, UGAMNH1074, 1111; right dentary with P1? P2 , P3 , M2 , M3 , UGAMNH1081; left dentary with Mlt UGAMNH1131; right dentary with Mh M2 , M3 , UGAMNH1108, 1126; right dentary with P2, P3 , M1? M2 , M3 , UGAMNH1191; right proximal femoral fragment, UGAMNH1184; femoral diaphysis, UGAMNH1064; left femoral head, 32 Timothy S. Young and Joshua Laerm UGAMNH1129; left femoral diaphysis, UGAMNH2163; right distal femur, UGAMNH11139; left distal femoral fragment, UGAMNH1085, 1088; left femoral lesser trochanter, UGAMNH1130; right femoral diaphysis, UGAMNH1139, 2164; right frontal with antler, UGAMNH1068, 1099; left frontal with antler, UGAMNH1012, 1082, 1118, 1121, 1666; frontal with antler pedicle, UGAMNH1196, 1203; right humeral fragments, UGAMNH1075, 1100, 1122, 1133, 1142, 1661, 1668; left humeral fragments, UGAMNH1070, 1093, 1136, 1137, 1143, 1185, 1186, 1235, 1679; right ilial fragments, UGAMNH1076, 1090, 1681; left ilial fragments, UGAMNH1079, 1086, 1147; left ischial fragments, UGAMNH1105, 1119, 1682; right lunate, UGAMNH1112; right maxilla with P4 , M1 , UGAMNH1206; left metacarpal fragments, UGAMNH1087, 1102, 1114, 1115, 1146, 1180, 1198, 1663; right metacarpal fragments, UGAMNH1072, 1084, 1092, 1097, 1116, 1665; metacarpal diaphysial fragments, UGAMNH1104, 1193-1195, 1670; right metatarsal fragments, UGAMNH1091, 1183, 1201, 1659, 1669, 1675, 1759; metatarsal diaphysial fragments, UGAMNH1117, 1199, 1200, 1208, 1212, 1656, 1672, 1673, 1676; left metatarsal fragments, UGAMNH1120, 1192, 1654, 1655, 1662, 1664, 1671; right M1 , UGAMNH1109, UGAMNH1151; left M3 , UGAMNH1289; right M3 , UGAMNH1190; left petrous, UGAMNH1110, 1128, 1202, 1214; medial phalanx, UGAMNH1080, 1141, 1209, 1213, 1753, 1754; proximal phalanx, UGAMNH1078, 1109, 1127, 1182, UGAMNH1211, 1755; left radial fragment, UGAMNH1065; right radial diaphysial fragment, UGAMNH1067; left radial fragments, UGAMNH1207, 2165; right radial fragments, UGAMNH1144; sacrum, UGAMNH1089; left scapular fragments, UGAMNH1140, 1188; right scapular fragments, UGAMNH1094, 1145; right distal tibial fragments, UGAMNH1095, 1123, 1658, 1680; left distal tibial fragments, UGAMNH1096, 1657; left proximal tibial fragment, UGAMNH1674; right proximal ulnar fragment, UGAMNH1132; left proximal ulnar fragment, UGAMNH1197; thoracic vertebral fragment, UGAMNH1760; lumbar vertebral fragment, UGAMNH1187; cervical vertebral fragment, UGAMNH1066; atlar fragments, UGAMNH1083, 1149; axial fragment, UGAMNH1660; antler, UF21289; five mandibles, UF21298. Remarks—The deer material shows a considerable range of mineralization. A significant portion is poorly mineralized and probably represents modern contaminants. The remaining material, however, is well mineralized, but mineralization alone is a poor indicator of possible Pleistocene age. The stratigraphic range of species is middle Blancan to Recent (Kurten and Anderson 1980). Odocoileus is a woodland and forest edge species (Golley 1962, Lowery 1974, Lundelius et al. 1983). St. Marks River Fauna 33 Family Bovidae Bovidae gen. et sp. indet. Material—Proximal phalanges, UGAMNH1621, 1627, 1631, 1632; left lunate, UGAMNH1633, 1634; left lunar, UGAMNH1625; left scapual spine, UGAMNH1635; rib head, UGAMNH1636; right scapula, UGAMNH1622; right P4, UGAMNH1623; left distal humerus, UGAMNH1624; left proximal femur, UGAMNH1626; right distal humeral epiphysis, UGAMNH1628; metatarsal diaphysial fragment, UGAMNH1629; left proximal tibial fragment, UGAMNH1630; tooth fragments, UF21239, 21281, 21282, 21285, 21288; distal humerus, UF21283; horn core tip, UF21284. Remarks—These elements are definitely bovid but the available material does not permit specific distinction. Bison bison (Linnaeus) Bison Material—-Right M2 , UGAMNH1620; left P3 , UGAMNH1619; left P2, UGAMNH1618; right M2 , UGAMNH1617; molar UF2299. Remarks—While Jones et al. (1992) have employed Bos bison for the American bison, we continue the traditional use of Bison bison. Two species of bison are known from Florida. The giant bison, B. latifrons, is known from Illinoian and Sangamonian and survived up until the late Wisconsinan. The American buffalo or bison, B. bison, was widespread throughout the Wisconsinan through the Recent (Kurten and Anderson 1980). Bison is typically associated with grasslands, though in the Southeast may well have ranged into woodlands (Golley 1962, Stock 1963). It became extinct in the southeastern United States early in the 19th Century. Bos taurus Linnaeus Cow Material—Right proximal humeral diaphysis, UGAMNH10863; left scapula, UGAMNH1155; left ilium, UGAMNH1152, 1154; right metatarsal diaphysis, UGAMNH117; right proximal humerus, UGAMNH1177; right distal humerus, UGAMNH1176; orbital portion of right maxilla, UGAMNH1 153; right proximal tibia, UGAMNH1171; right distal femoral epiphysis, UGAMNH1172; right astragalus, UGAMNH1173; distal phalanx, UGAMNH1174; metapodial, UF21300. Remarks—Bos taurus was introduced into North America sometime after 1492. All elements were poorly mineralized. The presence of cow indicates the site has modern contaminants. 34 Timothy S. Young and Joshua Laerm CLASS AVES Order Podicipediformes Family Podicipedidae Podiceps auritus (Linnaeus) Horned Grebe Material—Distal portion of left ulna, USNM209968. Remarks—Today the species winters in coastal areas and infrequently occurs in freshwater (Sprunt 1954). Podilymbus podiceps (Linnaeus) Pied-billed Grebe Material—Right humerus, USNM210293, 210294, 210301, 210302; humerus, USNM210311; right proximal humerus, USNM210304; left humerus, USNM210307; left tibial fragments, USNM210292, 210308, 210309, 210315, 210322, 210325, 210327; right tibia, USNM210297, 210300, 210312, 210316, 210317; left ulna, USNM210296, 210303, 210305, 210306, 210310, 210328, 210329; right ulna USNM210313, 210321; radius, USNM210298; right coracoid, USNM210319; left coracoid, USNM210320; left femur, USNM210298; right carpometacarpus, USNM210321; left carpometacarpus, USNM210323; scapula, USNM210324; pedal phalanx, USNM210326; tarsometatarsus, USNM210314, 210318; left tibia, USNM210306. Remarks—The species inhabits freshwater marshes and ponds, but also is associated with saltwater in winter (Sprunt 1954). Order Pelecaniformes Family Phalacrocoracidae Phalacrocorax auritus (Lesson) Double-crested Cormorant Material—Left