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A New Species of Hemiauchenia (Camelidae;Lamini) from the Plio-Pleistocene of Florida

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A New Species of Hemiauchenia (Camelidae;Lamini) from the Plio-Pleistocene of Florida
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2008

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A NEW SPECIES OF Hemiauchenia (CAMELIDAE;LAMINI) FROM THE PLIO-PLEISTOCENE OF FLORIDA By JULIE ANNA MEACHEN A THESIS PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE UNIVERSITY OF FLORIDA 2003

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I dedicate my thesis to my step-mother, Laura Oleshko Meachen, who passed away on October 12, 2002. She never got to see the finish ed product, but I am sure she would have been very proud of me. Laura, wherever you are, this is for you.

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ACKNOWLEDGMENTS First and foremost, I would like to thank my major advisor, Dr. S. David Webb. Although a valuable professional mentor, he was also like a caring member of the family. All the good advice he gave me, and all the interesting conversations I had with him I will never forget. I am sincerely glad that I had the pleasure of working with him before he retired this year. I will miss him dearly. I would also like to acknowledge the other members on my committee, Dr. Bruce MacFadden, who has also helped me in many ways and Dr. Richard Kiltie as well. There are many graduate students with whom I have had the pleasure of working over the years. I would like to thank Diana Hallman for being like a big sister in many respects and also being my collaborator on projects and a good friend. I would like to thank Matt Mihlbachler, Brian Beatty and Andy Hemmings for all their advice and help. I extend my thanks to the staff of the vertebrate paleontology collection at the FLMNH, Russ McCarty, who has been a great friend and mentor, Dr. Richard Hulbert, Art Poyer and especially Dr. Pennilyn Higgins for all the advice, proofreading and just being a darn good buddy. My master’s thesis was greatly improved by the FLMNH Lucy Dickinson fellowship in vertebrate paleontology. Last, but certainly not least, I would like to thank all my friends and family. Without their help and support I would never be where I am today! iii

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TABLE OF CONTENTS page ACKNOWLEDGEMENTSiii LIST OF TABLESv LIST OF FIGURES.vi ABSTRACT..viii CHAPTER 1 INTRODUCTION... 1 Family Camelidae1 A New Species 4 2 MATERIALS AND METHODS.............6 3 SYSTEMATIC SPECIES DESCRIPTION...........11 Age and Occurrence...........11 Comparisons from the American Museum of Natural History.. 13 Systematic Paleontology 14 4 PALEOECOLOGY OF THE NEW SPECIES..43 Dentition45 Mesowear...46 Stable Carbon Isotope Analysis.47 Limb Proportions...49 5 CONCLUSIONS............52 LITERATURE CITED......54 BIOGRAPHICAL SKETCH.58 iv

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LIST OF TABLES Table page 1. Specimens of the new species of Hemiauchenia by locality...........................................7 2. Measurements of the new species' dentition in mm......................................................16 3. Measurements of the new species' humeri in mm.........................................................26 4. Measurements of the new species' radio-ulnae in mm..................................................28 5. Measurements of the new species' femora in mm.........................................................32 6. Measurements of the new species' astragali in mm.......................................................35 7. Measurements of the new species' calcanea in mm.......................................................37 8. Measurements of the new species' metapodials in mm.................................................41 9. Measurements of the new species' proximal phalanges in mm.....................................41 10. Concurrent species at the Inglis 1A and DeSoto Shell Pits fossil localities................44 v

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LIST OF FIGURES Figure page 1. A modern phylogeny of the Camelinae.. 3 2. Map of Florida fossil sites where the new species has been found. 12 3. New Hemiauchenia species holotype.. 17 4. Maxillary fragment of new species with deciduous P3 and P4... 21 5. Maxillary fragment of the new species with P4 and M1. 21 6. Lower m3 measurements for five lamine species .. 23 7. Lower right m3 of new species 24 8. Lower left m3 with mandibular fragment of new species.. 24 9. Lower m3s of new Hemiauchenia species and Hemiauchenia edensis .. 25 10. Hemiauchenia macrocephala and Palaeolama mirifica m3s.... 25 11. Distal humeri from new Hemiauchenia species, posterior view 12. Distal humeri of new species, anterior view. 27 13. Comparison of the breadth of the distal end against the depth of the distal end of the humerus . 28 14. Radio-ulnae of new Hemiauchenia species, lateral view.. 29 15. Radio-ulnae of new species, anterior view 29 16. Comparison of lengths and widths of radio-ulnae 31 17. Femora of new Hemiauchenia species, posterior view. 34 18. Comparison of breadth of the proximal end to minimum shaft width of femora ....... 35 vi

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19. Astragali of new Hemiauchenia species, plantar view. 36 20. Astragali of new species, dorsal view.. 36 21. Size comparisons of the astragali of the new Hemiauchenia species to other lamine species and deer, Odocoileus virginianus 37 22. Calcanea of new Hemiauchenia species, medial view.. 39 23. Calcanea of new species, anterior view. 39 24. Metatarsal of new Hemiauchenia species, anterior view.. 40 25. Comparison of metapodial lengths and widths ..... 41 26. Proximal phalanges of new Hemiauchenia species, anterior view 42 27. Proximal phalanges, posterior view 28. Plot of 13 C values versus fraction C4 contained in varied kinds of plant communities, and the corresponding enriched values of tooth enamel 48 29. Stable carbon isotope data. 49 vii

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Abstract of Thesis Presented to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Master of Science A NEW SPECIES OF Hemiauchenia (CAMELIDAE;LAMINI) FROM THE PLIO-PLEISTOCENE OF FLORIDA By Julie Anna Meachen August 2003 Chair: S. David Webb Major Department: Zoology In 1974 paleontologists from the Florida Museum of Natural History (FLMNH) discovered a gracile llama that was originally thought to be Hemiauchenia macrocephala. It was discovered at the Inglis 1A fossil site in Citrus County, Florida. Upon further inspection and comparison of this material with known H. macrocephala material, Florida museum paleontologists decided that this was not H. macrocephala, but a new species altogether. More of this new lamine was also found at the De Soto Shell Pit fossil locality in De Soto County, Florida. Both fossils sites are latest Blancan, earliest Irvingtonian in age. For the description of this new species, I conducted a thorough morphological analysis of the crania and postcrania, which I then compared to the morphology of Hemiauchenia macrocephala, Palaeolama mirifica, “Hemiauchenia” minima, Hemiauchenia edensis and extant lamines, Lama and Vicugna. The holotype for this new species was a right mandibular fragment including p4, m1, m2 and m3 from De Soto viii

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Shell Pit in De Soto County, Florida (UF 210707). This specimen is housed in the FLMNH vertebrate paleontology collection. This new lamine shares many homologous character states with Hemiauchenia macrocephala and Hemiauchenia edensis. However, the new lamine has postcrania that are always more slender than those of H. macrocephala and usually larger than H. edensis. It also has some unique dental features, such as the lack of a p3. This new species is placed in the genus Hemiauchenia based on several morphological characters: The length and gracility of the postcranial skeleton, the shape and form of the teeth and the striking resemblance that this new species shares with H. macrocephala and H. edensis. From dental characters, post-cranial analysis and stable carbon isotope analysis it seems that this new species ate mainly browse with occasional grass mixed in, and preferred to dwell in savanna type environments. It was probably an agile cursor, using that ability as its main mode of predator evasion. It coexisted with Hemiauchenia macrocephala and other artiodactyls, but not with fellow lamine Palaeolama mirifica. This new species may have had an affiliation with the origin of the true (extant) llamas that crossed the Isthmus of Panama in the early Pleistocene, as did Palaeolama. ix

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CHAPTER 1 INTRODUCTION Family Camelidae The family Camelidae includes the living camels, llamas, vicuas, guanacos and alpacas. They originated in North America in the middle Eocene (approximately 54 MA) and remained endemic to North America until the late Miocene, when they dispersed to Asia and Africa and gave rise to the modern camels (Moya-Sola and Agusti 1989; Pickford et al. 1993; Pickford et al. 1995). In the early Pleistocene, they dispersed to South America to give rise to the modern lamines (Webb 1974). Camelids became extinct in North America in the late Pleistocene, along with most of the other endemic megafauna (Honey et al. 1998). There were four major episodes of camelid diversification in North America. The first episode took place in the late Eocene through the middle Oligocene and consisted of the camelids Poebrotherium and Paratylopus. The second radiation took place from the late Oligocene through the early Miocene with the radiation of the stenomylines (Blickomylus, Rakomylus). These species became extinct in the middle Miocene. The third radiation of camelids took place in the same time span as the second radiation, and consisted of “higher camelids” or those that possessed metastylids on the lower molars. This radiation more than doubled the number of camelid genera. The fourth radiation of camelids occurred in the early to middle Miocene and produced the Camelinae (Lamini and Camelini) including all extinct and modern American lamines (Honey et al. 1998). 1

