FOSSIL CARNIVORES FROM THE LEISEY SHELL PITS, HILLSBOROUGH COUNTY, FLORIDA Annalisa Bertal ABSTRACT Thirteen species ofcarnivores have been recovered from the Inisey Shell Pits, late early Irvingtonian of Florida These species include Armbruster's wolf Canis armbrusteri, the wolf coyote Canis edwardii, the gray fox Urocyon cinereoargenteus, the tremarctine bear Arctodus pristinus, the gracile sabercat Smilodon gracilis, the machairodont Homotherium sp., the bobcat Lynx r:08, the cheetah-like cat MiracinonB inexpectatus, the raccoon Procyon lotor, the long-tailed weasel Mustela Renata, the spotted skunk Spilogale putorius,lhe Tiver daer Lutra canadensis, and the Casibbean monk sed Monachus tropicalis. Of significance, the Leisey carnivore fauna includes (1) the largest most complete sample of Smitodon gracilis, and (2) the first Irvingtonian record from Florida of Lutra canadensis. The Leisey carnivores rank with those from Coleman 2A and Inglis IA asthe most diverse carnivore faunas known from the Irvingtonian ofFlorida. RESUMEN Se han recuperado trece especies de carnivoros de los dep6sitos de Conchuelas de Lcisey de fines del Irvingtoniano temprano de Florida. Estas especies incluyen el lobo de Armbruster Canis armbrusteri, el coyote lobo Canis edwardii, el zorro gris Urocyon cinereoargenteus, el oso tremarctino Arcrodus pnstinus, el tigre dientes de sable grkil Smilodon gracilis, el macairodonte Homotherium sp., el lince Lynx mfus, el felino tipo guepardo Miracinonyx mexpectatus, el mapache Procyon lotor, la comadreja de cola larga Mustela*enata, el zorrillo moteado Spilogale putorius, la nutria de rio Lutra canadensis y la foca monje del CaribeMonachus tropicalis. Significativamente, la fauna de carnivoros de Lcisey incluye: (1) La mayor y mdE completa muestra de Smilodon gracilis y (2) el primer registro de Lutra canadensis para el Irvingtoniano de Florida. Los camivoros de Leisey son comparables con los de Coleman 2A e Inglis IA en ser la fauna conocida de carnivoros mas diversa del Irvingtoniano de Florida 1 The author is a Professorin the Department of Biology, San Diego State University, San Diego CA 92182. . BERTA, A 1995. Fossil carnivores from theLeisey Shell Pits, Hillsborough County, Florida. Bull. Florida Mus. Nat }list 37 PL II(14):463-499. 464 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, Fr. II, No. 14 INTRODUCTION The Leisey carnivores, a relatively large and diverse sample of well preserved specimens, include cranial, dental, and postcranial remains from six families: Mustclidae, Felidae, Procyonidae, Canidae, Phocidae, and Ursidae. Thirteen species of carnivores are represented with a combined minimum number of 36 individuals (Table 1). In comparison to other Florida Irvingtonian localities Leisey ranks with Inglis lA and slightly behind Coleman 2A in diversity of carnivores (Webb 1974a: table 2.1; Martin 1974: table 3.1) At Leisey large carnivores are more numerous and diverse than small carnivores (Table 1). The large carnivores include a large and a small wolf, Canis armbrusteri and C edwardii; a tremarctine bear, Arctodus. two machairodont cats, Smilodon and Homotherium; and the cheetah-like cat Miracinonyx. The Inglis lA fauna shares all these tan, except it has only the small wolf, C edwardii. One additional large carnivore not represented at Leisey but present at Inglis lA is the hyaena Chasmaporthetes. The Coleman 2A large carnivore fauna comprises C armbrusteri,Arctodus, andthe jaguar, Panthera onca. Small carnivores at Lcisey include a fox, Urocyon; a raccoon, Procyon; a weasel, Mustela; a skunk Spilogale; an otter, Lutra; and a bobcat, Lynx rufus. This assemblage is similar to both Inglis lA and Coleman 24 although less diverse in lacking the skunks Conepatus and Mephitis, and unique in the presence of an otter, Lutra. With the exception of the Ursidae (Emslie this volume), representatives of the other five carnivore families will be discussed in this paper. The format for each taxon includes a list of referred specimens, description and/or discussion of this material, and the chronology and stage of evolution of the species as indicated by other occurrences. The method of measurement for Smilodon follows Berta (1987). All measurements are in millimeters (mm) unless indicated otherwise. Abbreviations cited in the tables are as follows: N, number of specimens measured (sample size); X mean; and OR observed range. ACKNOWLEDGEMENTS I thank Toni TKjada-Flores and Christopher Shaw ofthe George C. Page Museum 16(- their help in my study of the Rancho La Brea Smilodon. I am especially grateful to Toni for measuring specimens in these collections Ralph (Tony) Estevez, Frank A. Garcia~ and James L Pendergraft donated important specimens to the Florida Museum of Natural History from Leisey 3. BERTA: LEISEY SHELL PIT FOSSIL CARNIVORES 465 TABLE 1. & Lcisey Shell Pit local fauna. Minimum number Lcisey carnivores ofindividuals Family Canidac Canis armbrusteri 3 Canis edwardii 3 Uro€yon cinereourgenteus 1 Family Ursidae Arctodus pristim,3 8 Family Progyonidae Procyon lotor Family Mustelidae Lutra canadends 2 Spilogale putorius 1 Mustelafrenata 1 Family Felidae Lynx rufus 2 Smilodon gracilis 8 Homotherium sp. 3 Miracinon>x inexpectatus 1 Family Phocidae Mc)nachus tropicalis TOTAL. 