radius, USNM209845; left ulna, USNM209844; scapula, USNM209859; anterior sternum, USNM209843; left coracoid, USNM209858; phalanx 1 of digit II, USNM209861, 209852; phalanx 2 of digit II, USNM209856; left humerus, USNM209846; right humerus, USNM209851; proximal radius, USNM209860; right ulna, USNM209852; distal tibia, USNM209848; right femur, USNM209854; right mandible, USNM209855; left mandible, USNM209850; sternal fragment, USNM209857; left coracoid, USNM209849; right coracoid, USNM209847. Remarks—This species is distributed in large rivers and lakes as well as brackish and saltwater systems (Sprunt 1954). St. Marks River Fauna 35 Order Ciconiformes Family Ardeidae Ardea herodias Linnaeus Great Blue Heron Material—Cervical vertebrae, USNM210282, 210283, 210285, 210287; right mandible, USNM210281; mandible fragments, USNM210280, 210286, 210288; maxilla fragment, USNM210279; right coracoid, USNM210291; right proximal humerus, USNM210289; distal tarsometatarsus, USNM210290; right carpometacarpus, USNM210279. Remarks—The great blue heron has wide ecological tolerances, occurring in freshwater swamps and riparian habitats as well as saltwater marshes (Sprunt 1954). Butorides striatus (Linnaeus) Green-backed Heron Material—Right humerus, USNM209966. Remarks—Butorides striatus and B. virescens, sometimes regarded as separate species, are recognized as geographic races of B. striatus by the American Ornithologists Union (1983). It occurs along lake margins, streams, ponds, and freshwater and saltwater marshes (Sprunt 1954). Egretta caerulea (Linnaeus) Little Blue Heron Material—Mandibular tip with right ramus, USNM209862. Remarks—Freshwater swamps and saltwater marshes are the preferred habitats (Sprunt 1954). Family Threskiornithidae Eudocimus albus (Linnaeus) White Ibis Material—Right humerus, USNM209971; left proximal coracoid, USNM209972. Remarks—Eudocibus albus is associated with swampy forests, marshy sloughs, and saltwater marshes (Sprunt 1954). Order Anseriformes Family Anatidae Aix sponsa (Linnaeus) Wood Duck Material—Right carpometacarpus, USNM209931, 209938, 209939, 209944; left carpometacarpus, USNM209934; right ulna, 36 Timothy S. Young and Joshua Laerm USNM209927, 209946; left ulna, USNM209926, 209940, 209945; left humerus, USNM209928, 209930, 209932, 209933; right humerus, USNM209941; radius, USNM209929, 209936; scapula, USNM209942, 209943; proximal tibia, USNM209935; right coracoid, USNM20992; right femur, USNM209925; right tarsometatarsus, USNM209937. Remarks—The species is common today in freshwater woodland rivers, ponds, and marshes (Sprunt 1954). Anas sp. indet. Material—Right ulna, UGAMNH2078. Anas acuta Linnaeus North Pintail Material—Left coracoid, USNM209965. Remarks—The pintail is associated with freshwater marshes, ponds, and lakes (Sprunt 1954). Anas americana Gmelin American Wigeon Material—Left humerus, USNM210270; left ulna, USNM210267, 210273; right scapula, USNM210278; scapula USNM210272, 210274; right coracoid, USNM210268, 210275, 210276; right ulna, USNM210277; phalanx 1 of digit II, USNM210269; radius, USNM210271. Remarks—This species is an inhabitant of freshwater marshes, ponds, and shallow lakes (Sprunt 1954). Anas discors Linnaeus Blue-winged Teal Material—Left carpometacarpus, USNM209865, 209866, 209872- 209874; right carpometacarpus, USNM209870, 209871; right humerus, USNM209869; right coracoid, USNM209868; left ulna, USNM209864. Remarks—Sprunt (1954) reports the species from freshwater ponds and lakes. Anas platyrhynchos Linnaeus Mallard Material—Left humerus, USNM209910, 209911, 209914; right humerus, USNM209913, 209918; right scapula, USNM209916; left scapula, USNM209917; left coracoid, USNM209919; right coracoid, USNM209912; right carpometacarpus, USNM209920; furcula, USNM209915. St. Marks River Fauna 37 Remarks—The mallard prefers freshwater lakes and marshes (Sprunt 1954). Aythya sp. indet. Material—Right carpometacarpus, UGAMNH2073; left distal tibiotarsus, UGAMNH2077. Aythya collaris (Donovan) Ring-necked Duck Material—Humerus shaft, USNM209899; left humerus, USNM209884, 209878, 209886, 209890, 209894, USNM209910; right humerus, USNM209877, 209893, 209903; left ulna, 209885, 209888, 209897; right ulna, USNM209878, 209892, 209905-209907; left tibia, USNM209880, 209908; right tibia, USNM209909; left carpometacarpus, 209898, 209900; right tarsometatarsus, USNM209881, 209889; left tarsometatarsus, USNM209887; tarsometatarsus, USNM209902; right coracoid, 209895, 209896; proximal radius, USNM209882; distal radius, USNM209883; radius, USNM209904; right scapula, USNM209891; cervical vertebra, USNM209901. Remarks—This species is associated most commonly with wooded lakes, ponds, and rivers, but also is reported from saltwater systems (Sprunt 1954). Branta canadensis (Linnaeus) Canada Goose Material—Right coracoid, UGAMNH2074; right tarsometatarsus, USNM209875; right distal carpometacarpus, USNM209876. Remarks—Both USNM specimens from the 1970s are noted by Storrs Olson (personal communication) as small and possibly represent either a small subspecies or juveniles. The UGAMNH specimen from 1987 is large. Sprunt (1954:53) states the center of abundance in Florida for modern Branta canadensis is the St. Marks Refuge. This coracoid could possibly be assigned to Branta cf. B. dickeyi on the basis of size. Steven Emslie (Point Reyes Bird Observatory, personal communication) examined the St. Marks River specimen and thought it could be assigned to B. dickeyi. Measurements of the coracoid are larger than modern B. canadensis, but there is some overlap. Emslie (personal communication) reported a large B. dickeyi from the early Pleistocene of Florida. We refer the coracoid conservatively to B. canadensis. The species prefers freshwater lakes, rivers, and marshes (Sprunt 1954). 