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2 The first lamine fossils, represented by the genus Alforjas or Pleiolama (Webb and Meachen in press) originated in the Great Plains of North America approximately 11 million years ago. Pleiolama is thought to have given rise to the genus Hemiauchenia approximately ten million years ago. Hemiauchenia is thought to have engendered the extinct genus Palaeolama, and also the modern South American lamines. This transition is estimated to have taken place two million years ago (Wheeler 1995) (See Figure 1 for current lamine phylogeny). The genus Hemiauchenia (syn. Tanupolama) was named by H. Gervais and Ameghino in 1880. This genus is in the Class Mammalia (Linnaeus 1758), Order Artiodactyla (Owen 1848), Suborder Tylopoda (Illiger 1811), Family Camelidae (Gray 1821), Subfamily Camelinae (Gray 1821) and Tribe Lamini (Webb 1965). Cranial characteristics of this genus include a dental formula of I1/3, C1/1, P3/2-3, M3/3, a markedly low hypsodonty index (tooth crown height/tooth width or length), with only the genus Palaeolama having a lower hypsodonty index, laterally compressed and recurved canines, a sharp diastemal crest on the mandible, a swollen, narrow rostrum, and arched and retracted nasals, but not as far as in the genus Lama. The palatine notch is sharply V-shaped extending to the M2. Postcranial characters include, very long, slender limbs and cervical vertebrae (Harrison 1979), fused metapodials that are longer than the basal length of the skull, and a proximal phalanx with a W-shaped suspensory ligament scar (Honey et al. 1998). Most Hemiauchenia species were comparatively small camelids, barely larger than the domestic llama (Dalquest 1992), but some became larger than the modern camels.

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3 The four species currently attributed to this genus are: Hemiauchenia vera (Matthew) from Florida, Texas, the Texas/Oklahoma panhandle, Kansas, Nebraska, Arizona, New Mexico, California, and Central Mexico; Hemiauchenia blancoensis (Meade) from Florida, Texas, the Texas/Oklahoma panhandle, Kansas, Nebraska, Arizona, New Mexico, Colorado, and Washington state; Hemiauchenia macrocephala (Cope) is known from Florida, Texas, California, and South America; and “Hemiauchenia” minima from Florida (Honey et al. 1998). Webb, MacFadden and Baskin (1981), however, suggested that a new genus name might be more appropriate for “Hemiauchenia” minima. Figure 1: A modern phylogeny of the Camelinae, adapted from Honey et al. 1998. Numbered slash marks indicate synapomorphies of the clade above. 1.) LAMINI: arched nasals. 2.) I1-2 lost; P2/p2 lost; anteroexternal style present on lower molars. 3.) Small P1/p1; p3 small or absent; reduced lacrimal vacuity; shortened rostrum. 4.) HEMIAUCHENIA: extremely elongate metapodials, neck.

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4 Hemiauchenia vera can be distinguished from other members of the Hemiauchenia genus by relatively brachydont (low-crowned) teeth, a large caniniform upper P1, and the retention of the lower p3. This species was found in the Hemphillian North American Land Mammal Age (Dalquest 1992). Hemiauchenia blancoensis is characterized by a shorter mandibular diastema than in H. macrocephala, but longer than H. vera; Caniniform upper P1, absent upper P2, upper P3 present or absent, and lower crowned molars than H. macrocephala (Breyer 1977). Hemiauchenia blancoensis is known from the late Blancan (2.5 to 2.0 MA)(Morgan and Hulbert 1995). Hemiauchenia macrocephala is the Hemiauchenia species that has been found at the widest range of fossil sites in Florida, and throughout North America. It occurs at localities that range in age from the late Blancan through the late Rancholabrean, 2.5 MA to 10 KA; (Morgan and Hulbert 1995; Webb and Stehli 1995). H. macrocephala is characterized by long, robust limbs, large skeletal size, presence of a deciduous upper P2, a fully molariform deciduous upper P3, hypsodont (high-crowned) molars (hypsodonty index of 1.4 to 2.2), a thick layer of cementum on the teeth and a broad mandibular symphysis with incisors oriented in a vertical fashion, reminiscent of the lower incisors of a horse (Hallman and Meachen in review). It overlaps in time span and locality ranges in Florida with both H. blancoensis, in the late Blancan and a new species of lamine in the late Blancan to early Irvingtonian (Morgan and Hulbert 1995; Webb and Stehli 1995). A New Species In 1974 S. David Webb and other paleontologists from the Florida Museum of Natural History (FLMNH), then the Florida State Museum, discovered a gracile llama

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5 that was originally thought to be Hemiauchenia macrocephala. It was discovered at the Inglis 1A fossil site in Citrus County, Florida (Webb 1974). Upon further inspection and comparison of this material with known H. macrocephala material stored in the UF vertebrate paleontology collection, Florida museum paleontologists decided that this was not H. macrocephala, but a new species altogether. In the 1980s and 1990s, volunteers Steve and Suzan Hutchens and Reed and Barbara Toomey discovered more of this new llama material from the De Soto Shell Pit fossil locality in De Soto County, Florida, which they donated to the FLMNH vertebrate paleontology collection (Ruez 2001). The similarity of the fossils from these two sites as well as the same approximate age of the fossil localities, indicate the existence of a new llama species in the Plio-Pleistocene of Florida.

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CHAPTER 2 MATERIALS AND METHODS For the description of this new species, I conducted a thorough morphological analysis of the crania and postcrania, which I then compared to the morphology of Hemiauchenia macrocephala, Palaeolama mirifica, “Hemiauchenia” minima, and extant lamines, Lama and Vicugna. I also compared it to a lesser-known lamine species, Hemiauchenia edensis (Webb et al. in press). These comparisons formed the basis for two different series of interpretations, one systematic and the other ecological. The first purpose was to determine how this sample differed from other lamine samples and how its characteristics placed it in a phylogeny with other lamine clades. The second purpose of this comparison was to better understand the relationship between form and function in this and other lamines. The Anatomy of the Dromedary (Smuts and Bezuidenhout 1987) provided an authoritative basis for many of the descriptive features used in the comparisons. All structures in this new lamine were recognized as homologous structures in the dromedary camel. A Guide to the Measurement of Animal Bones from Archaeological Sites (Von den Dreisch 1976) was used to guide the postcranial measurements taken for this study. This study did not concentrate only on the cranial material of this new species because of the small amount of cranial and dental material recovered from the two fossil localities in which this species is found. A large quantity of postcranial material was also incorporated to describe this new animal (Table 1). 6

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7 All fossil specimens in this study are currently housed in the Florida Museum of Natural History Vertebrate Paleontology collection unless otherwise specified. All extant specimens are housed in the Florida Museum of Natural History Environmental Archaeology collections. The Florida Museum of Natural History specimens are designated with the prefix UF. Cranial, dental, tarsal, carpal, and phalanx measurements were taken with digital calipers to the nearest 0.1 mm, as were the measurements of the widths and ends of the long bones. All other measurements were taken with manual sliding calipers to the nearest millimeter. The abbreviation “M.” denotes Musculus, for a particular muscle, followed by the rest of the anatomical Latin name, for example: M. anconeus. Lower-case letters indicate lower teeth and upper-case letters indicate upper teeth. To supplement the Florida fossil material used in this study, material from the American Museum of Natural History’s Frick collection (abbreviated F:AM) was also utilized. Table 1: Specimens of the new species of Hemiauchenia by locality. Skeletal Element Inglis 1A (Citrus County) De Soto Shell Pit (De Soto County) Cranial and Dental Material UF 45493 maxillary fragment with M1 and M2 UF 210707 mandibular fragment with p4-m3, right. UF 210714 maxillary fragment with P4 and M1, right. UF 210715 mandibular fragment with m3, left. UF 210716 maxillary fragment with DP3 and DP4, left. UF 210717 lower m3, right.

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8 Humeri UF 176915 distal end of humerus. UF 210702 distal end of humerus, right. Radio-ulnae UF 8917 complete radio-ulna. UF 179636 distal end of radio-ulna. UF 210701 reconstructed radio-ulna, both ends intact, but no visible contact, left. Femora UF 176925 partial distal femur. UF 45275 and 45276 proximal femora including majority of shaft, but no distal ends. Astragali UF 210722 (Inglis 1F) complete astragalus, right. UF 10281 complete astragalus. UF 210706 complete astragalus, left. Calcanea UF 210709, left and 210710, right. Metapodials UF 176935 complete metatarsal. UF 18230 unfused metacarpal. UF 18236 distal end of metapodial. UF 210720 proximal end of metacarpal, right. UF 210711 proximal end of metacarpal, left. UF 210726 proximal metacarpal fragment, right. Phalanges UF 179639, 18237, and 179638 three proximal phalanges. UF 97203 (Inglis 1B?) proximal phalanx. UF 177024 distal phalanx. UF 210704, 210708, 210712, and 210703 four complete proximal phalanges. UF 210705 medial phalanx.