36 SYSTEMATIC PALEONTOLOGY Order CARNIVORA Bowdich 1821 Family CANIDAE Gray 1821 Canis annbrusteri Gidley 1813 Referred Specimens.- Leisey lA: UF 81654, left maxillary fragment with P4-Ml; 81655, right maxillary with Ml-2; 81152, right edentulous mandibular ramus; 67091, upper canine; 80662, 81661, left P4; 81656, right P4; 81657, right Ml; 81658-81659, right p4; 81660, left ml; 87283, fragment of left mandible with p4. Leisey 3: UF 142230, right proximal radius; 132049, right metatarsal IV. Discussion.- Two large fossil wolves are known from Florida, Armbruster's wolf, Canis armbrusteri, and the dire wolf, Canis dirus (Berta 1981). The smaller of the two, C armbrusteri, also is known from three other Irvingtonian faunas in Florida, Coleman 24 McLeod Limerock Mine, and Haile 21A. The Leisey sample of C annbrusteri resembles specimens from Coleman 2A described as C lupus by 466 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, Fr. II, No. 14 Martin (1974). Although C annbrusteri is closely related to gray wolves, it can be distinguished from C lupus by several characters, one of which is apparent among specimens from Leisey, a pronounced buccal cingulum on the Ml (Fig. lA-B). Comparison of measurements of fossil wolves from the Florida Pleistocene (Martin 1974, table 3.11) with the Leisey sample (Table 2) indicates that the latter sample falls within the size range of Coleman 2A. C armbrusteri is distinctly smaller than either Florida or Rancho La Brea C dirus. The smaller size of C armbrusteri, especially as regards carna~ial size, was previously noted and figured by Kurt6n and Anderson (1980). Canis edwardii Gazin 1942 ' Referred Specimens.- Leisey lA: UF 67092, posterior skull fragment; 81665, right maxilla with P4-M2; 80372, right premaxilla with I3; 81666, right maxilla with Ml-2; 81663, left maxilla with Pl-2; 81664, right maxilla with P3; 63667, left mandible with pl-ml; 64399, left mandible with ml; 81670, juvenile right mandible with p3-ml in crypt; 87291, edentulous maxilla; 81685, edentulous mandible; 67090, 81667, right upper canine; 81682, partial left P3; 80662, left P4; 81668, right P4; 81669, right Ml; 81671, right lower canine; 81675, right p2; 81674, right p3; 87285, right ml, 81673, partial right ml; 81672, 87297, left ml; 81677, atlas; 87284, 63671, 87296, axis; 81677, cervical vertebrae; 81127, 81509, 81679, left distal humeri; 81680, right distal radius; 81686, right metacarpal II; 87286, right metacarpal II (juvenile); 87292, left proximal metacarpal II; 87294, right metacarpal III; 87293, 63668, right metacarpal IV; 81219, right distal femur; 67093, 67094, right distal tibiae; 81681, left distal tibia; 63669, right metatarsal III; 63670, right metatarsal IV; 87295, medial phalanx; 87288, 87289, proximal phalanx. Discussion.- Canis edwardii is the best represented canid at Leisey and includes adult and juvenile individuals. In addition to fragmentary maxillae and mandibles, skeletal elements are known. Other Florida Irvingtonian occurrences of this species include Inglis 1 4 Haile 21 4 and Rigby Shell PiL This species of small fossil wolf is thought to be related to the red wolf, Canis rufus (Tedford pers. comm.). According to Tedford, whose usage is followed here, material referred to Canis priscolatrans by Kurt6n and Anderson (1980) should be referred to this species. Among those characters which distinguish C edwardii from C rudus are longer, narrower carnassials (Fig. IC), and smaller M2 with broad hypocone and weak anterobuccal cingulum (Tedford pers. comm.). Measurements of C edwardii from Leisey are compared with some other Florida occurrences of this species in Table 3. The Leisey C edwardii resembles in size and morphology BERTA: LEISEY SHELL PIT FOSSIL CARNIVORES 467 A if- fil -r /../ - c TE- Figure 1. Canidae and hocyonidae from kisey Shell Pit 1 A (A) Canis armbrusteri, UF 81654, occlusal view of left maxilla with P4-Ml; (B) C armbrusteri, UF 81655, occlusal view of right maxilla with Ml- M2; (C) C. edward,i, I_IF 81665, occlusal view of right maxilla with P#*12; (D) occlusal and (IE) lateral views ofProcyon lotor, UF 80588, right mandible with ml. Scale bar 20 mm in length. TABLE 2. Upper tooth measurements of Canis armbrusten from Leisey Shell Pit UF UF UF UP 81654 81655 81656 81657 P4 length 25.4 - 26.3 - width 12.2 -- 12.1 -- Ml length 16.4 15.9 - 14.5 width 22.4 22.2 - 20.9 M2 length - 8.4 - - width - 13.6 - - 468 BULLEI'IN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, Fr. II, No. 14 TABLE 3. Dental and mandibular measurements for Canis edward,7. Lcisey Haile 21A Rigby Shell Pit N X OR (UF 62561-62562) (UF 40090-40091) P4 length 2 21.5 20.5-22.5 22.5 19.0 width 2 10.3 10.1-10.5 11.4 9.0 M 1 length 3 19.3 18.4-20.1 20.0 width 3 14.6 14.5-14.8 14.3 13.5 M2 length 2 12.3 12.2-12.3 12.0 - width 2 7.4 6.8-8.0 7.6 - Depth jaw below middle m 1 1 25.8 - - pl length 1 6.1 5.2a - width 1 4.0 - - p2 length 1 9.Be 13.1 10.7 width - - 5.4 4.4 p3 length 2 - 14.0 - width 2 - 5.7 p# length 1 13.5 14.5 - width 1 6.4 7.2 - ml length 2 - 24.7 - width 2 - 10.5 - specimens from Haile 21A. Study of skeletal material, including tibiae, femur, radius, cervical vertebrae, and metapodials, referred to this species from Leisey also compared best with the Haile 21A sample. Urocyon cinereoargenteus (Schreber 1115) Referred Specimens.