38 Timothy S. Young and Joshua Laerm Bucephala albeola (Linnaeus) Bufflehead Material—Right carpometacarpus, USNM209969. Remarks—The bufflehead is most common in saltwater bays and estuaries, and rarely in freshwater lakes and ponds (Sprunt 1954). Lophodytes cucullatus (Linnaeus) Hooded Merganser Material—Right proximal humerus USNM209975; right distal humerus USNM209976, 209980; right humerus, USNM209977; left humerus, USNM209978; left ulna, USNM209979. Remarks—The species occurs in freshwater wooded ponds, rivers, and lakes (Sprunt 1954). Mergus merganser Linnaeus Common Merganser Material—Left distal tarsometatarsus, USNM209863. Remarks—The common merganser inhabits wooded freshwater rivers and ponds but winters in saltwater bays (Sprunt 1954). Order Falconiformes Family Accipitridae Pandion haliaetus (Linnaeus) Osprey Material—Right distal tarsometatarsus, USNM209967. Remarks—The species prefers fresh and saltwater marshes, lakes, and bays (Sprunt 1954). Buteo jamaicensis (Gmelin) Red-tailed Hawk Material—Left distal humerus, USNM209970. Remarks—The red-tailed hawk is most common in deciduous forests adjacent to open grasslands (Sprunt 1954). Order Galliformes Family Phasianidae Meleagris gallopavo Linnaeus Wild Turkey Material—Left tarsometatarsus, UGAMNH2075; right proximal tibiotarsus, USNM209921; right proximal femur, USNM209922; tarsometatarsus shaft, USNM209923. Remarks—The species is known from drier swamps, open pine, and hardwoods as well as prairies (Sprunt 1954). St. Marks River Fauna 39 Order Gruiformes Family Rallidae Fulica americana (Gmelin) American Coot Material—Left distal tarsometatarsus, UGAMNH2076; left tibia, USNM209947, 209951; left distal tibiotarsus, USNM209956, 209958; right distal tibiotarsus, USNM20949, 209952, 209962; right tibiotarsus, USNM209961; tibiotarsus shaft, USNM209955; left ulna, USNM209954, 209966; right carpometacarpus, USNM209950, 209963; right distal femur, USNM209948; distal humerus, USNM209953; right scapula, USNM209959; left coracoid, USNM209960. Remarks—The American coot is primarily associated with open freshwater ponds and marshes (Sprunt 1954). Gallinula chloropus (Linnaeus) Common Moorhen Material—Right tarsometatarsus, USNM210259, 210260, 210255; right tibiotarsus, USNM210257; radius, USNM210256; left phalanx 1 of digit II, USNM210258. Remarks—This species prefers freshwater marshes and ponds with heavy aquatic vegetation (Sprunt 1954). Family Aramidae Aramas guarauna (Linnaeus) Limpkin Material—Left tarsometatarsus, USNM210262, 210266; right tarsometatarsus, USNM210261; right distal tarsometatarsus, USNM210265; left distal tibiotarsus, USNM210263; right distal tibiotarsus, USNM210264. Remarks—The limpkin is associated with open, freshwater swamps and marshes (Sprunt 1954). Order Strigiformes Family Strigidae Strix varia Barton Barred Owl Material—Right proximal femur, USNM209973; right tibiotarsus shaft, USNM209974. Remarks—The barred owl occurs in low, wet woodlands and swampy forests (Sprunt 1954). 40 Timothy S. Young and Joshua Laerm CLASS REPTILIA Order Testudines Family Kinosternidae Kinosternidae gen. et sp. indet. Material—Nuchal, UGAMNH2038, 2047; right peripheral 1, UGAMNH2041; left peripheral 2, UGAMNH2042; left peripheral 4, UGAMNH2052; right peripheral 4, UGAMNH2044; left peripheral 9, UGAMNH2053; right peripheral 10, UGAMNH2048; plastron fragment, UGAMNH2050, 2051; right pleural 1, UGAMNH2039; left pleural 1, UGAMNH2045; right pleural 2, UGAMNH2043; right pleural 6, UGAMNH2040; pleural fragments, UGAMNH2046, 2049. Remarks—None of the kinosternid material could be referred to genus or species. Family Chelydridae Chelydra serpentina (Linnaeus) Snapping Turtle Material—Right peripheral, UGAMNH2034, 2037; left peripheral 4, UGAMNH2035; peripheral UGAMNH2036. Remarks—This material compares well with modern Chelydra serpentina. The species prefers permanent freshwater systems (Conant 1975). Family Emydidae Emydidae gen. et spec, indet. Material—Right epiplastron, UGAMNH1350, 1351, 1355, 1356, 1366, 1368, 1370, 1402, 1403, 1405, 1444, 1462, 1482, 1510, 1530, 1535, 1538, 1546, 1548-1550, 1554, 1872; left epiplastron, UGAMNH1235, 1268, 1282, 1285, 1307, 1343, 1345, 1346, 1349, 1359, 1362, 1375, 1380, 1390, 1394, 1445, 1511, 1551; left humerus, UGAMNH1498; right hypoplastron at inguinal notch, UGAMNH1217, 1237, 1241, 1242, 1281, 1308, 1316, 1322, 1324, 1325, 1357, 1379, 1382, 1418, 1419, 1467, 1469, 1501, 1601, 1871; right hypoplastron at axillary notch, UGAMNH1238, 1240, 1301, 1352, 1358, 1404, 1452, 1474, 1503, 1547, 1553, 1555, 1565; right hypoplastron, UGAMNH1164, 1167, 1215, 1216, 1219, 1221, 1236, 1283, 1333, 1344, 1354, 1376, 1388, 1389, 1429, 1433, 1470, 1473, 1480, 1495, 1521, 1533, 1572, 1575, 1584, 1600, 1867, 1870; left hypoplastron at axial notch, UGAMNH1290, 1361, 1369, 1566, 1602; left hypoplastron at inguinal notch, UGAMNH1168, 1220, 1269, 1280, 1284, 1300, 1302, 1413, 1447, 1449, 1516, 1522, 1527, 1559, 1567, 1582; left hypoplastron, UGAMNH1218, 1222-1225, 1239, 1278, 1279, 1293, 1320, 1342, 1377, 1396, 1423, 1456, 1464, 1471, St. Marks River Fauna 41 1475, 1509, 1513, 1519, 1526, 1532, 1560, 1564, 1569, 1571, 1574, 1578, 1583; neural 1, UGAMNH1328, 1439, 1545, 1558; neural 2, UGAMNH1568, UGAMNH1588; neural 3, UGAMNH1573, 1581; neural 6, UGAMNH1461, 1577; neural 7, UGAMNH1260, 1341, 1579, 1874; neural 8, UGAMNH1271; neural 9, UGAMNH1410; neural, UGAMNH1233, 1258, 1259, 1277, 1291, 1309, 1310, 1312, 1363, 1364, 1372, 1384, 1392, 1393, 1406, 1409, 1414, 1440, 1441, 1494, 1524, 1563, 1570, 1576, 1580, 1589, 2139, 2141; nuchal, UGAMNH1261- 1264, 1313, 1411, 1417, 1427, 1457, 1486, 1504, 1508, 1518, 1595; right periphal 1, UGAMNH1165, 1321, 1399, 1398, 1451, 1489, 1525, 1528, 1592, 1866; left peripheral 1, UGAMNH1231, 1245, 1303, 1319, 1454, 1505, 1562; right peripheral 2, UGAMNH1381, 1397, 1407; left peripheral 2, UGAMNH1275, 1298, 1400; right peripheral 3, UGAMNH1169, 1294, 1442, 1531; left peripheral 3, UGAMNH1416, 1421, 1472; right peripheral 4, UGAMNH1540; left peripheral 4, UGAMNH2167; right peripheral 5, UGAMNH1395; left peripheral 5, UGAMNH1425, 1591; right peripheral 6, UGAMNH1244, 1517; left peripheral 6, UGAMNH1274, 1296, 1446; right peripheral 7, UGAMNH1246, 1428, 1594; left peripheral 7, UGAMNH1329, 1432, 1453, 1455; right peripheral 8, UGAMNH1232, 1552, 2143; left peripheral 8, 1424, 1542; right peripheral 9, UGAMNH1166, 1373, 1484, 1490; left peripheral 9, UGAMNH1249, 1299, 1492; right peripheral 10, H1273, 1276; left peripheral 10, UGAMNH1248; right peripheral 11, UGAMNH1326, 1332, 1429, 1587, 1597; left peripheral 11, UGAMNH1297, 1304, 1408, 1435; peripheral UGAMNH1000, 1243, 1247, 1311, 1420, 1426, 1434, 1442, 1449, 1450, 1536, 1593, 1604; right pleural 1, UGAMNH1234, 1334, 1336, 1374, 1385, 1437, 1485, 1493, 1502, 1554, 1875; left pleural 1, UGAMNH1365, 1371, 1378, 1391, 1438, 1468, 1487, 1507, 2142; left pleural 2, UGAMNH1292; right pleural 2, UGAMNH1465, UGAMNH1491; left pleural 3, UGAMNH1431; right pleural 3, UGAMNH1436, UGAMNH1430; left pleural 4, UGAMNH1460; right pleural 5, UGAMNH1596; left pleural 5, UGAMNH1492; right pleural 6, UGAMNH1488; right pleural 7, UGAMNH1340; Pleural, UGAMNH1265-1267, 1270, 1286-1289, 1295, 1305, 1306, 1314, 1318, 1330, 1339, 1348, 1352, 1367, 1383, 1387, 1401, 1483, 1537, 1543, 1585, 1586, 1590, 1598, 1599, 1603, 1868, 1873; pygal, UGAMNH1317, 1323, 1422, 1458, 1476, 1556, 1561; left scapula, UGAMNH1496, 1497; right scapula, UGAMNH1665; suprapygal, UGAMNH1499; right xiphiplastron, UGAMNH1226- 1230, 1250, 1251, 1253, 1255, 1257, 1331, 1347, 1360, 1415, 1463, 1472, 1479, 1481, 1869, 1876; left xiphiplastron, UGAMNH1252, 1254, 1256, 1315, 1335, 1337, 1338, 1386, 1512, 1514, 1515, 1523, 1539, 1544, 1459, 1478. 