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9 I performed stable carbon isotope analysis on the tooth enamel of this new species to better estimate feeding strategy. I then incorporated previous studies of artiodactyl postcrania to assess habitat usage. For stable carbon isotope analysis, I used the following procedure. Tooth enamel from the P 4 or M 2 was sampled for carbon isotopic composition following Feranec and MacFadden (2000). The teeth were prepared by cleaning the enamel. This entailed removal of the outermost layer of enamel and any other material that adhered to the enamel surface. An enamel layer was then removed with a carbide dental drill bit in a Foredom Drill at low RPM to minimize vibration. Isotope samples were removed under a binocular stereomicroscope by manually passing the drill bit along the length of the buccal crown until approximately 10 mg of powdered enamel was obtained. To each tooth enamel sample, 1 ml of 30% H 2 O 2 was added to remove organic compounds. The samples were sealed and agitated until the enamel dissolved. The samples were left in the H 2 O 2 solution overnight. Samples were then centrifuged at 10,000 RPM for five minutes. Following centrifugation, the solvent was discarded and the samples rinsed with deionized water. The rinse entailed addition of 1 ml of deionized water, agitation, centrifugation for five minutes at 10,000 RPM, and finally removal of the solvent. The samples were rinsed three times. After the final rinse, 1 ml of 0.1 N acetic acid was added to each sample to remove carbonates. Samples were left overnight. After treatment with acetic acid, the samples were centrifuged and the acetic acid discarded. The samples were given three deionized water rinses and a final rinse with ethanol. Vials were left open overnight in order for the samples to dry.

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10 For isotope analysis samples were shipped to the University of Michigan and analyzed in the mass spectrometer lab at Ann Arbor. The following procedure was followed: 20 g of sample was placed within a capped vial with a septum. Samples were placed in an autosampler that analyzes carbon isotope composition with the addition of phosphoric acid. Isotope concentrations were measured on a Finnigan mass spectrometer with measurements listed in -notation relative to a standard (VPDB). A correction of .10 was applied to the carbon results based on carbon isotope values of an internal UF standard (MEme, an Elephas molar) relative to VPDB (Hallman and Meachen in review).

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CHAPTER 3 SYSTEMATIC SPECIES DESCRIPTION Age and Occurrence The two sites in which this new species occurs are Inglis 1A and the DeSoto Shell Pits, both on the west coast of south-central Florida (Figure 2). The Inglis 1A site is a sinkhole deposit with excellent fossilization. The site is approximately five meters below present sea level, but lacks any marine fossils, therefore it would seem that this deposition occurred at a time of low sea level (Morgan and Hulbert 1995). Larger animals may have fallen or climbed into the sinkhole and starved to death. The faunal assemblage indicates a more arid and open habitat than is present today in Citrus County. Large grazers dominated the savanna habitat of Inglis 1A. This fauna includes both the new Florida llama species and Hemiauchenia macrocephala. The nearby Inglis 1C site had a wetter, forested habitat with large numbers of browsers including the lamine species, Hemiauchenia macrocephala and Palaeolama mirifica with no evidence of the new llama species (Emslie 1998; Ruez 2001). Inglis 1A was an important site because of its unique biochronologic and geographic position. The time that the site was deposited coincides with the acme of the Great American faunal interchange as well as a surge in native faunal diversity, and it lies along the broad subtropical Gulf Coastal Corridor. As a result, this site contains many fossils that give us clues into the Plio-Pleistocene faunal history of Florida as well as the faunal exchange with South America (Ruez 2001; Webb 1976). The presence of Blancan 11

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12 fauna and the absence of Mammuthus at Inglis 1A place it in the latest Blancan mammal age with a date of approximately 2.0-1.6 MA (Bell et al. in press; Morgan and Hulbert 1995). igure 2: Map of Florida fossil sites where the new lamine species has been found. F

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13 T he DeSoto local fauna is the designation for the vertebrate fauna from the organicial o useum of Natural History One specimen ide. les and astragali from the Panaca fossil site in Nevada showed similarller cations along the southern/southwestern U.S. border in the Frick collection that were slightly smaller in layers in the Caloosahatchee Formation, which is exposed in three commercshell mines in DeSoto County, Florida (Morgan and Hulbert 1995). The DeSoto site shares the late Blancan and early Irvingtonian type of fauna that Inglis 1A has and alslacks Mammuthus. However, at Inglis 1A the deposition occurred below present-day sealevel, possibly during a glacial event, whereas DeSoto was deposited five to ten meters above present sea level in an interglacial period. Corals that were collected from the Caloosahatchee Formation gave Helium/Uranium dates of approximately 1.89 to 1.73MA (Bender 1973; Morgan and Hulbert 1995). Comparisons from the American M important question is whether the new species from Florida is also represented in other undescribed samples of Plio-Pleistocene lamines. Only onewas found in the Frick American Museum collection that could possibly belong to the new species, an uncataloged astragalus from the Collins, Texas area, Blancan in age. Approximately the same size as the new species, it was 39.1 mm long and 24.7 mm w Recently a single proximal phalanx was found at the 111 Ranch locality in Arizona that appears to be the new species. This fossil locality is Blancan in age at approximately 2 MA. Several mandib ity in the teeth, mandibles and metapodials. These specimens were slightly smathan the new species, and they also differed by being Hemphillian in age. There were several of these Hemphillian specimens from different lo

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14 size than the new species. Still other specimens of similar character are also found in Florida at the Bone Valley fossil site. They are all Hemphillian in age (approx. 5-7 MA) and seem to represent a small lamine originally named Procamelus edensis (Frick 192from the Mt. Eden fauna of southern California. Frick describes it as having lost the lower p2 and having a reduced lower p3. He describes the tooth crowns as fairly short and broad, and mentions the presence of the “camelid buttresses” (Frick 1921), a diagnostic dental character of lamines. According to Webb et al. (in press) this lamine ireferable to Hemiauchenia, and not to Procamelus. They describe the lower molar“transversely compressed with angular lingual crescents, strong ‘llama buttresses’ and substantial midlingual stylids.” They also mention that the enamel is mildly crenulated and lacks cementum (Webb et al. in press). This species from the Panaca site is also mentioned in more detail in Macdonald and Pelletier (1965). Systematic Paleontology Class Mammalia, Linnaeus 1758 1) s s as Order Artiodactyla, Owen 1848 Suborder Tylopoda, Illiger 1811 Family Camelidae, Gray 1821 Genus Hemhino 1880 Holotype: UF 21070ding p4, m1, m2 and m3 from De Soto Sheia macrocephala and a lesser-known species Hemiauchenia edensis. Subfamily Camelinae, Gray 1821 Tribe Lamini, Webb 1965 iauchenia, H. Gervais and Ameg 7, Right mandibular fragment inclu ll Pit in De Soto County, Florida. Referred material: See Table 1 for a complete listing of all dental and postcranial specimens. Diagnosis: This new lamine shares many homologous character states with Hemiauchen

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15 HowevH. rter. f nus the ost n r the new species is an incomplete right mandibis a er, the new lamine has postcrania that are always more slender than those of macrocephala. In some cases the postcrania are more elongate, in some cases, shoThe postcrania of the new species are always longer and usually more robust than that oH. edensis. It also has some unique dental features, such as the lack of a p3. This new species is placed in the genus Hemiauchenia based on several morphological characters: The length and gracility of the postcranial skeleton, the shape and form of the teeth andthe striking resemblance that this new species shares with H. macrocephala and H. edensis. Evidence for this generic assignment will be given in the description below. Only one speculative character of this new species is troubling with regard to the geassignment, the possible lack of a p1. According to Honey et al. (1998), a character ofgenus Hemiauchenia is that it possesses a p1. Since the evidence for the loss of this character in the new species is speculative at best, it will not be given much weight in this analysis. Also, it is possible, if this new species does truly lack a p1, that it still was mclosely related to a Hemiauchenia species and should be assigned to that genus, despite the currently accepted genus description. Dentition: (Table 2). No incisors, canines, first or second premolars have yet beefound for the new species. The holotype fo le with p4-m3 (Figure 3). The length of the diastema in the holotype is 39.2 mm. This specimen lacks a p3, and has no remnant of one. The lack or reduction of a p3 diagnostic feature of the genera Lama and Vicugna and a variable feature for the genera Hemiauchenia and Palaeolama (Honey et al. 1998). This feature is interesting because the lack of a p3 is a late Blancan character. The Hemiauchenia and Palaeolama individuals from the Irvingtonian seem to posses the p3 more often than not.