- Leisey lA: UF 81653, right calcaneum. Discussion.- This specimen agrees in detail with Recent Florida specimens of Urocyon cinereoargenteus. Other Florida Irvingtonian records of this species occur at Inglis lA (Webb 1974a) and Coleman 2A (Martin 1974). Although Martin originally described the Coleman specimens as a new fossil species of gray fox, U. minicephalus, I follow Kurt6n and Anderson (1980) in recognizing it as a junior synonym of the living gray fox, U cinereoargenteus. BERTA. LEISEY SHELL Prr FOSSIL CARNIVORES 469 Family PROCYONIDAE Bonaparte 1850 Procyon lotor (Linnaeus 1758) Referred Specimens.- Leisey lA: UF 80588, right mandible with posterior crown ofp2 and ml. Leisey 1: UF 91016, left distal humerus. Discussion.- The Leisey raccoon is tentatively referred to the modern species Procyon lotor. Comparison of Recent specimens indicates that the fossil material differs in its considerably larger size and differing morphology of ml, in which an additional cusp is positioned behind the metaconid (Fig. 1C-D; Table 4). Further study of the fossil m,terial, especially the large Inglis lA sample, may warrant establishment of a new species (see also Morgan and Hulbert this volume). This species was previously recorded from a single Florida Irvingtonian fauna Coleman 2A (Martin 1974). In addition to Leisey, other Florida Irvingtonian faunas that record the presence of Procyon are Inglis 14 Haile 214 and Haile 16A. The stage of evolution of the Leisey specimen is most similar to that represented by the Inglis jaws. Table 4. Mandibular and tooth measurements of fossil and Recen:Proown lotor from Florida. Recent Leisey l A Coleman 2A Inglis l A Haile 16A UF 80588 UF 13163 UF 49251 UF 46665 N X OR Toothrow length (p 1-m2) 48.Oe 39.9 45.7 48.0 5 37.2 36.7-37.6 Depth of mandible below m 1 14.8 - 8.3 - - - - m 1 length 12.5 - 12.1 - 5 9.4 9.1-9.5 width 7.7 - 7.7 - 3 6.2 6.0-6.3 Family MUSTELIDAE Swainson 1835 Muste/aj~enata Lichtenstein 1831 Referred Specimens.- Leisey 3A: UF 95765, right mandible with p4-ml. Discussion.- This specimen (Fig. 2A) is very similar to the Recent Florida long-tailed weasel, Muste/a «#enata. It is larger than the Devil's Den M kenata, the only other Pleistocene record of this species from Florida (Table 5). 470 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, Fr. H, No. 14 Lutra canadensis (Schreber 1776) Referred Specimens.- Ieisey IA: UF 81651, left maxillary with P3; 81652, left Ml. Leisey 3: UF 124633, left mandible with p3-ml. Discussion.- The Leisey otter resembles the fossil species Lutra parvicuspis Gidley and Gazin (1933) in its large size (Fig. 2D-IE) and P3 with conspicuous posteromedial basin. However, the Leisey specimen differs from L. parvicuspis in having a broadly proportioned paracone and metacone (Fig. 2D; Table 6). I follow Kurt6n and Anderson (1980) and other workers in synonymizing L. parvicuspis with the modern species. The Leisey occurrence represents the first In'ingtonian record of this species in Florida. Other Irvingtonian occurrences of L. canadensis include the slightly younger Cumberland Cave and Port Kennedy Cave faunas of middle Irvingtonian age. Spilogale putorius (Unnaeus 1758) Referred Specimens.- Leisey 3: UF 142229, right mandible with ml. Discussion.-- The Leisey specimen (Fig 2B-C) compares favorably with a large sample of mandibles from Inglis lA listed as Spilogale putorius by Webb A~ D~ B E UF# 42229ir C Figure 2. Mustelidae from the Lcisey Shell pits. (A) Mustela *enata, UF 95765, lateral view, right mandible with p*m 1, (B) lateral and (C) occlusal views, Spilogale puronus, UF 142229, right mandible with ml; (D) Lutra canadensis, UF 81652, occlusal view, left Ml, (E) L. canadensis, UF 124633, lateral view, left mandible with p3-ml. Scale bar 14 mm in length for A-C, 16 mm for D, and 20 mm for E BERTA: LEISEY SHELL PIT FOSSIL CARNIVORES 471 Table 3. Dcntal measurements offossil and RecentMustela,»enata from Florida Recent I.cisey 3A Devil's Den UF 95765 UF 16542 NX OR p4 length 3.3 - 4 3.4 3.0-3.7 width 2.0 - 4 1.7 1.5-1.9 ml length 6.4 5.9 4 5.9 5.1-6.4 width 2.8 2.2 4 2.3 2.2-2.5 Table 6. Upper tooth measurements of fossil and Recent Lurra from Florida Lutra canadensis Lutra parvicuspial UF 81631- 81652 NX OR NX OR P3 length 9.0 5 7.6 6.8-8.0 3 7.8 7.5-8.2 width 5.8 5 5.2 5.0-5.5 3 5.4 5.0-6.0 Ml length 11.7 5 9.4 9.1-9.9 - - - width 13.6 5 12.2 11.6-12.7 - - 1 Meam,ements aftcy Gidley and Gazm (1933). (1974a). However, the ml of the Leisey specimen is slightly larger (anteroposterior length 9.3 mm) than most of the Inglis mls. Both the Leisey and Inglis specimens are larger than mandibles of the extant spotted skunk from Florida, but are similar in size to those of larger western subspecies of S putorius. Kurt6n and Anderson (1980) referred all North American Irvingtonian spotted skunks to S. putorius. Family FELIDAE Gray 1821 Lynx ndus (Schreber 1776) Referred Specimens.