42 Timothy S. Young and Joshua Laerm Remarks—Most of the emydid material could only be identified to the familial level. Species level identification is difficult and requires nearly complete elements. Almost all the material was well mineralized. We are confident that the majority represents Pleistocene deposition as opposed to Recent. Pseudemys concinna (LeConte) River Cooter Material—Left peripheral 3, UGAMNH1882; left peripheral 4, UGAMNH1885; right peripheral 7, UGAMNH1884; right peripheral 11, UGAMNH1883. Remarks—Pseudemys concinna is distinguished by its distinctive carapace. It is most common in slow streams and rivers (Conant 1975). Pseudemys floridana (LeConte) Cooter Material—Left peripheral 3, UGAMNH2030; left pleural 3, UGAMNH2031; left pleural 4, UGAMNH2033; nuchal, UGAMNH2032. Remarks—The species is most commonly associated with permanent bodies of freshwater including swamps and rivers (Conant 1975). Pseudemys nelsoni Carr Florida Redbelly Turtle Material—Entoplastron, UGAMNH1904; right epiplastron, UGAMNH1920; right hypoplastron axial notch, UGAMNH1889, 1913, 1938; left hypoplastron axial notch UGAMNH1928, 1940; right hypoplastron inguinal notch, UGAMNH1908, 1943; left hypoplastron inguinal notch, UGAMNH1897, 1905; neural 7, UGAMNH1901; neural, UGAMNH1887; nuchal, UGAMNH1914, 1953; right peripheral 1, UGAMNH1899, 1906; right peripheral 2, UGAMNH1929; right peripheral 3, UGAMNH1937; left peripheral 3, UGAMNH1917; right peripheral 4, UGAMNH1930; left peripheral 5, UGAMNH1900; left peripheral 7, UGAMNH1890, 1898; right peripheral 8, UGAMNH1942, 1950; left peripheral 8, UGAMNH1945, 1948; right peripheral 9, UGAMNH1915; left peripheral 9, UGAMNH1886; left peripheral 10, UGAMNH1946, 1947; right peripheral 11, UGAMNH1506, 1918, 1919, 1941; left peripheral 11, UGAMNH1944; peripheral, UGAMNH1907, 1909, 1949; right pleural 1, UGAMNH1534, 1892, 1910; left pleural 1, UGAMNH1922, 1951, 2140; left pleural 2, UGAMNH1917; left pleural 3, UGAMNH1912; left pleural 4, UGAMNH1933; right pleural 5, UGAMNH1934; pleural, UGAMNH1891, 1893-1895, 1903, 1916, St. Marks River Fauna 43 1921, 1923-1927, 1931, UGAMNH1932, 1935, 1936, 1939; suprapygal, UGAMNH1888; right xiphiplastron, UGAMNH1896; left xiphiplastron UGAMNH1902, 1952. Remarks—This is a species associated with freshwater sloughs, marshes, streams, and ponds (Conant 1975). Trachemys scripta (Schoepff) Slider Material—Entoplastron, UGAMNH1763, 1792, 1801, 1831; right hypoplastron axial notch, UGAMNH1780; left hypoplastron axial notch, UGAMNH1828, 1833; right hypoplastron inguinal notch, UGAMNH1774; left hypoplastron inguinal notch, UGAMNH1819; neural 1, UGAMNH1790, 1834; neural 3, UGAMNH1789; neural 8, UGAMNH1846; neural, UGAMNH1766, 1769, 1819, 1837, 1842; nuchal, UGAMNH1520, 1764, 1773, 1776, 1778, 1782, 1784, 1787, 1791, 1823, 1841, 1844; right peripheral 1, UGAMNH1768, 1770, 1826, 1827; left peripheral 1, UGAMNH1783, 1840; right peripheral 2, UGAMNH1765, 1798, 1802; left peripheral 2, UGAMNH1776, 1806; right peripheral 3, UGAMNH1794; left peripheral 3, UGAMNH1796; left peripheral 5, UGAMNH1775; left peripheral 8, UGAMNH1835, 1836, right peripheral 9, UGAMNH1779, 1793; left peripheral 10, UGAMNH1767, 1820, 1839; right peripheral 11, UGAMNH1781, 1843; left peripheral 11, UGAMNH1762, 1803-1805, 1816, 1825, 1832, 1845; peripheral, UGAMNH1785, 1824, 1829; left pleural 1, UGAMNH1807; right pleural 2, UGAMNH1799; left pleural 2, UGAMNH1800; left pleural 4, UGAMNH1788; pleural, UGAMNH1771, 1772, 1786, 1808- 1815, 1821, 1822, 1838; pygal, UGAMNH1795, 1797, 1818, 1830. Remarks—This material has the distinctive sculpted appearance of Pleistocene Trachemys scripta. All the material is well mineralized. It occurs in freshwater ponds, streams, and rivers (Conant 1975). Terrapene Carolina (Linnaeus) Eastern Box Turtle Material—Right and left epiplastron, UGAMNH1703; left hypoplastron, UGAMNH2144; left and right hypoplastron and xiphiplastron, UGAMNH1697, 1698; right hypoplastron at hinge, UGAMNH1686, 1727; left hypoplastron at hinge, UGAMN1685, 1687, 1690, 1705, 1731; right hypoplastron at inguinal notch, UGAMNH1713; right hypoplastron, UGAMNH1714; left and right hypoplastron, UGAMNH1696, 1715; hypoplastron, UGAMNH1716; neural 1, pleural and peripheral 1 and 2, UGAMNH1699; neural 1 and left and right peripheral 1, UGAMNH1732; neural 5 and 6, UGAMNH1730; neural, UGAMNH1707; nuchal, UGAMNH1704, 1725; right peripheral 1 and 44 Timothy S. Young and Joshua Laerm 2, UGAMNH1728; left peripheral 1 and 2, UGAMNH1726; right peripheral 1, 2 and 3, UGAMNH1692; left peripheral 1, 2, 3 and pleural 1, UGAMNH1720; left peripheral 3, UGAMNH1722; left peripheral 3 and 4, UGAMNHA1721; right peripheral 3 and 4, UGAMNH1708; right peripheral 5, UGAMNH1688, 1706; left peripheral 5, UGAMNH1691; right peripheral 6 and 7 and pleural 4 and 5, UGAMNH1710; right peripheral 6, 7, and 8, UGAMNH1712; left peripheral 7, UGAMNH1684; left peripheral 8, UGAMNH1702; left peripheral 8, 9, and 10, UGAMNH1694; right peripheral 9, 10, and 11, UGAMNH1733; left peripheral 10, UGAMNH1689; right peripheral 10 and 11, UGAMNH1695, UGAMNH1718; left peripheral 10 and 11, UGAMNH1719, 1734; right peripheral 10 and 11 and pygal, UGAMNH1711; left peripheral 11, UGAMNH1701; left peripheral 11 and pygal, UGAMNH1709; left and right peripheral 11 and pygal, UGAMNH1717; right peripheral 11, UGAMNH1724; left pleural 2 and peripheral 4 and 