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16 Table 2: Measurements of the new species' dentition in mm. Catalog number Tooth orientation Length (anterior-Width (lingually-y) Crown height posterior) labiall U F 210714 P4 13.8 10.9 14.5 UF 45493 M1 17.8 12.7 7.1 M1 21.8 14.3 21.5 16.2 UF 210716 DP4 22.6 15.4 12.8 UF 45493 M2 22.2 15.7 19.5 UF 210707 m1 13.9 9.9 Too w orn UF 210707 m2 19.9 12.1 8.4 UF 210707 m3 29.4 11.7 10.9 UF 210715 m3 26.5 11.8 8.9 UF 210717 m3 28.2 11.1 27.2 UF 210716 11.4 UF 210714 DP3 11.0

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17 A B Figure 3: New Hemiauchenia species holotype, right mandible with p4-m3, UF 210707 from DeSoto Shell Pits, A. labial view, B. occlusal view.

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18 Breyer’s (1977) study on the interspecific jaw variation in Hemiauchenia blancoensis from the late Blancan Broadwater fossil locality in Morrill County, Nebraska, found that fewer than one in ten adults had a lower p3, or evidence of one. Juveniles possessed the Dp3 in most cases and after it was shed, the alveolus closed up leaving no trace that a Dp3 was ever present. Webb and Stehli found, in their 1995 study on Hemiauchenia macrocephala and Palaeolama mirifica, from the Irvingtonian aged Leisey Shell Pits in Hillsborough County, Florida, that over half of the adult individuals possessed p3s. The Leisey Shell Pit fauna is slightly younger than the Broadwater fauna, Inglis 1A fauna or DeSoto Shell Pit fauna (Webb and Stehli 1995). The age of the Leisey fauna is early Irvingtonian, approximately 1.4-1.7 MA, from paleomagnetic dating of shells located within the vertebrate fossil layer (MacFadden 1995). Therefore, it seems likely that the new species possessed a deciduous p3, but when it was shed the alveolus closed, leaving no trace it was ever present, much like the Blancan species Hemiauchenia blancoensis. This presumption can be tested when juvenile mandibles of this new species are recovered. This new species is distinguishable from H. edensis in this regard, because H. edensis seems to always possess a small p3 with two nearly fused roots (Webb at al. in press). Although it is difficult to discern with certainty, it appears that the new species also lacked a p1. On the holotype, the mandible is broken in half at the point where the two halves of the mandibular symphysis meet. Therefore, it can be determined whether this animal had an alveolus for the p1 where the two bones of the symphysis met. The new species evidently lacks this alveolus, arguing for an absent p1. Absence of the p1 is a character of the genera Palaeolama, Lama and Vicugna. In all previous studies

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19 Hemiauchenia has been thought to possess the p1 (Honey et al. 1998). In this respect the new species differs from all other known Hemiauchenia species. However, larger samples may show that this feature of the type specimen is variable. At most it is a minor loss and it should not hold much weight in the decision about where in the lamine phylogeny to place this animal. The mandible of this new species is deeper and more robust than the mandible of H. edensis. H. edensis has a very shallow mandible with an average width at the posterior end of the m3 of approximately 34 mm. This measurement is compared with 43 mm in the new species. The dental arcade is also comparably shorter in H. edensis, with an average length of approximately 72 mm, including the p3, compared to 75 mm in the new species without a p3. The deciduous teeth, DP3 and DP4 are very molariform (Figure 4), and upon first examination, were thought to be M1 and M2. The DP3 presents the trilobate quality of artiodactyl deciduous premolars, but not very strongly. It almost appears to be a deformation of an M1. The DP4 is bilobate (molariform) and resembles an M2. However, these teeth do show the diagnostic root splay of deciduous premolars. The P4 has a distinctly rounded and laterally flattened shape with an open “U-shaped” hollow crescent (Figure 5). In my observations, this “U-shaped” crescent is diagnostic of the genus Hemiauchenia. The genus Palaeolama has sharply pointed lophs with “V-shaped” crescents. The P4 appears to have little wear, and is in excellent condition. The P4 in H. edensis is considerably smaller with a maximum length of 11.6 mm and a maximum width of 9.3 mm. The P4 of the new species retains a rounded

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20 appearance from labial to lingual sides, whereas the P4 of H. edensis tapers toward the lingual side, forming a V-shaped premolar. The molars of the new species are less robust in appearance than the molars of either H. macrocephala or Palaeolama. The enamel of the new species is thin, but the molars are coated in a complete layer of cementum in all specimens, much like the molars of H. macrocephala. By contrast, Palaeolama and H. edensis only have a thin patchy coating of cementum. The teeth of the new species show little crenulation. This small degree of crenulation is also indicative of Hemiauchenia macrocephala. The teeth of Palaeolama show comparatively more crenulation than is present in Hemiauchenia. Crenulation of the teeth is generally a browsing adaptation (Webb and Stehli 1995). In the new species the M1 is shorter and smaller than that in H. macrocephala, as is the m3 (Figure 6). However, it maintains the same shape. Again the “U” shape of the hollow crescents is conserved. The shape of the tooth crown, although less hypsodont, also maintains the same pattern. The ribs of the molars (or the parastyle, mesostyle, and metastyle) are close in height to the cusps of the crescents as in H. macrocephala. I will discuss the functional significance of this concept in the next section. In Palaeolama, by contrast, the ribs appear to be considerably shorter than the cusps. In the new species, the anterior and posterior labial crescents are raised, but not sharply so, and the anterior and posterior lingual crescents are close in height to the former two structures. This contrasts strikingly with the condition in Palaeolama in which the anterior and posterior labial crescents are sharply pointed and form high peaks on the upper molars and there is a considerable size discrepancy in the heights of the lingual and labial crescents.

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21 Figure 4: Maxillary fragment of new species with deciduous P3 and P4, UF 210716, occlusal view. B A Figure 5: Maxillary fragment of the new species with P4 and M1, UF 210714, A. labial view, B. occlusal view.

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22 T he M2 of the new species has a very similar appearance to the M1, except that it is slightly larger and more robust. The lower dentition of the new species is much more transversely compressed than that of either H. macrocephala or Palaeolama, but less so than that of H. edensis. On the lower m3 (Figures 7 and 8), the anterior enamel folds on the molars, referred to as “llama buttresses” are very prominent in the new species, which helps it sustain a great deal of interdental wear in an anterio-posterior direction. In H. edensis the m3 is not only smaller, but the "llama buttresses" are also less robust and have a posteriorly curved orientation, as opposed to the vertically oriented buttresses of the new species (Figure 9). The crown height in the unworn m3 suggests that the new species may have incorporated some grass into its diet, but not as much as H. macrocephala (Figure 10). The posterolophid on the m3 is rather robust in this species, and has a completely longitudinal orientation when compared with other lamine teeth, which have a more transversely curved orientation.

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23 152025303540152025303540Unworn length in mm.Unworn crown height in mm. New Species H. macrocephala Palaeolama H. edensis L. glama 152025303540579111315Unworn width in mm.Unworn crown height in mm. New species H. macrocephala Palaeolama H. edensis L. glama Figure 6: Lower m3 measurements for five lamine species in mm, plotting unworn crown heights against unworn anterio-posterior lengths and unworn widths lingually-labially.

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24 Figure 7: Lower right m3 of new species, UF 210717, A. labial view, B. occlusal view B A Figure 8: Lower left m3 with mandibular fragment of new species, UF 210715, A. labial view, B. occlusal view

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25 B A Figure 9: Lower m3s of A. new Hemiauchenia species (right) and B. Hemiauchenia edensis (left) for comparison. B A Figure 10: A. Hemiauchenia macrocephala and B. Palaeolama mirifica m3s for comparison, labial view. Note the cementum layer and crown height on the H. macrocephala tooth, and the cusp reliefs (mesostyle to cusp) and cusp tip shape on both, Palaeolama cusps are slightly broken, dotted lines indicated original relief.