- Leisey lA: UF 81690, right calcaneum; 81691, metatarsal III; 87300, proximal right ulna; 87299, proximal right metacarpal III. 472 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, Fr. II, No. 14 Leisey 3: UF 124631, left mandible with c, p3-ml; 130012, right proximal humerus; 102668, right distal radius; 130013, left proximal ulna; 102669, left distal femur. Discussion.- All elements compare well in most features with Recent Lynx rufus from Florida (Fig. 3). Differences seen in the Leisey calcaneum are a rounded posterior articular surface and a relatively short shaft. In the modern species the posterior articular surface is nearly triangular, and the shaft is long relative to total length. The Leisey metatarsal differs from Recent specimens of L. rufus in having a more extensive proximal posterior projection that is not as broad transversely (Table 7). A1 Figure 3. Bobcat (Lynx rufus) from Leisey 3. (A) lateral and (B) occlusal views. UF 124631, left mandible with c and p)-ml. Scale bar 20 nam in length. This species occurs commonly in Florida Rancholabrean faunas and has been recorded previously from two Irvingtonian localities, Inglis lA and Coleman 2A (Kurt6n 1965; Webb 1974a). In his review of small Pleistocene felines of North America Werderlin (1985) referred all pre-Wisconsin species of Lynx rufus (with the exception of the type of L. r. koakudsi) to Lynx nifus calcaratus. He listed the temporal range of this subspecies as Blancan (Cita Canyon, Texas) to late Sangamonian (Sabertooth Cave, Florida). As Werderlin commented (1985:199), "...it seems possible that these specimens may eventually be referrable to a species distinct from Lynx rufus... this can only be substantiated through discovery of complete skulls, or ... postcranial material that would permit calculation of limb propoidons: BERTA- LEISEY SHELL PIT FOSSIL CARNIVORES 473 Table 7. Measureme[Its ofhindlimb elements of fossil and Recent Lynx nBs from Florida Recent Ldry lA Calcaneum UF 81690 N X OR length 36.6 2 39.5 38.041.0 width 12.1 2 11.6 9.8-13.4 shaA length 23.0 2 24.8 23.945.7 Recent Leisey lA Metatarsal III UF 81691 N X OR length 61.4 2 66.2 65.0-67.4 proximal width 8.3 2 9.1 8.6-10.6 proximal depth 10.0 2 11.2 10.5-11.9 Smilodon gracilis (Slope 1880) Referred Specimens.- Ikisey lA: UF 81700, nearly complete skull with left I2-3, P34, Ml and right Il-3, P.3-4; 87238, partial skull with P34 and complete braincase; 87239, partial skull with left P34 and complete braincase; 87242, occipital region of skull; 87240, right masmid region; 87237, right premaxillary with Il-3; 81714, 87236, left premaxilla with I2-3; 63666, 81617, left premaxilla with I3; 84187, right premaxilla with I3 ; 83050, right maxilla with P4; 87243, 87245, left maxilla with P34; 63652, left maxilla with P4; 87246, right maxilla with P34; 80463, edentulous left maxilla fragment; 87241, 87261, left m,ndible with p4-ml; 81723, nearly complete right mandible with i3-c, p4-m 1, 81724, left mandible with c, p3-ml; 86746, nearly complete right mandible; 82529, right mandible with p3-ml; 87258, associated right mandible with p4-ml and left mandible with p4; 63653, right mandibular ramus with p4; 87275, left mandible with p4; 63655, 63654, 81722, 87244, 80182, 87259, 82251, right upper canines; 86843, 80111, 87276, 84189, left upper canines; 87251, 87260, left I2; 81220, right Il, 87251, 87260, left, I2; 87254, right I2; 80221, leR B; 86692, right fragmentary P4; 67086, 81472, 81017, 83087, left lower cnnines; 84188, right lower canine; 83088, left p4; 63656, left ml; 63657, partial right ml; 81829, 81726, 81725, 87303, atlas; 67084, 87255, 81507, 67085, 82834, axis; 81828, cervical vertebra; 80028, 2 thoracic vertebrae; 81154, 80162, 81294, 80088, 474 BULLEnN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, Fr. Il, No. 14 lumbar vertebrae; 63665, 65397, sacrum; 65074, 81715, 80183, proximal right scapula; 67083, 87304, proximal left scapula; 63659, 81727, 87277, right humerus; 63661, 80184, 67087, right distal humerus; 83581, 81153, 87278, 87307, left humerus; 81371, 81730, 80962, right radius; 81729, left radius; 63663, proximal left radius; 87281, left ulna (distal end missing); 87280, right ulna (proximal and distal end missing); 83429, 63660, 81370, proximal right ulna; 85321, 81728, left ulna; 81221, right innominate; 87256, 87259, associated right and left innominate; 80319, 80087, 80663, 80222, 67082, right partial innominate; 80223, 63664, 81618, 87228, left partial innominate; 67081, 82652, right femur, 87272, proximal left femur, 81732, proximal right tibia; 81731, left tibia; 80917, patella; 87274, proxi,nal left fibula; 67088, distal left fibula; 87259, left astragalus; 87264, left calcaneum; 87267, left magnum; 87248, left scapholunar; 86963, left pisiform; 87269, left metacarpal III; 87273, left metatarsal IV; 86747, right metacarpal IV; 81712, right metatarsal IV; 86804, right metatarsal V; 87253, 87262, 87247, 88048, proximal phalanx; 81713, 81508, 87263, 81711, 80161, 87270, medial phalanx; 81710, 81295, 87271, 