5, UGAMNH1700; pygal, UGAMNH1683, UGAMNH1723; left xiphiplastron, UGAMNH1327, 1729; left and right xiphiplastron, UGAMNH1693. Remarks—Terrapene Carolina can be distinguished from its extinct relative T. Carolina putnami based on smaller size. It is a terrestrial woodland species (Conant 1975). Terrapene Carolina putnami Hay Giant Box Turtle Material—Right epiplastron, UGAMNH1860; left hypoplastron at inguinal notch, UGAMNH1855; left hypoplastron and epiplastron and entoplastron, UGAMNH1863; right hypoplastron and xiphiplastron, UGAMNH1864; neural 1 and pleural and peripheral 1, UGAMNH1856; nuchal, UGAMNH1865; left peripheral 3 and 4, UGAMNH1858; right peripheral 6 and 7, UGAMNH1859; right peripheral 10 and 11 and pygal, UGAMNH1862; right peripheral 1, UGAMNH1849; left peripheral 2, 3, and 4, UGAMNH1848; left peripheral 4, 5, and 6, UGAMNH1861; right peripheral 6, UGAMNH185; left peripheral 7 and 8, UGAMNH1852; left peripheral 8 and 9, UGAMNH1853; right peripheral 9, UGAMNH1847; left peripheral 10 and 11, UGAMNH1854; left pleural 2 and 3 and peripheral 4 and 5, UGAMNH1857; right pleural 2 and 3, UGAMNH1850. Remarks—This extinct giant subspecies is common in late Pleistocene deposits of Florida where it occurred in coastal marshes and lowland savannahs. (Auffenberg 1958, Kurten and Anderson 1980). It is readily distinguishable on the basis of its large size. St. Marks River Fauna 45 Family Testudinidae Testudinidae gen. et sp. indet. Material—Osteoderms, UGAMNH1645, UGAMNH1646. Remarks—These specimens represent a large tortoise, but the osteoderms are not diagnostic. Geochelone sp. indet. Material—Pleural, UGAMNH1638; left hypoplastron, UGAMNH1639; right pleural 2, UGAMNH1640; left pleural 4, UGAMNH1641. Remarks—The available material, while certainly Geochelone, could not be referred to a species with confidence. Geochelone incisa (Hay) Material—Right peripheral 7, UGAMNH1642; nuchal UGAMNH1643; right peripheral 5, UGAMNH1644. Remarks—This material compares well with the series of G. incisa in the collections of the Florida Museum of Natural History and corresponds to Auffenberg's (1963) description. The was apparently an open grassland inhabitant thought to require a frost free winter (Kurten and Anderson 1980); however, Martin and Guilday (1967) disagree. Gopherus polyphemus (Daudin) Gopher Tortoise Material—Nuchal, UGAMNH1637. Remarks—This material compares well with modern Gopherus polyphemus which ranges in dry sandy soils (Conant 1975). Family Trionychidae Trionyx sp. indet. Material—Carapacial fragment, UGAMNH1761. Remarks—The available material, while certainly Trionyx because of the distinctive pattern on the bone, could not be referred to a species with confidence. Order Squamata Family Colubridae Colubridae gen. et spec, indet. Material—Vertebrae, UGAMNH2054-2061. 46 Timothy S. Young and Joshua Laerm Elaphe obsoleta (Say) Rat Snake Material—Vertebra, UGAMNH2055. Remarks—This material compares well with modern Elaphe obsoleta which may be found in woodlands and grasslands (Conant 1975). Order Crocodilia Family Alligatoridae Alligator mississippiensis (Daudin) American Alligator Material—Left angular, UGAMNH1015; distal phalanx, UGAMNH1001, right dentary (without teeth), UGAMNH1012; dermal scutes, UGAMNH1003-1011 (1010 and 1011 exhibit crossmends), UGAMNH1023; right femur, UGAMNH1020, 1022; left humerus, UGAMNH1019; fused parietals, UGAMNH1025; left scapula, UGAMNH1014; right scapula, UGAMNH1016, 1018; teeth, UGAMNH1002, 1024, 1026, 1028, 1029; vertebra, UGAMNH1017; frontal, UGAMNH1013; left jugal, UGAMNH1021. Remarks—This material has the distinctive Alligator mississippiensis morphology and it compares well with modern examples. Alligators occur in both fresh and brackish waters (Conant 1975). CLASS AMPHIBIA Order Caudata Family Sirenidae Siren sp. indet. Material—Vertebrae, UGAMNH2129-2131, 2161. Remarks—The available material compares well with modern Siren. Order Anura Anura gen. et sp. indet. Material—Vertebrae, UGAMNH2132-2134-right humerii. Remarks—The available material, while certainly frog, could not be referred to a genus or species with confidence. CLASS OSTEICHTHYES Order Lepisosteiformes Family Lepisosteidae Lepisosteus sp. indet. Material—Scales, UGAMNH2109-21 11. St. Marks River Fauna 47 Remarks—The scales, while certainly Lepisosteus, could not be referred to a species with confidence. Lepisosteus occurs in freshwater and estuarine habitats (Hoese and Moore 1977, Lee et al. 1980). Order Amiiformes Family Amiidae Amia calva Linnaeus Bowfin Material—Left dentary, UGAMNH2088; left frontal, UGAMNH2089; cervical vertebra, UGAMNH2090. Remarks—This material compares well with modern specimens of Amia calva. The bowfin is a freshwater and estuarine species (Hoese and Moore 1977, Lee et al. 1980). Order Siluriformes Family Ictaluridae Ictaluridae gen. et sp. indet. Material—Spine, UGAMNH2112; vertebra, UGAMNH2113. Remarks—The available material, while certainly catfish, could not be referred to a genus or species with confidence. Pylodictis cf. P. olivaris (Rafinesque) Flathead Catfish Material—Left proximal coracoid, UGAMNH2119. Remarks—The morphology of the single element is very similar to modern specimens of P. olivaris and distinct from the other known regional ictalurids available for comparison. The specimen at hand shows some evidence of mineralization, but mineralization is not extensive. The species occurrence in the St. Marks River is outside its reported range which extends from northeastern Mexico east throughout Gulf of Mexico drainages to Mobile Bay (Lee et al. 1980 et seq.). However, in recent times the species has undergone introductions and populations are now known from at least the Appalachicola-Chatahoochee System (M. and B. J. Freeman, University of Georgia, personal communication). Uyeno and Miller (1962) reported some specimens of P. olivaris from the Trinity River Terrace, Texas. The deposit was dated to the Sangamon (late Pleistocene); however, that site is within the present range of the species. It is a freshwater species (Hoese and Moore 1977, Lee et al. 1980). 