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26 Postcranial: There was a great deal of material found representing the postcranial skeleton of the new species. The long, slender nature of the limb bones is the most striking feature of this new species. Only two distal ends of the humerus were recovered for the new species (Table 3 and Figures 11 and 12). The size of the humerus differentiates it from both H. macrocephala and Palaeolama (Figure 13). These distal humeri were approximately one-half the depth and two-thirds the width of the latter two species. Table 3: Measurements of the new species' humeri in mm. Catalog number Breadth of distal end (BD) UF 176915 36.8 UF 210702 40.9 The dimensions of the humeri of the new species were almost identical to those of extant lamines and H. edensis. The humeri of the new species were slightly more robust then either of the other species, especially in the olecranon fossa, the fossa radialis, and the thickness of the humeral shaft. There also appears to be more surface area for the attachment of the M. flexor carpi ulnaris, M. flexor digitorum profundus, M. extensor digitorum lateralis, and M. extensor digitorum communis. With regards to H. edensis, the epicondylus lateralis and the epicondylus medialis were both more flattened and robust in the new species. The humeri of the new species were much less robust than those of either H. macrocephala or Palaeolama. The radio-ulna is the most complete postcranial element for the new species, and also seems to be the most diagnostic postcranial element for this species (Table 4 and Figures 14 and 15).

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27 A B Figure 11: D istal humeri from new Hemiauchpecies, A. UF 210702, B. UF 176915, posterior view Figure 12: Distal humeri of the new species, A. UF 210702, B. UF 176915, anterior view. enia s A B

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28 30354045505560657075203040506070Depth of the distal end in mm.Breadth of the distal end in mm. New Species H. macrocephala Palaeolama H. edensis L. glama/L. guanaco Figure 13: Comparison of the breadth of the distal end against the depth of the distal end of the humerus in several species of lamines. Only distal ends were compared Table 4: Measurements of the new species'dio-ulnae in mm. Catalog number Greatest Length (GL) Length of Olecranon (LO) Smallest depth of olecranon (SDO) Depth across the Processus Anconaeus (DPA) Smallest diameter of the diaphysis (SD) Depth of distal end (DD) Breadth of distal end (BD) because that was the only available complete measurement for the new species. ra UF 210701 No contact 64.5 38.5 48.4 27.4 38.9 44.8 UF 8917 472 64.5 40.3 50.7 31.5 42.2 49.8 UF 179636 N/A N/A N/A N/A N/A 35.9 41.4

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29 Figure 14: Radio-ulnae of new Hemiauchenia species, A. UF 210701, B. UF 8917, lateral Figure 14: Radio-ulnae of new Hemiauchenia species, A. UF 210701, B. UF 8917, lateral view. Figure 15: Radio-ulnae of the new species, A. UF 210701, B. UF 8917, anterior view. A B A B A 29 view. Figure 15: Radio-ulnae of the new species, A. UF 210701, B. UF 8917, anterior view. A B

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30 The most striking feature of the radio-ulna of the new species is the length. It is about 1.2 times longer than the radio-ulnae of H. macrocephala and 1.3 times longer than that of Palaeolama (Figure 16). Out of the two radio-ulna specimens that were available, only one has a usable length. One has been restored, and there is no visible point of contact to validate the length as accurate. However, the overall shape of the specimens are identical and there is no doubt that they belong to the same species. The radio-ulna is long and slender with small proximal ends and a short olecranon process with a tapered appearance. The trochlear incisure of the radio-ulna where it articulates with the humerus is more tightly curved and neatly rounded in the new species than in any other lamine specimen avaliable for analysis. This feature distinguished the dio-ulna of this species from other possible species matches in the Frick collection at the American Museum of Natural History. In Hemiauchenia the shaft of the radio-ulna remains thin and functionally uniform all the way down to the styloideus process where the radio-ulnae has an abrupt thickening to facilitate articulation with the carpals. The new species displays this pattern. In Palaeolama, the distal end of the radio-ulnar shaft gradually thickens toward the medial styloid process, and produces a very robust appearance in the distal half of the radio-ulna. The radio-ulna of the new species can be distinguished from H. edensis by size. There were no whole bone specimens of H. edensis to compare lengths. However, the smallest width of the radio-ulnar shaft in H. edensis was 22 mm, and the breadth of the istal end was only 32.4 mm (compare to the new species measurements in Table 4). ra d

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31 . vera and H. blancoensis specimens are from the F:AM. No complete radioea for attachment of the M. triceps brachii and M. area ulna to the radius at the distal portion of the bone. In H. macrocephala and e earlier species of Hemiauchenia, such as H. vera and H. blancoensis. This complete fusion is present to a degree in H. edensis where a pin-sized aperture is present. The Figure 16: Comparison of lengths and widths of radio-ulnae in six species of lamines. Hulna specimens were available from H. edensis. In the new species there is limited ar 10203040506070Width in mm. H. blancoensis 200 Palaeolama L. glama 300400500600Length in mm. New Species H. macrocephala H. vera anconeus. In both H. macrocephala and Palaeolama there is a large surfacefor the attachment of both these muscles. Since these muscles extend the elbow joint andflex the shoulder joint, this reduction would have given the new species a very gracile appearance in the leg and shoulder and may have compromised power for agility. One very interesting character of the radio-ulna of the new species is the complete fusion of the Palaeolama as well as extant lamines, there is a small aperture where the ulna has not completely fused to the radius. Neither of the two whole bone specimens of the new species show this aperture and a portion of the distal end of a third radio-ulna does not show it either. This complete fusion of the radius and ulna appears to be present in som

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32 purpose of this complete fusion could possibly be an adaptation to cursory. The reduction of apertures in the bone could have strengthened the radio-ulna, making it more amenable to the impacts of cursorial activity. The radio-ulna from the De Soto site appears to articulate with the humerus from that same site. The femora of the new species bear a striking resemblance to those of extant lamines (Table 5 and Figure 17). The diameter of the femoral shaft is only slightly greater than that of Lama (Figure 18). The distal and proximal ends are also highly comparable in size and shape. The length of the shaft of the new species is unknown because there are no complete femora available. A good estimate of length can be made from the new specieTable 5 end (BP) circumference of the the diaphysis (SD) proportions of the other elements of the skeleton and it is thought that the femora of the s are considerably longer than those of Lama. : Measurements of the new species' femora in mm. Catalog number Breadth of proximal Smallest diaphysis (CD) Smallest diameter of UF 45276 66.8 30.3 23.2 UF 45275 67.0 28.8 23.8 The femur presents a very rugose lateral supracondylaris tuberosity, which M. adductor. The trochanter major in the De Soto specimen is quite rugose, suggesting the M. vastus lateralisH. macrocephalaPalaeolamaM. vastus lateralis scars than does the new species. The fovea capitus is only a small notch la. In the extant lamines, the fovea capitus extends from the middle of the caput femoris to appears to facilitate a large M. flexor digitorum superficialis and to some extent a large strong attachments for the M. gluteus accessorius, M. gluteus profundus and possibly. However, both and show stronger in the middle of the caput femoris, this is a shared character state with H. macrocepha

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33 the suture of the epiphysis. Palaeolama shares this modern condition. The trochanter minor is well worn in both femur specimens, however it appears to have been sharply pointed in life. Although the new species has a number of pronounced rugosities whmuscle attachments were present, the femora of bo ere th H. macrocephala and Palaeolama are mor the astragali of the new deep notch, on the plantar es wic. This notch lamines, but not deer. This notch is possibly aism in lamines, and most camelids, tcilitates standingt for extended periods of time. The flange on the medial surface of the astragalus is greatly reduced in the new species, allowing the navicular bone to reach the deep notch. The large concave facet located medially on the dorsal side of the astragalus also seems to help facilitate the locking mechanism in the lamine astragalus. The articular facets of the astragalus of the new species are well defined, but not as well as those of H. macrocephala or Palaeolama. The astragalus of this species is also less robust than either of the other two species. The astragali of the new species are indistinguishable from those of H. edensis. e rugose overall. No femora of H. edensis were available for comparison. Notibia material was recovered for the new species. The astragali (Table 6 and Figures 19 and 20) of the new species are significantlysmaller than the astragali of H. macrocephala or Palaeolama (Figure 21). In fact, the astragali of the new species are approximately the same size as deer (Odocoileusvirginianus) astragali (Table 6 and Figure 21), and are sometimes confused with the latter. However, there are a few very important features on species that diagnose them as lamine. One feature is a very side that articulat th the cuboid and navi ular bones locking mechan is present in all other likely other hat fa uprigh

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34 A B Figure 17: Femora of new Hemiauchenia species, A. UF 45276, B. UF 45275, posterior view.