87252, ungual phalanx. Leisey 3: UF 142231, proximal two-thirds ofright radius. Description- The Florida record of Smilodon gracilis has been recently reviewed (Berta 1987). The following supplemental description is based on the Leisey sample. Comparisons are made between this sample, previously known material of this species, and the more advanced Rancholabrean species, Smilodon populator (Berta 1985). Skull: Smi/odon gracilis was described by Cope (1880) largely on the basis of a crushed and distorted skull and mandible. To this can now be added from Leisey a nearly complete skull (UF 81700) and two partial skulls with complete basicrania (UF 87238, 87239). The Leisey skulls are smaller and more lightly built than S populator (Fig. 4, Table 8). The muzzle is shorter and namwer in this species (Table 8). The anterior narial opening is narrower than in S populator and rises more steeply along its lateral margin owing to relatively less inflated canine tracts. The zygomata are deep and arched with the posterior root of the arch ventrally extended, as is characteristic of the genus. The sagittal crest, although abraded and worn in the most complete specimen, is prominent and elevated above the parietals. The pOStOIbital processes are relatively large, blunt protuberances. The braincase is posteriorly expanded, and its surface pitted by several large fossae near its junction with the lambdoidal crests. Lambdoidal crests are strongly posteriorly and ventrally directed and enclose a rounded occiput. Several pairs of deep depressions for insertion of the rectus capitus muscles are positioned on either side of the strong venical keel. The occiput is shorter relative to skull length in this species (Table 8). BERTA: LEISEY SHELL PIT FOSSIL CARNIVORES 475 j A . .Fli... l.6/¢r A r U B Figure 4. Sabercat skull from Lkisey lA (A) lateral and (B) ventral views of Smilodon gracihs, UF 81700, skull with rigid and left 12-13, P)-P4, and right Il. The left canine belongs to a different individual. The teeth were coated for photography. Scale bar 50 mm in length. The optic foramen is small, and the large sphenoidal fissure and foramen rotundum are positioned along the lateral side of the alisphenoid. A large bony strut separates the foramen rotundum from the foramen ovale. The foramen ovale is directed ventrally, as in S populator. 476 BULLEI'IN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, Fr. II, No. 14 Table 8. C . ofcranial meagurements of Smilodon from kisey lA and Rancho La Brea. Smilodon gracilis Smilodon populatorl FLORIDA CALIFORNIA 1£isey lA Rancho La Brea UF 81700 UF 87239 N OR Length, premaxillary- condyles 250 - 23 271.4-344.1 IEngth, postglenoid cavity-condyles 73.9 73.0 25 83.4-117.8 Greatest width across muzzle at C 70.0 - 25 93.1-117.8 Greatest width across postorbital processes 671 - 25 104.1-130.2 Least width behind posto,bital processes 44.8 - 25 52.9- 65.9 Palatal width at C 38.le - 25 48.9- 63.0 Palatal width at P4 63.8 - 24 114.7-135.4 Greatest transverse diameter, - auditory bulla 36.7 - 23 46.7- 63.9 Greatest width across magtoid processes 98.5 94.3 24 122.0-154.3 Greatest widlh across con(lyles 51.5 51.8 25 57.4- 72.3 Minimum distance from oociput-~amen magnum notch 52 - 10 63.3- 90.8* The palate is posteriorly broad. Large round embrasure pits are located between P4 and Ml for occlusion with ml. On either side of the midline of the palate are a pair of narrow palatal grooves that extend anteriorly. Lateral to these grooves is a large median trough bounded by another ridge. The ear region of S gracihs presents more significant differences thgn other portions of the skull and is discussed in greater detail. The mastoid is not as enlarged relative to the bulla as in S populator, and it does not extend as far anteriorly as in the late Pleistocene species. The paroccipital processes are defined as pointed projections on either side of depressions in the exoccipital. In comparison to S. populator, the tubular external auditory meatus forms a more open channel owing to a less expanded magtoid farther separated from the postglenoid process (Fig. 4). The postglenoid foramen is positioned at the base of the postglenoid process near the entrance of the external auditofy meatus. It is BERTA: LEISEY SHELL Prr FOSSIL CARNIVORES 477 variable in size; very reduced in UF 81700, large in UF 87238, and very large in UF 87239. The posterior portion of the postglenoid process is enlarged and extends farther anteriorly than in S populator. In ventral view the mastoid terminates near the level of the glenoid fossa and does not extend medially around the bulla as in most S populator. The mastoid and bulla in S. gracilis are farther separated from one another than in S populator. This is especially true for the posterior portion of these elements which remain separate rather than coalescing, as is common in S. popu/ator. The bullae are large and globose. The ventral walls of the bullae are broken in all specimens, exposing the internal structure of the ear, and it is comparable to S populator in structure. The septum dividing the tympanic cavity into a small ectotympanic (tympanic) and