48 Timothy S. Young and Joshua Laerm Family Ariidae Ariidae gen. et sp. indet. Material—Spine, UGAMNH2114-2116; cervical vertebrae, UGAMNH2117, UGAMNH2118. Remarks—These specimens show the characters of the marine catfishes, although species identification is not possible. Ariusfelis (Linnaeus) Hardhead Catfish Material—Spine, UGAMNH2091. Remarks—This spine compares well with the distinctive Arius felis morphology. This species is restricted to saltwater and estuaries (Hoese and Moore 1977, Lee et al. 1980). Order Salmoniformes Family Esocidae Esox sp. indet. Material—Right dentary, UGAMNH2101, left dentary UGAMNH2095, 2096, 2098-2100, 2102-2105; dentary, UGAMNH2097; parasphenoid, UGAMNH2106; pharyngeal grinding plates, UGAMNH2107, 2108. Remarks—These specimens closely resemble both E. americanus Gmelin and E. niger Lesueur. Both are considered freshwater species (Lee et al. 1980 et seq.) and occur in regional waters today. Order Perciformes Family Percichthyidae Morone saxatilis (Walbaum) Striped Bass Material—Right maxilla, UGAMNH2082; right premaxilla, UGAMNH2083; right quadrate, UGAMNH2084, 2085; left quadrate, UGAMNH2086; atlas, UGAMNH2087. Remarks—This material compares well with modern examples of Morone saxatilis which occurs in both coastal saltwater and estuaries (Hoese and Moore 1977, Lee et al. 1980). Family Sparidae Archosargus probatocephalus (Walbaum) Sheepshead Material—Right dentary, UGAMNH2079; left preoperculum, UGAMNH2080; tooth, UGAMNH2081. Remarks—This material compares well with modern examples of St. Marks River Fauna 49 Archosargus probatocephalus. The sheepshead is a coastal salt-water and estuary species (Hoese and Moore 1977, Lee et al. 1980). Family Sciaenidae Sciaenops ocellatus (Linnaeus) Red Drum Material—Quadrate, UGAMNH2092. Remarks—This material compares well with modern examples of Sciaenops ocellatus. It is a coastal saltwater species, but is also associated with estuaries (Hoese and Moore 1977, Lee et al. 1980). Family Mugilidae Mugil sp. indet. Material—Vertebrae, UGAMNH2093, UGAMNH2094. Remarks—The available material, while certainly Mugil, could not be referred to a species with confidence. Mugil is a coastal saltwater species (Hoese and Moore 1997, Lee et al. 1980). RESULTS AND DISCUSSION Chronology and Environment of Deposition Of several thousand separate skeletal elements recovered from the St. Marks River, 1,162 were referable to specific taxa. Included are 37 species of mammals, 3 birds, 13 reptiles, 2 amphibians, and 9 fish. An additional 23 species of birds were identified from the 1972 collection made by Storrs Olson. Of all species we reported, 14 mammals and 2 reptiles are restricted to the Pleistocene. The remaining are representative of the modern extant regional fauna. With the exception of modern contaminants, the latter are acceptable Pleistocene species; however, they more probably represent a mixture of Holocene and Pleistocene material. This is reflected in the range of mineralization observed in many species. In all cases those species known only from the Pleistocene are well mineralized. However, several species with both a Pleistocene and Recent occurrence such as horse and deer exhibit both well mineralized and, what appears to be, very recent unmineralized condition. Modern contaminants such as cow and pig are unmineralized. In general, mineralization is no criterion of Pleistocene deposition. The problem of apparent heterochronous deposition and separation of Pleistocene and Holocene materials is exacerbated by the apparent rapid mineralization that can occur in reducing environments. Neill (1957) noted that rapid mineralization of organic remains in Florida creates the illusion that Recent material is of older age. Nonetheless, the St. Marks River 50 Timothy S. Young and Joshua Laerm fauna is clearly mixed and reflects heterochronous deposition over time beginning no later than the late Pleistocene (Wisconsinan) and extending through the Recent. We compared the St. Marks River faunal list and a modern regional faunal list of the Apalachicola River system (Means 1976). Of the 344 species listed by Means, 29% of the mammals, 10% of the birds, 19% of the reptiles, 5% of the amphibians, and 2% of the fish are represented in the St. Marks River fauna. This bias toward mammals probably reflects taphonomic factors associated with the larger size of mammalian elements in a fluvial environment. Small, more fragile vertebrates (birds, reptiles, amphibians, and fish) are clearly under-represented in the St. Marks River fauna. This bias is reflected also in the mammalian fauna where chiropteran, insectivoran, and small rodent remains are conspicuously absent. While many of the species recovered from the St. Marks River are eurytopic and provide only limited information regarding the environment of deposition, a number are stenotopic and are considered good environmental indicators. Mammals—The mammalian fauna, in particular, is very useful in assessing the chronology and paleoenvironment of the St. Marks River. The reason for this is two-fold. First, mammals are the most numerous and have the largest component of extinct forms. Second, Florida has an extremely rich and well-documented late Pleistocene as well as modern mammalian fauna upon which comparisons to the St. Marks River fauna can be made. Thirteen (35%) of the mammalian fauna of the St. Marks River is represented by extinct forms. These include Holmsina septentrionalis, Megalonyx jeffersonii, Glossotherium harlani, Canis dims, Smilodon sp., Synaptomys australis, Tapirus, sp., Equus sp., Platygonus compressus, Hemiauchenia macrocephala, Paleolama mirifica, Mammut americanum, and Mammuthus jeffersonii. This closely approximates the relative percentage of extinct mammals from a number of Rancholabrean faunas from elsewhere in Florida (Martin and Webb 1974). The temporal span of the extinct forms ranges from Blancan through Recent. However, they all share a late Wisconsinan chronology. Those species representing extant forms, although individually some exhibit a longer stratigraphic history, also share a late Wisconsinan chronology. With few exceptions, all the extant species are represented in the local fauna today. Comparison of the known and inferred habitat preferences or requirements of the extant and extinct mammalian species suggests the depositional environment was heterogeneous. On one hand there are a number of essentially woodland species: Didelphis, Holmsina, St. Marks River Fauna 51 