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35 6065707580859095100105152025303540Smallest width of the femur shaft in mm.Breadth of the proximal end in mm. New Species H. macrocephala Palaeolama L. glama Figure 18: Comparison of breadth of the proximal end to minimum shaft width of femora in four lamine species. These measurements were chosen due to limited specimen availability in the new species. Table 6: Measurements of the new species' astragali in mm. Catalog number Length of astragali Width of astragali UF 210722 38.0 24.1 UF 10281 40.4 25.4 UF 210706 38.6 24.3 Deer astragalus UF 210713 37.3 23.4

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36 Figure 19: Astragali of new Figure 20: Astragali of new species, A. UF 10281, B. UF 210722, C. UF 210706, dorsal view. Hemiauchenia species, A. UF 210706, B. UF 210722, C. UF 10281, plantar view. A B C A B C

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37 30354045505560651525354555Width of astragali in mm.Length of astragali in mm. New Species H. macrocephala andPalaeolama H. edensis L. glama Odocoileus virginianus Size comparisons of the astragali of the new Hemiauchenia species to otine species and deer, Odocoileus virginian Figure 21: her lamus. H. macrocephala and alaeolama astragali are indistinguishable. The calcanea (Table 7 and Figures 22 and 23) of the new species are smaller than e transversely narrower than la or Palaeos noticeabe new ticular facets on theneum of the new species appear to mirror those on alacets in the new species as prominent. The new species has very similar calcanea to “Hemiauchenia” minima in gracility, however, they are considerably smaller overall than those of “H.” minima. Like the astragali, the calcanea of the new species are indistinguishable from those of H. edensis. Table 7: Measurements of the new species' calcanea in mm. Catalog number Greatest length (GL) Length across the Coracoid process Length across the Sustentaculum tali P other Florida lamine species. The proximal tuber and shaft ar in H. macrocepha lama, and the tuber i ly more gracile in th species. Ar calca H. macrocephala, however, l the f re les UF 210709 broken 26.8 39.8 UF 210710 89.2 27.8 37.2

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38 One cuneiform and one unciform were recovered from the Inglis 1A site. However, these elements were so badly worn that there is little information to be gleaned from them besides their small size. The most remarkable aspect of the metapodials is their gracility as is indicated in Figure 24 and Table 8. The metapodials of the new species were absolutely shorter than those of H. macrocephala, longer than those of Palaeolama, and had a smaller diameter than metapodials of either of the other species (Figure 25). The metapodials of the new sp ecies are longer than those of H. edensis, but approximately the same width, making e metapodials of the new species appear more gracile. This intermediate length and Several proximal phalanges were found for the new species, as shown in Table 9 and Figure 26. These phalanges are characterized by being shorter and more gracile than the phalanges of H. macrocephala. The proximal phalanges have a triangular shaft shape with a wide posterior side and a tapering anterior side. They have the greatest length to width ratio (length/width 7.1, compared to 5.4 in H. macrocephala, 6.1 in H. edensis and 6.0 in extant lamines) out of all fossil Florida lamines and extant lamines, so the phalanges have a very gracile appearance. One medial phalanx and one distal phalanx were also recovered, they too, were smaller than either H. macrocephala or Palaeolama. The proximal phalanges of the new species have the W-shaped suspensory ligament scar at is diagnostic of the genus Hemiauchenia (Figure 26). th extremely narrow width are distinctive features of this new lamine. th

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39 A B Figure 22: Calcanea of new edial view. Figure 23: Calcanea of new species, A. UF 210710, B. UF 210709, anterior view. Hemiauchenia species, A. UF 210709, B. UF 210710, m A B

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40 Figure 24: Metatarsal of new Hemiauchenia species, UF 176935, anterior view.

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41 Table 8: Measurements of the new species' metapodials in mm. Catalog number Greatest length (GL) Breadth of proximal end (BP) Breadth of distal end (BD) Smallest diameter of diaphysis (SD) UF 176935 320 33.0 39.4 19.1 UF 18236 N/A N/A 38.2 N/A UF 210720 N/A 31.1 N/A 16.8 UF 210711 N/A 34.9 N/A N/A UF 210726 N/A 34.2 N/A N/A Figure 25: Comparison of metapodial lengths and widths in several species of lamines. H. vera and H. blancoensis specimens are from the F:AM. Unidentified extant lamines may be L. glama or L. guanicoe. Table 9: Measurements of the new species' proximal phalanges in mm. See abbreviations from Table 8. Catalog number GL BP BD SD UF 97203 74.5 17.8 14.9 11.3 UF 179638 82.6 18.5 16.0 10.7 UF 179639 84.2 19.6 14.6 11.1 UF 18237 71.2 17.8 14.4 10.8 UF 210704 66.2 17.7 14.1 10.4 15020030035040045050010203040Width in mm.Length in mm. New Species H. macrocephala Palaeolama Unidentified extant L.glama H. vera H. blancoensis 250 H. edensis U F 210708 78.5 18.3 15.8 10.3 UF 210712 67.9 18.1 14.9 10.6 U F 210703 67.7 18.1 14.5 10.4

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42 A B C D E F G Figure 26: Proximanus sory ligament scar on phalanges A and B, All 6 proximal phalanges display this scar. A. UF 210708, B. UF 210712, C. UF 210703 al phalanges of new Hemiauchenia species, anterior view, A-C mphalanges, D-G pes phalanges. A. UF 179638, B. UF 179639, C. UF 210708, D. UF 210703, E. UF 18237, F. UF 210712, G. UF 210704 A B C Figure 27: Proximal phalanges, posterior view highlighting the W-shaped suspen

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CHA PTER 4 LEOECOLOF THE NEECIES ew s is found as a part of a rich maian fauna. Atme two lorida fossil localities lived other terrestrial mammalian species including a wide range of carnivorans: Canidae, Ursidae, Procyonidae, Mustelidae, Felidae and Hyaenidae; other mammalian orders including Xenarthra, Insectivora, Chiroptera, and Lagomorpha; at least three families of Rodentia: Castoridae, Geomyidae, Muridae; and many other ungulates including families: Equidae, Tapiridae, Tayassuidae, Cervidae, Antilocapridae, Bovidae, Mammutidae and other similar sized artiodactyls (see Table 10 for a complete species listing of concurrent fauna at both Inglis 1A and DeSoto Shell Pits). Other artiodactyls that coexisted with the new species include: Hemiauchenia macrocephala, Odocoileus virginianus, and Capromeryx arizonensis (Ruez 2001). Body size can be a very useful indicator of the habitat in which an animal lives that have b in ungulates are long bone anis 1990; MacFadden and Hulbert 1990). Estimated body mass, used in conjunction anial pions, can givccurate asset of a speciesrred ever, long bone le to estimate body mass in camen be and tre should beed (Scott 19 PA OGY W SP The n pecies mmal the sa F and its role in that habitat (MacFadden and Hulbert 1990). Some good skeletal indicators een used to assess approximate body mass elements, especially the diameter of the femur (Scott 1985), and certain dental characters (J with postcr roport e an a ssmen prefe habitat. How using ngths lids ca problematic, herefo avoid 85). 43

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44 Table 10: Concurrent species at the Inglis 1A and DeSoto Shell Pits fossil localities. Xenarthra Dasypodidae Dasypus bellus Pampatheriidae Holmesina floridanus Glyptodontidae Glyptotherium arizonae Megalonychidae Megalonyx leptostomus Megatheridae Eremotherium eomigrans Insectivora Soricidae Blarina carolinensis C arnivora Pro Mu Trigonictis macrodon Hyaenidae Chasmaporthetes ossifragus Lagomorpha Leporidae Sylvilagus webbi Rodentia Sciuridae Sciurus sp. Geomyidae Orthogeomys propinetis Muridae Sigmodon curtisi Reithrodontomys sp. Ondatra idahoensis Canidae Canis edwardii cyonidae Procyon, sp. nov. stelidae

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45 Table 10: continued. Artiodactyla Camelidae Hemiauchenia macrocephala Hemiauchenia (sp. nov.) Antilocapridae Capro Cervidae Tapiridae Equus Mammutidae Table adapted from Ruez (2001). Dentition Dentition is often the best indicator for determining the type of diet to which a mammal is best adapted. In ungulates, it is possible to differentiate between a browsing and grazing diet by the morphology of the teeth. A grazing diet generally consists of 90% grasses. A browsing diet consists of 90% leaves and soft foliage. Mixed feeders fall in between these two extremes (Janis 1990). A dentition suitable for a grazing diet usually consists of hypsodont molars, with reinforcements of enamel or cementum, and broad incisor breadth. A dentition that is more indicative of a browsing diet has a narrow mandibular symphysis and brachydont molars with crenulated enamel that have little to no cementum. They also have narrower incisors that may be “feathered” for dexterity in stripping leaves from branches. Mixed feeders tend to share a suite of traits from both groups (Perez-Barberia and Gordon 2001;Webb and Stehli 1995). meryx arizonensis Odocoileus virginianus Perissodactyla Tapirus, sp. nov. Equus sp. Proboscidea Mammut americanum