large entotympanic (caudal entotympanic of Hunt 1974) is apparent, as is the septum separating the entotympanic into an anterior and posterior chamber. The posterior chamber is much larger and extends into the m=gtoid process. Upper Dentition: Judging from the varying stages of tooth wear represented, the Leisey sample includes individuals that ranged in age from young to very old adults. Upper incisors preserved in the skull (UF 81700) and an edentulous maxilla (UF 80463) are arranged transversally in an arc and are very similar in morphology to S. populator. The canine is proportionally smaller, shorter and less curved in S. gracilis than in S. populatee (Fig. 5; Table 9). This tooth is set in the maxilla nearly in line with the cheek toothrow (Fig. 4). The swollen tracts of the canine roots extend to the interorbital constriction. In S populator the canine is medially positioned in the jaw and offset from the cheek toothrow. Other than the smaller size, P3-4 in the Leisey sample compare well with previous known material (Table 10). Very fine serrations are present along the anterior and posterior tooth margins 6f P3. P4 presents several distinctive features of S gracilis, a small ectoparastyle, a distinct protocone, and a separate protocone root. Mandible and Lower Dentition: The mandible and lower dentition of S gracilis are well represented in the Leiscy sample and compare favorably in size and morphology with previously known samples (Fig. 6; Table 11). In all Leisey specimens p3 is separated from p4-ml by a small diastema. In contrast to the variability of p3 seen in other samples of S. gracilis, only one specimen of p3 in the Leisey sample, UF 87258, has two distinctly separate roots. In all other specimens p3 is single rooted with a lateral sulcus at crown level dividing the tooth part way down the root and fusing into a single root socket. Since absence of this tooth is common among late Pleistocene S popu/ator, the more common single rooted condition of this tooth in the Leisey sample, as compared with previously 478 BULLEI'IN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, Fr. II, No. 14 A B J\P Figure 5. Two Smilodon gracilis upper canines from 1.£isey 1 A. (A) UF 81722; (B) UF 82251. Scale bar 30 mm in length. known samples, is considered evidence of its more advanced stage of evolution. In UF 87258, p4 differs from previously known specimens of this species in lacking an anterior cusplet. Comparison of the length of the lower carnassial indicates that the Leisey sample is intermediate in size between that of Inglis lA and Port Kennedy specimens (Table 11). Skeleton: The remainder of the axial skeleton is represented by elements from the major regions of the vertebral column including the cervical, thoracic, and sacral BERTA: LEISEY SHELL PIT FOSSIL CARNIVORES 479 Table 9. Comparison of upper Lasey lA and PR,¥*0 La Brea. A-B lengthl C-D length2 Arc3 S. gracilis FLORIDA Izisey lA UF 87244 142.6 44.6 3.197 UF 87122 162.0 32.0 3.114 UF 86692 153.4 52.1 2.943 UF 80111 161.0 53.5 3.012 UF 87276 145.7 47.2 3.086 UF 84189 147.9 44.0 3.361 S. populatorA CALIFORNIA Rancho La Brea 2000*48 247.0 72.0 3.430 2000-R-24 225.0 63.0 3.571 2000-L-43 216.0 69.0 3.130 2000-R-21 229.0 68.0 3.367 2000-R-5 226.0 63.0 3.587 2000-R-47 206.0 62.0 3.322 20004.16 232.0 66.0 3.515 2000-R-31 234.0 63.0 3.600 20004,26 262.0 80.0 3.275 2000-L-2 200.0 58.0 3.448 1 Total tooth lensth measured Rom canme t* to bue of mot 3 Arc 1,~th meamied no,mal to-amol ma*L 3A 4 M ' ba-d on th= JJdy. vertebrae. With the exception of two cervical vertebrae from Inglis IA, none of these elements was previously known. Other than their smaller size, these elements are virtually indistinguishable from those of Smilodon populator (Table 12). Both the atlas and axis display characteristic Smilodon features. In the atlas the transverse process extends posteriorly well behind the aniculations for the axis. In the axis, the neural spine projects posteriorly behind the posterior zygopophyses. The sacrum is like S populator in having small anterior zygopophyses that do not extend high above the dorsal surface, and a broad dorsal margin lateral to the anterior zygopophyses. Table 10. Measurements of upper dentition ofimilodon. * Il n B P3 P4 Ml L W L W L W L W L W L W S. gracilis FLORIDA (Leiscy) UF 87242 7.3 8.6 10.3 - - - - UF 87243 - 14.0 6.2 31.3 11.9 UF 87246 - 13.5 6.3 31.6 11.4 UF 87238 - - - - - - 15.3 7.4 32.4 14.3 UF 81700 07.3 7.9 10.0 9.0 10.7 10.9 14.7 7.2 32.0 13.9 (Inglis IAl) 13.9 7.1 32.0- 11.ge- (Mcimd) - 16.5 7.8 33.7e 12.9(2) 5.1 7.5e (Bass Point Watcnvay) I2 8.5- 10.1- 16.2- 7.0- 30.4 11.5e - - (El Jobean Pit2) 9.1(2) 10.2(2) 17.0(2) 7.*2) 34.2 11.9e 17.3 7.1 30.5 12.7 B U LLE rIN FLO R ID A M U S E U M N AT U R A L H lb O R Y V O L. 37, F r. II, N o. 14 PENNSYLVANIA 0 15.4- 6.5- 32.0- 11.8- (Port Kennedyl) 7.5 5.4 8.7 7.4 10.3 9.0 16.0(2) 7.2(2) 33.7(2) 12.9(2) 8.4 4.7 S. populator FLORIDA (Ichetucknee3) UF 3470 - 41.1 16.0 (Melbourne3) USNM - 35.7 CALIFORNIA (Rancho IA Brea4) Minimum - 5.0 7.6 11.0 15.0 8.4 37.5 14.2 Maximum - 7.6 9.7 13.0 19.7 10.6 46.0 19.9 (N=23) 1 2M foll,~,ir Ch:rir (1985).3M fallowin, Kunen (1965). 