Megalonyx, Lutra, Mephitis, Urocyon, Ursus, Tapirus, Platygonus, Odocoileus, and Mammut. However, grassland species are well represented also: Glossotherium, Mephitis, Geomys, Equus, Hemiauchenia, Paleolama, Bison, and Mammuthus. From a simple listing it might appear that grassland species are about as common as woodland species. However, when compared by the number of identified specimens per taxon, woodland species are more prevalent. Despite criticism, this method is reliable for a comparison of relative abun-dances of species (Grayson 1984). In addition, a number of species indicate proximity of water: Didelphis, Lutra, Procyon, Ursus, Castor, Neofiber, Ondatra, Synaptomys, and Tapirus are all typically associated with moist, riparian, or standing water habitats. Birds—Storrs Olson's collection from the St. Marks River have never been published. He was kind enough to provide a list of the birds identified and has permitted us to include it in the present discussion. Olson (personal communication) felt that "there was very little of interest among the birds" mainly because the list of avian species recovered from the St. Marks River is essentially similar to the modern fauna (Means 1976). As a whole, birds are uninstructive concerning the dating of the St. Marks River fauna. They do, however, provide considerable information relating to the environment of deposition. The St. Marks River avian fauna is clearly biased toward large species with predominantly salt and freshwater marshland habitat preferences: Podiceps, Podilymbus, Phalacrocorax, Ardea, Butorides, Egretta, Eudocimbus, Aix, Anas, Aythya, Branta, Bucephala, Lophodytes, Padion, Fulica, Gallinula, and Aramus. In addition, a number of the species are typically associated with woodlands or woodland riparian habitats: Aix, Mergus, Buteo, and Strix. Conspicuously absent are the passeriforms. This probably represents the taphonomic bias referred to above. While a significant number of the birds are often present in saltwater marsh habitats, there are no shorebird (charadriform) species present. Reptiles and Amphibians—Many turtles, but few other reptiles, are reported from the St. Marks River. Emydid turtles, in particular, are well represented and make up approximately 90% of the recovered reptilian material. In fact, in numbers alone they make up well over one third the individual elements in the fauna. The emydid turtle species identified from the 1987 collection were Pseudemys concinna, P. floridanus, P. nelsoni, Trachemys scripta, and Terrapene Carolina, all of which are found in the area today. Pseudemys and Trachemys are indicative of a freshwater environment, while Terrapene is terrestrial. An extinct, large, late Pleistocene subspecies of Terrapene 52 Timothy S. Young and Joshua Laerm Carolina, T. c. putnami, is represented in the St. Marks River fauna by a number of elements. It was probably limited to the Coastal Plain and Savannah habitats (Auffenberg 1958) and is represented in many late Pleistocene sites in Florida. Other aquatic turtles recovered include one chelydrid, C. serpentina, and a number of unreferrable kinosternid fragments. Terrestrial testudinoid turtles present at the site are Geochelone incisa, Geochelone sp., and Gopherus polyphemus. Geochelone incisa represents a definite late Pleistocene species, as does Terrapene Carolina putnami. Gopherus polyphemus occurs in the area today. Only two snakes, Nerodia sp. and Elaphe obsoleta, were identified from the 1987 collection. Both snakes occur in the area today. No lizards were identified from any of the fossil collections. Two amphibians were recovered, one caudate and one anuran, neither of which could be identified to species. With the exception of the two late Pleistocene components, the herpetofauna is representative of the modern regional fauna and includes both lower Coastal Plain riverine and marshland species, as well as terrestrial forms. Fishes—The fish fauna described includes both freshwater and marine forms. Ariopsis felis, Morone saxatilis, Archosargus probatocephalus, Sciaenops ocellata, and Mugil sp. although typically marine are also estuarine tolerant. The freshwater fishes include Pylodictis cf. P. olivaris, Lepisosteus sp., Esox sp., and Amia calva. Of these, Lepisosteus sp., Esox sp., and A. calva tolerate estuarine, but not marine, conditions (Hoese and Moore 1977). In conclusion, the aquatic community suggests a mixed freshwater and marine, or more likely an estuarine environment, similar to the lower half of the St. Marks River drainage today. The terrestrial fauna indicates a wooded riparian environment also similar to that found in the St. Marks River drainage today. However, the presence of Hemiauchenia, Bison, Equus sp., and Mammumthus coupled with Geomys, Geochelone, and Gopherus suggests that more open, semi-forested savannah habitats were also represented. This is consistent with other late Pleistocene (Rancholabrean) faunas from the panhandle of Florida, some of which are considered below. Faunal Comparison The Chipola River sites (IA and HA)—This is a river deposit similar to the St. Marks River and contains similar species including Didelphis virginiana, Holmesina septentrionalis, Castor canadensis, Procyon lotor, Bison sp., Equus sp., Mammut americanum, Odocoileus virginianus, and Hemiauchenia macrocephala (Webb St. Marks River Fauna 53 1914a). Although no formal paleontological description of the site exists, the species present in that assemblage indicate a mixed woodland/grassland environment (Webb 1974a). The Aucilla River IA site—The site is also similar to the St. Marks River in depositional and temporal characters. No published paleontological description exists for this site either, but from the fauna a habitat of woodland and marsh can be assumed. It includes Didelphis virginiana, Holmesina septentrionalis, Glossotherium cf. G. harlani, Ondatra zibethicus, Castor canadensis, Neochoerus pinckneyi, Sylvilagus floridanus, Canis dims, and Tremarctos floridanus (Webb 1974a). Wakulla Springs—This, too, is similar to the St. Marks River in depositional and temporal characters. Included are Mammuthus sp., Mammut americanum, and Bison bison antiquus (Webb 1974a). No formal paleontological description of the site exists. Generally there are only slight differences