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46 It is known from direct observation that Odocoileus virginianus is a browser, whereas, th e teeth of H. macrocephala suggest that it was probably a grazer to a mixed cies show properties of both browsers and grazers. The psodont than a deer, but less hypsodont than H. te coating of cementum, which tends to correlate 95). esowear yzing mesowear patterns in ungulate teeth is a new method to determine onty indices. Mesowear patterns refer to the esult of tooth attrition and abrasion (tooth on tooth and tooth on food wear) usp relief and cusp shape are the two major attributes that s. Cusp relief is the difference in height between cusp zers, there is little difference in these heights. hese heights. Cusp shape refers to the apex e or metacone in ungulates. The shape is described as sharp, rounded or based on qualitative characters and can be performed aked eye, or only with a hand lens (Fortelius and Solounias 2000). ia macrocephala and the new species of Hemiauchenia the s it is in all extant camelids. Likewise, the cusp ded or blunt in most cases. This combination of ed feeder, as are extant lamines. In Palaeolama and . edensis, on the other hand the cusp relief is high, but the cusp shape is sharp, feeder. The teeth of the new spe new species has teeth that are more hy macrocephala. They also have a comple with a grazing diet (Webb and Stehli 19 M Anal paleodiet in conjunction with hypsod combined r (Butler 1972; Fortelius 1985). C are analyzed in mesowear analyse tips and inter-cusp valleys. In pure gra Browsers have a large discrepancy between t of the paracon blunt. These analyses seem to be with the n In both Hemiauchen cusp relief could be referred to as high, a shape could be referred to as roun mesowear characters suggests a mix H

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47 indicating a browser ( Fortelius and Solounias 2000) (Figures 7 and 8 for new species, hala and Palaeolama, Figure 9 for H. edensis). on Isotope Analysis a useful tool for reconstructing herbivore l because the carbonate apatite found in tooth s high isotopic fidelity during fossilization (Feranec and MacFadden way utilized by consumed plants is preserved in the e 28). In the Pleistocene of Florida, plants that utilize the C4 rier grasses and shrubs and when ingested, produce tooth enamel signatures o 5, whereas, plants that use the C3 pathway are aves and softer foliage and when ingested, produce tooth enamel signatures with 13C values of approximately -20 to -10. Therefore, grazers will have a C4 signature in their tooth enamel and browsers will have a C3 signature in their tooth enamel; whereas, mixed feeders will have intermediate isotopic values (Hallman and Meachen in review; Koch et al. 1998). H. macrocephala from the Irvingtonian, appear to be mixed feeders, with 13C values of -2 to -7 (Feranec and MacFadden 2000; Hallman and Meachen in review). Feranec (2003) found that H. macrocephala was predominantly a browser to mixed feeder in the latest Blancan, however, the tooth morphology suggests these animals were mixed feeders to grazers, and warrants further analyses on this issue. The new Hemiauchenia species appears to be mainly a browser with a small amount of grass in the diet (13C of -8). It should also be noted that at all sites where Palaeolama mirifica is present, the new species is absent and vice versa (Morgan and Hulbert 1995; Ruez 2001). Figure 10 for H. macrocep Stable Carb Stable carbon isotope analysis is paleodiet (Koch et al. 1998). It is usefu enamel maintain 2000) and the photosynthetic path carbon compounds (Figur pathway are d with 13C values of approximately -2 t le

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48 This could possibly be due to direct competition for food, however, the 13C values fofor P. mirifica are between -10 and -14 , which have sufficiently more of a browsing signature than that of the new species (Figure 29)(Hallman and Meachen in review). This feeding strategy would have been effective for the new species, because it would avoid sharing a complete food source with either Odocoileus virginianus or H. macrocephala. The 13C values for the new s und pecies fall in between the 13C values for H. macrocephala and a pure browser. 13 Figure 28: Plot of C values versus fraction C4 contained in varied kinds of plant communities, and the corresponding enriched values of tooth enamel. From MacFadden and Cerling (1996).

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49 Figure 29: Stable carbon isotope data for three species of lamines, circles represent the new Hemiauchenia species, squares represent Hemiauchenia macrocephala and ovals represent Palaeolama mirifica. Data were taken from Hallman and Meachen (in review) and Feranec and MacFadden (2000). Diagram modeled after MacFadden and Cerling (1996). y mass, Limb Proportions According to Scott (1985), the limb proportions of extant ungulates will sort themin to five habitat categories: Flat, open grasslands or arid areas; mixed woodland andopen areas; heavy forest; true mountainous habitat; or rolling hill country. This may hold true for extinct ungulates. Although metapodials are not a good indicator of bodthey are an accurate indicator of habitat adaptation. The extant ungulates that live in open country have long metapodials (23-27 cm long); the ungulates from mountainous regions or thick forests have shorter metapodials (14-22 cm long); and the species living

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50 in scattered woodland or hilly habitats have intermediate length metapodials (Scott 1985). Based on the metapodial lengths of this new species and the comparison of the metapodial lengths of species concurrent in time span (H. macrocephala and Palaeolama) and the data in appendices 1 and 2 of Scott (1985), it seems that this new species was a scattered woodland dweller. The length of the metapodials of the new species fell in between those of H. macrocephala and Palaeolama, this suggests that it would be more likely to dwell in ope n habitat than Palaeolama, but less likely than H. macrocephala. Based on stable carbon isotope data this animal would be more likely to dwell in a forested habitat. The validity of either prediction can be assessed when the proposed habitats that this animal lived in are examined. Inglis 1A was thought to be a scrubby savanna habitat. Inglis 1C was thought to be a heavily forested neighboring habitat, but the new species was only present at Inglis 1A. One might expect to find this species in forested habitats as well, considering the amount of browse in its diet, the fact that it is absent from these forested sites suggests that it preferred scrubby savanna or open environments. The almost identical faunal composition at the DeSoto Shell Pits indicates a very similar type of environment Morgan and Hulbert 1995). The reason for this open habitat preference may have temmed from its mode of predator evasion. ong-limbed artiodactyls generally have heightened cursorial ability and use a cursorial vasion technique, whereas short-limbed artiodactyls prefer to take cover and freeze. The extremely long limbs of the new species suggest that it may have had heightened cursorial ability. This heightened cursorial ability would have made it more efficient for ( s Scott (1985) separated artiodactyls by their distinct modes of predator evasion. L e

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51 the new species to evade predators by running from them in an open habitat. Running in a forested habitat would be clumsy and the new species’ height would have made it difficult to hide from predators. For animals with shorter legs and smaller body sizes it is more efficient to use the hide and freeze mode of predator evasion, thereby necessitating that they dwell in a forested area where they could take cover (Scott 1985). This new species, perhaps dwelled in a location where a forested habitat bordered a scrubby habitat so it could take advantage of both forest and open environments. It probably preferred a savanna type habitat with scattered trees. The length of its limbs may also have facilitated the type of feeding used by the gracile African gerenuks, which stand on their hind legs to reach otherwise inaccessible leaves.

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CHAPTER 5 ing from the early Pliocene of Floridaay rred unning ala, another Florida lamine, possibly to avoid compet CONCLUSIONS The new species of Hemiauchenia is a gracile lamine that lived in Florida durthe late Pliocene and early Pleistocene. It is relatively rare and appears to have been sparsely distributed along the west coast of Florida. It may be a sister species to Hemiauchenia macrocephala, with which it was originally mistaken and with which it shares many morphological synapomorphies. Alternatively it may be a sister species toHemiauchenia edensis, a little known fossil lamine species and the southwestern United States, with which it also shares many morphological synapomorphies, such as gracile limb bones and a very similar dentition. Until a more detailed cladistic analysis is completed, the new gracile lamine species mbe considered closely related to both. This new species ate mainly browse with occasional grass mixed in, and prefeto dwell in a savanna type environment. It was probably an agile cursor, using its rand leaping abilities to avoid predators. It coexisted with Hemiauchenia macrocephbut has never been found at the same site with ition for a food source. The fossil sites at Inglis 1A and 1C were closed long ago, but the De Soto Shell Pit site is still active. More material of the new species from De Soto would help augment our understanding of the form and function of this animal. 52

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53 Placing new species on record will help future discoveries. It would also seem worthwhile to investigate Blancan/Irvingtonian fossil sites north and south of Ingliand DeSoto to see if the range of this lamine expanded farther than we now know. The possible astragalus found in the F:AM collection from Texas and the phalanx from Arizona hint that this species may have ranged into the tropical latitudes of M s 1A esoamerica. This nel l sil record, it may be a closer ancestor than any other known species. It would also be worthwhile to investigate further the Texas site from which the F:AM astragalus was found, to determine whether the new species was actually there or whether the astragalus belonged to the similar species, Hemiauchenia edensis. w species may have had an affiliation with the true (extant) llamas that crossed the isthmus of Panama in the early Pleistocene, as did Palaeolama. The morphologicasimilarities it shares with extant lamine species (e.g. small body size and reduced dentaformula) may not be a coincidence, but a true insight into the ancestral nature of the extant lamines. Although this lamine has at present a limited fos