4 Meamucmeots follao,ing Merriam mt Stock (1932). foll.4.5 B.'ta (iggl). BERTA LEISEY SHELL PIT FOSSIL CARNIVORES 481 A r 8 - 1 - L~ Figure 6. Mandibles of Smilodon gracihs from Leisey 1A in lateral view. (A) UF 81724, left mandible with c and p)-m 1; (13) UF 81723, right mandible with 13, c, and 1,4-m 1. Specimens coated for photography. Scale bars 20 mm in length. The scapula is not represented by any complete specimens; only the articular end is preserved. The lateral border of the glenoid fossa below the base of the spine is incised by a prominent notch, as in S. populator, that is lacking or very small in Panthera onca. Also, the glenoid fossa is narrow transversely at the cranial border (Table 13). As was true for previously known Florida samples of S. gracihs, the humerus is abundantly represented at Leisey. In most dimensions this element is larger than the humeri in the Inglis lA sample, although considerably smaller than those of the McLeod sample (Table 14). The proximal end of the humerus is 482 BULLETIN FLORIDA MUSEUM NATURAL HISrORY VOL. 37, Fr. U, No. 14 Table 11. Measurements of mandibular rami and lower dentition of Smilodon. Smilodon gracilia FLORIDA Leisey 1 A Inglis IA' UF UF UF UF UF UF 81724 87258 81723 82529 81723 87241 Greatest depth. nange 55.6e 52.5e 52.8 Lcast depth ramus below diastema 28.2 26.2 27.8 29.0 29.3 28.1 26.4-29.3(3) Width ramus at diastema 11.0 11.5 11.6 11.3 11.5 11.6 11.0-11.7«3) Width ramus at ml 15.4 15.2 16.8 15.2 15.2 17.3 14.2-15.9(4) Mandibulartoothrow length 54.0 53.3 50.4e 54.3 54.5 54.5 48.6-51.9(4) Height. angle- coronoid 54.1 Imigth, symphysis- condyle 149.5e - - c length 10.2 10.5 10.0 9.30-11.3«2) width - - 7.5 7.le-8.3«2) p3 length 9.5 9.0 9.4 8.le- 12.0(4) width 5.0 - - 5.0 5.3 5.le-6.1(4) p4 length 19.0 18.7e 18.7 18.2-20.3(4) width 8.9 8.1 9.Oe 7.8-8.5(4) ml length 23.6 23.0 23.7 20.8-23.0(5) 10.7 10.0 9.4-10.0(5) characterized by an open channeled bicipital groove. The lateral profile is diagnostic of Smilodon in having the junction of the greater tuberosity and the anterior border of the shaft nearly orthogonal, whereas in true cats these surfaces form arcuate connections. The distal end is broad relative to total length (Fig. 7A); the ratio of length to distal width ranges from 3.5 to 4.0. By comparison, this same ratio approximates 3.2 in Smilodon populator, 3.2 in Megantereon cultridens, and 4.0 in Panthera tigris. The supracondyloid ridge displays several well developed muscle scars for the origin of extensors of the forelimb and manus. BERTA LEISEY SHELL PIT FOSSIL CARNIVORES 483 Table 11 Extended Smilodon gracilis Smilodon populator PENNSYL- CALI- FLORIDA VANIA FLORIDA FORNIA Bass Rancho El Point Port Aucilia Hog La Jobean Waterway 12 Kcm,edy' 1¢1-kneej River~ Creek~ Breas 50.7e-65.0(3) 26.4e 30.3-31.7«3) 29.6-30.*2) 30.8 28.4 27.3-38.7 11.7e 11.8-13.3«3) 15.5e 15.4©-17.3«3) 57.5-59.0«3) 48.3-60.9 55.5-67.3(2) 58.0-76.3 - 171-197(2) - - 178.3-230.0 7.Oe-11.2(3) 14.4 13.8 13.4e 13.0-16.6 7.2-7.6(3) 9.8 9.5 11.9e 9.7-12.2 12.Oe 10.6e- 11.5«2) 6.5-10.2 4.6e 4.Oe-4.3«2) - - - 5.7-7.1 19.3 - 20.0-20.*4) 23.0-27.0(2) 25.0 23.4 23.2-27.7 8.0 8.8 8.4-9.5(4) 10.8-12.8(2) 11.3 10.4 10.9-14.6 21.3 23.Oa 22.0-25.2(3) 27.5-30.0(2) 23.4 27.0 25.0-32.1 9.7 10.Oa 9.8-10.5(3) 12.9-14.5(2) 12.4 13.2 12.4-17.6 I Mca=ments follow~ Bsta (1987) 2 Meamucments folow#W Chwcha (1983). 34 Measurements fo[lowin*Kintdn (1965). Measurements folowing Webb (197*). 5 Measurements folowing Mmiam and Stock (1932). Except for its larger size, the ulna agrees closely with those previously described from Inglis lA and McLeod (Table 15). It is relatively short with a robust shaft and a broad distal end. In anterior view (Fig. 7B), a large ovate scar 484 BULLEnN FLORIDA MUSEUM NATURAL HISrORY VOL. 37, PT. II, No. 14 Table 12. Measurements of atlas and axis vertebrae and sacrum ofSmilodon. Greatest Greatest Anterior Posterior Greatest Allan Width Width Lr~*h H~g)It S. gracilis (~isey) UF 81725 53.6 - 56.7 23.1 UF 81726 54.2 52.0 55.4 25.4 UF 81829 52.5 54.5 59.9 S. populator (Rancho La Brea') OR (N=10) 72.7-88.5 70.1-88.4 69.846.0 46.0-56.7 Greatest Greatest Greatest Depth Araaior Posterior Axis Lmth Centrum Width Width Height S. gracilis (Lcisey) UF 67084 76.8 - 54.6 - 41.5 UF 82834 74.5 15.1 51.6 41.7 - UF 87255 75.0 18.1 51.3 39.4 37.9 UF 81507 73.0 - 52.8 UF 67085 - 55.6 S. populator (Rancho La Brea') OR ~4=10) 86.9-108.5 21.1-31.9 63.6-84.2 45.1-71.5 81.4-103.3 Greatest Depth Greatest Anterior Anteflor Sacrum I~ngth Width Centrum S. gracilis (Leisey) UF 63665 55.4 72.7 8.5 UF 65397 56.7 72.5 9.6 S. populator (Arredondoz) UF 2562 126.0 93.0 24.0 (Rancho La Bres') OR(N=10) 76.2-90.6 77.0-110.3 23.0-33.9 1 Meannments fo[lowi,41 Mel,iam and Stock (1932).2 MeaRuements followlig Kimtin (1965). BERTA LEISEY SHELL PIT FOSSIL CARNIVORES 485 Table 13. Measurements ofs¢apula and pelvis ofSmilodon. Greatest Width. Greatest Lmgth, Least Width. Scapula Glenoid Cavity Glenoid Cavity Neck S. gracilia (Leisey) UF 80183 29.5 40.3 40.6 UF 81715 30.7 41.7 45.0 UF 67083 30.9 39.4 45.0 UF 65074 - 40.5 - S. populator (Arredondot) UF 