between the St. Marks River and other Florida panhandle, riverine deposits. These differences can probably be attributed to a number of causes including collection by amateurs, undersampling, taphonomic events, or other collecting biases. Compared to the other Rancholabrean faunas from peninsular Florida (Martin and Webb 1974, Webb 1974a, Webb and Wilkins 1984), the St. Marks River assemblage probably is not representative of the full late Pleistocene fauna that existed in the area. For example, more than 50 species of mammals are known to have been present in Florida during the time of accumulation of the Ichetucknee River fauna, Columbia County, Florida (Martin and Webb 1974). As shown by Martin and Webb (1974) mammalian faunal diversity was considerably elevated in peninsular Florida during Rancholabrean time, and it is highly likely that is was the case along the rich fluvio-estuarine environment of the panhandle during the same period. ACKNOWLEDGMENTS—Thanks are due Chris McKensie, Locke Rogers, Brad Newsom, Tim Gaudin, and Luis Insignares for their efforts below and above the surface of the St. Marks River. Chris McKensie deserves added thanks for the many long hours he spent screening and sorting bones. The United States Department of Agriculture provided an advance copy of their publication on the soils of Wakulla County. Gary Morgan and Russ McArty at the Florida Museum of Natural History were more than just helpful. The gracious offer of Storrs Olson to make available his unpublished records of the avian material he collected and analyzed is greatly appreciated. Robert Martin, Robert Frey, and Elizabeth Reitz provided 54 Timothy S. Young and Joshua Laerm many helpful critical comments on earlier drafts. Funds for this study were provided through Department of Zoology and the Museum of Natural History, the University of Georgia. LITERATURE CITED American Ornithologists Union. 1983. Check-list of North American birds. Sixth edition. Allen Press, Lawrence, Kansas. Auffenberg, W. 1958. Fossil turtles of the genus Terrapene in Florida. Bulletin of the Florida State Museum 3:53-92. Auffenberg, W. 1963. Fossil Testudinine turtles of Florida, genera Geochelone and Floridemys. Bulletin of the Florida State Museum 7:54-97. Brodkorb, P. 1963. Catalogue of fossil birds, Part 1 (Archaeopterygiformes through Ardeiformes). Bulletin of the Florida State Museum 7:179- 293. Burt, W. H. 1928. 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Pages 370-383 in Contributions in Quaternary Vertebrate Paleontology: a volume in memorial to John E. Guilday (H. H. Geonways and M. R. Dawson, editors.). Carnegie Museum of Natural History, Pittsburgh, Pennsylvania. Winker, C. D., and J. D. Howard. 1977a. Correlation of technically deformed shorelines of the southern Atlantic Coastal Plains. Geology 5:123-127. Winker, C. D., and J. D. Howard. 1971b. Plio-Pleistocene paleogeography of the Florida Gulf Coast interpreted from relict shorelines. Transactions of the Gulfcoast Association of Geologists Society 27:409- 420. Yon, J. W. 1966. Geology of Jefferson County, Florida. Bulletin of the Florida Geological Survey 48:1-119. Accepted 29 July 1992 58 AUTUMN LAND-BIRD MIGRATION ON THE BARRIER ISLANDS OF NORTHEASTERN NORTH CAROLINA by Paul W. Sykes, Jr. For three consecutive years Sykes investigated the autumn migration of land birds in the Bodie Island and Pea Island area of coastal North Carolina. During a 102-day period in 1965, he recorded 110,482 individual birds of 148 species. He was able to correlate major influxes of migratory species with specific weather patterns. His data show seasonal peaks of southward movement for the land-bird species that pass along the North Carolina coast in large numbers. In addition, Sykes recorded five species native to the western United States. Three of these vagrants provided the first reports of Swainson's Hawk, Sage Thrasher, and Western Meadowlark for North Carolina. 1986 49 pages Softbound ISBN 0-917134-12-5 Price: $5 postpaid. North Carolina residents add 6% sales tax. Please make checks payable in U.S. currency to NCDA Museum Extension Fund. Send order to: LAND-BIRD MIGRATION, N.C. State Museum of Natural Sciences, P.O. Box 27647, Raleigh, NC 27611. No Decline in Salamander (Amphibia: Caudata) Populations: A Twenty-Year Study in the Southern Appalachians Nelson G. Hairston, Sr., and R. Haven Wiley Department of Biology, University ofNorth Carolina Chapel Hill, North Carolina 27599-3280 ABSTRACT—Identical observations, conducted 1-4 times per year for 15-20 years at two locations in the southern Appalachians, have yielded quantitative data on populations of six species of salamanders. Although the numbers have fluctuated for various reasons, there has been no trend in the numbers of any of the species. The "world-wide decline of amphibian populations" has not occurred in the two localities studied. Recently, much attention has been given to a decline in many populations of amphibians (Barringer 1990, Blaustein and Wake 1990, Phillips 1990). There is a suggestion by some authors that there is a general cause for a supposed "world-wide" decline. We do not deny that many amphibian species have decreased in abundance. Among the causes that have been suggested are acid precipitation (Harte and Hoffman 1989, Beebee et al. 1990) and ultraviolet increase due to ozone depletion (Barringer 1990, Blaustein and Wake 1990, Phillips 1990). The same authors have considered overcollecting and rejected it as a general cause. Habitat destruction is also widely mentioned. The last cause is common to all species, except for some pioneering ones, and would not apply only to amphibians. The situation is regard-ed by many herpetologists as very serious, so much so that the World Conservation Union (IUCN), Species Survival Commission, has activated a Declining Amphibian Populations Task Force. This group has estab-lished local subgroups throughout the United States and elsewhere in the Americas to promote research on the problem. If there has been a general cause for the decline in amphibian populations, all amphibian populations should be involved; if they are not, the original claim of a "world-wide decline" must be modified, either by eliminating some taxonomic groups, some ecolog |
OCLC Number-Original | 4904283 |