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LITERATURE CITED Bell, C., E. Lundelius, A. Barnosky, R. Graham, E. H. Lindsay, D. R. Ruez, Jr., H. Rancholabrean mammal ages. Cenozoic Mammalian Biostratigraphy of North 1229-1247. Breyer, J. 1977. Intraand interspecific variation in the lower jaw of Hemiauchenia. 483. Dalquest, W. W. 1992. Problems in the nomenclature of North American Pleistocene of the Florida Peninsula. Ornithological Monographs 50: 1-113. Feranec, R.S. 2003. Stable isotopes, hypsodonty, and the paleodiet of Hemiauchenia generalization. Paleobiology 29(2): 230-242. Feranec, R. S. and B. J. MacFadden 2000. Evolution of the grazing niche in Pleistocene Palaeoclimatology, Palaeoecology 162: 155-169. Fortelius, M. 1985. Ungulate cheek teeth: Developmental, functional, and evolutionary gica 180: 1-76. using the abrasion-attrition wear gradient: A new method for reconstructing Bautista Creek and San Timoteo Canyon, southern California. University of California Bulletin of the Department of Geology 12(5): 277-424. Semken, S. D. Webb and R. Zakrzewski in press. The Blancan, Irvingtonian and America M. O. Woodburne. New York, Columbia University Press. Bender, M. L. 1973. Helium-Uranium dating of corals. Geochim. et Cosmochim 37: Journal of Paleontology 51(3): 527-535. Butler, P. M. 1972. Some functional aspects of molar evolution. Evolution 26(3): 474camels. Annales Zoologici Fennici 28(3-4): 291-299. Emslie, S. 1998. Avian community, climate and sea-level changes in the Plio-Pleistocene (Mammalia: Camelidae): A morphological specialization creating ecological mammals from Florida: Evidence from stable isotopes. Palaeogeography, interrelations. Acta Zoolo Fortelius, M. and N. Solounias 2000. Functional characterization of ungulate molars paleodiets. American Museum Novitates 3301: 1-36. Frick, C. 1921. Extinct vertebrate faunas of the Badlands of 54

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55 Hallmapartitioning in two sympatric lamines from the Florida Pleistocene using stable Natural History. Harrison, J. A. 1979. Revision of the Camelinae (Artiodactyla, Tylopoda) and description Contributions paper 95: 1-19. Honey, J. G., J. A. Harrison, D. R. Prothero and M. S. Stevens 1998. Camelidae. L. L. Jacobs. Cambridge, Cambridge University Press: 439-462. Janis, C. M. 1990. Correlation of cranial and dental variables with body size in ungulates Biological Implications. J. Damuth and B. J. MacFadden. Cambridge, University mammals in North America, part 1, Florida. Chemical Geology 152: 119-138. Macdonald, J. R. and W. J. Pelletier 1965. The Pliocene mammalian faunas of Nevada, 120. MacFadden, B.J. 1995. Magnetic ploarity stratigraphy and correlation of the Leisey Shell rough County, Florida. Bulletin of the Florida Museum of Natural History: Paleontology and Geology of the Leisey Shell Pits, Early Pleistocene of Florida. R. C. Hulbert, G. S. Morgan and S. D. Webb. Gainesville, Florida Museum of Natural History. 37, Part I: 107-116. MacFadden, B. J. and T. E. Cerling 1996. Mammalian herbivore communities, ancient feeding ecology, and carbon isotopes: A 10 million-year sequence from the Neogene of Florida. Journal of Vertebrate Paleontology 16(1):103-115. MacFadden, B. J. and R. C. Hulbert 1990. Body size estimates and size distribution of ungulate mammals from the late Miocene Love Bone Bed of Florida. Body Size in Mammalian Paleobiology: Estimation and Biological Implications. J. Damuth and B. J. MacFadden. Cambridge, Cambridge University Press: 337-363. Morgan, G. S. and R. C. Hulbert 1995. Overview of the geology and vertebrate biochronology of the Leisey Shell Pit Local Fauna, Hillsborough County, Florida. Bulletin of The Florida Museum of Natural History: Paleontology and Geology of the Leisey Shell Pits, Early Pleistocene of Florida. R. C. Hulbert, G. S. Morgan and S. D. Webb. Gainesville, Florida Museum of Natural History. 37, Part I: 1-92. n, D. P. and J. A. Meachen in review. Reconstructing paleoecology: Resource isotopes and postcranial morphometrics. Bulletin of the Florida Museum of of the new genus Alforjas. The University of Kansas Paleontological Evolution of Tertiary Mammals of North America. C. M. Janis, K. M. Scott and and macropodoids. Body Size in Mammalian Paleobiology: Estimation and of Cambridge Press: 255-299. Koch, P. L., K. A. Hope and S. D. Webb 1998. The isotopic ecology of late Pleistocene U.S.A. Proceedings of the 20th International Geological Congress, 1956: 119Pits, Tampa bay, Hillsbo

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56 Moya-Sola, S. and J. Agusti 1989. Bioevents and mammal successions in the Spanish Miocene. European Neogene Mammal Chronology E. H. Lindsay, V. Fahlbusch and P. Mein. New york, Plenum: 357-373. Perez-Barberia, F. J. and I. J. Gordon 2001. Relationships between oral morphology and feeding style in the Ungulata: A phylogenetically controlled evaluation. Proceedings of the Royal Society of London 268(1471): 1023-1032. Pickfor Morales and D. Soria 1995. Fossil camels from the upper Miocene of 50. 2001. Early Irvingtonian (latest Pliocene) rodents from Inglis 1C, Citrus 85. Allometric trends and locomotor adaptations in the Bovidae. Bulletin . and A. J. Bezuidenhout 1987. Anatomy of the Dromedary. Oxford, h, A. 1976. A Guide to the Measurement of Animal Bones from C. Hulbert, G. S. Morgan and H. F. Evans in press. Terrestrial mammals a unty Museum Contributions. e ce us d, M., J. Morales and D. Soria 1993. First fossil camels from Europe. Nature 365:701. Pickford, M., J. Europe: Implications for biogeography and faunal change. Geobios 28: 641-6 Ruez, D. R., Jr. County, Florida. Journal of Vertebrate Paleontology 21: 153-171. Scott, K. M. 19 of the American Museum of Natural History 179(2): 197-288. Smuts, M. M. S Oxford University Press. Von den Driesc Archaeological Sites. Cambridge, Peabody Museum of Archaeology and Ethnology. Webb, S. D. 1974. Pleistocene llamas of Florida, with a brief review of the Lamini. Pleistocene Mammals of Florida. S. D. Webb. Gainesville, The University Presses of Florida: 170-213. Webb, S. D. 1976. Mammalian faunal dynamics of the Great American Biotic Interchange. Paleobiology 2: 230-234. Webb, S. D., R. of the Palmetto fauna (early Pliocene, latest Hemphillian) from the central Floridphosphate mining district. Los Angeles Co Webb, S. D., B. J. MacFadden and J. A. Baskin 1981. Geology and paleontology of thLove Bone Bed from the late Miocene of Florida. American Journal of Scien281: 513-544. Webb, S. D. and J. A. Meachen in press. On the origin of Lamini including a new genand species.

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57 Webb, S. D. and F. G. Stehli 1995. Selenodont artiodactyls (Camelidae and Cervidae) from the Leisey Shell Pits Hillsborough County, Florida. Bulletin of the FloridaMuseum of Natural History: Paleontology and Geology of the Leisey Shell PEarly Pleistocene its, of Florida. R. C. Hulbert, G. S. Morgan and S. D. Webb. ille, Florida Museum of Natural History. 37, Part II: 621-644. Wheeleamelidae. an Society 54: 271-295. Gainesv r, J. C. 1995. Evolution and present situation of the South American CBiological Journal of the Linne

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BIOGRAPHICAL SKETCH Julie Ann a Meachen was born in Cleveland, Ohio, on August 28, 1978 to Sarah Elizabeth Mul Meachen and Edward Willis Meachen. She lived in many states across ee in zoology at the University of Florida, where she decided that she wantedd in the to the master’s program in zoology at the University of Florida in 2001. In Septove to Los Angeles, California, to begin her Ph.D. of Dr. Blaire Van Valkenburgh. the U.S. until the age of 11 when she moved to Tampa, Florida. She completed her bachelor’s degr to become a mammalian paleontologist. She worked and volunteere vertebrate paleontology collection at the Florida Museum of Natural History until her matriculation in ember of 2003, she will m studies at UCLA, where she will work on fossil Carnivorans under the guidance 58