2562 - - 58.1 (Rancho La Breab OR (N=10) 40.8-57.9 67.047.1 55.1-76.4 Greatest Depth, Acetabulum Pelvis Lm*h Ilium Diameter S. gracills (Inisey) UF 81221 239.0 22.5 33.4 UF 67083 - 22.1 33.5 S. populator (Arredondot) UF 2562 300.Oa 70.0 45.5 (Rancho La Brea~) OR (N=10) 283a-368 73.8-94.0 44.3-34.8 1 Meamuements folowbig Kintdn (1965).2 Measurements foilowing Mmilm and Stock (1932) on the shaft provides an area of origin for carpal and digital flexor, M. flexor profundus digitorum. The broad olecranon process of this species resembles S populator as does the proximally and laterally directed coronoid process that is distinctly distally directed in tnie cats. The radius also resembles that of previously described samples. The short, slightly curved shaft is relauvely slender in comparison with S. populator (Fig. 7C- D; Table 15). The pelvis of S gracilis is known only from Leisey. Except for its smaller size and stender proportions, it is very similar to S populator (Table 13). The ilium differs from those of Pm:thera onca and other true cats in having a more prominent inferior iliac spine that is positioned fprther dorsally on the crest of the ilium creating a sharper ridge (Fig. 8). 486 BULLEIDI FLORIDA MUSEUM NATURAL HISrORY VOL. 37, Fr. U, No. 14 Table 14. Measurements of the humcrus ofSmilodon. Prox- Prox Shaft Shaft Distal Distal I~ast Imigth Depth Width Depth Width Width Depth S. gracilia Onsey) UF 87250 256 71.7 52.3 - - 41.4 67.7 23.7 UF 87249 235e 64.2 - 33.8 21.0 - - 20.6 UF 87277 238 65.5 53.0 32.7 21.9 38.3 65.3 19.7 UF 83381 233 68.0 55.2 31.0 22.1 37.7 63.5 20.6 UF 81727 257 71.6 55.4 31.6 23.5 51.9 69.5 21.8 UF 87307 245 69.5 52.9 33.5 24.1 37.5 70.1 22.1 UF 63659 232 66.6 46.7 28.0 20.5 38.8 65.7 21.2 UF 87278 238 69.8 58.3 33.6 23.5 42.5 69.1 22.0 UF 81153 236 66.6 51.3 28.8 19.9 38.1 66.4 21.1 (Inglis IA') OR 237(1) 68.5- 48.5- 31.8- 20.7- 38.3- 66.6- 20.5- 74.3(2) 55.3(2) 37.5(5) 24.7(5) 42.7(3) 76.1(3) 21.1(3) (McI,codi) OR 40.5- 25.6- 44.7(1) 41.7(2) 26.3(2) S. populator (Arredond02) UF 2562 307 94.6 67.3 29.0 96.9 (Rancho La Breab OR 309- 92.0- 75.4- 32.2- 98.7- 385 118.2 92.4 41.7 128.3 ~ Measurements folowig Berta (1987).2 Measurements fonowig Kw·tdn (1965).3 Measurements folovving Merriam and Stock (1932). In the only known patella of S. gracilis from Leisey, the distal end is prolonged owing to ossification of a small portion of the patellar ligament. In this specimen, UF 80917, the greatest proximodistal diameter of this element measures 23.2 mm, and the greatest transverse width measures 36.7 mm. By contrast, a patella from McLeod is considerably deeper proximodistally, these dimensions measure 41.9 mm and 34.8 mm, respectively. The femur from Leisey is very similar to that from Inglis lA (Table 16). This element is shorter and more slender at both ends than that of S. populator (Fig. 9A). Several structural peculiarities of the proximal end distinguish S gracilis from P. onca. In S. graci/is a tuberosity is apparent on the anterior rim of the BERTA: LEISEY SHELL PIT FOSSIL CARNIVORES 487 A BCD , 9%. . k i"it Figure 7. Forelimb elements of Sm,lodon gracilis from Leisey lA (A) UF 81727, right humerus in anterior view; (B) UF 81728, left ulna in anterior view, (C) antcrior and (D) posterior views of UF 81730, right radius. Scale bar 30 mm in length. intertrochanteric fossa; P. onca lacks this tuberosity. The proximal end of the femur in S. gracihs is further characterized by a more medial position of the lesser trochanter. In distal view, the intercondylar notch is more deeply incised than that of P. onca. 488 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, Fr. II, No. 14 Table 1 5. Measurements ofthe ulnae and radii of Smilodon. 0!e- Ole- Sig- Coro- cranon cranon moid noid Process Process Notch Process ShaR ShaR Distal Ulna L,gth Width Width Width Width Width S. gracilis (Leisey) UF 85231 237 27.5 30.7 27.8 45.5 28.2 14.4 24.6 UF 81728 - 26.9 39.2 51.4 29.5 17.4 - UF 63660 - 36.1 - - - UF 83429 24.9 31.8 32.4 47.5 - - 26.0 33.2 29.5 47.6 28.1 16.4 (Inglis IA') UF 18109 - 21.8 33.0 26.5 41.5 42.6 12.7 27.9 (Mci.codi) F:AM 95324 26.1 31.6 29.7 50.2 47.3 13.8 30.5 S. populator (Arredon,102) UF 2562 310.3 46.7 63.2 (Rancho & Breab OR (N=10) 287- 41.5 57.3 19.1- 372 -60.2 -78.8 33.8 Prox. Prox. Disial Distal Long Shod Shaft Shaft Long Shon Radius I.cngth Diameter Diameter Width Depth Diameter Diameter S. gracilis (Le~sey') UF 81730 185 26.5 20.3 22.1 13.1 40.1 26.3 UF 81729 185 26.3 20.4 21.9 12.9 39.3 25.4 UF 81371 209 29.0 22.6 23.2 12.0 41.3 26.1 (MCLeodl) OR 35.2(1) 27.8(1) 23.6- 15.4- 45.6- 31.3- 26.*2) 16.1(2) 49.7(2) 32.0(2) S. populator (Arredond02 5 UF 2562 240e 42.8 33.2 26.5 18.5 (Randu IA Breab OR(N=10) 235- 41.3- 32.2- 26.0- 16.5- 295 55.5 44.0 38.8 24.6 ' Mess=nents foOowing Berta(1987). 2 M . Mowi~Klutdn (1965) ) Measurements folkming Me:*am and Stock (1932). 490 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, Fr. II, No. 14 1 . - 3441/2491/,3./Wil 1--Ii 4 M' .AY:~.,1,~3 Figure 8. Lateral view of UF 81221, right innominale of Smilodon gracilis from Lkisey 1 A. Scale bar 30 mm in length. A B ; f t. t :1 Figure 9. Hindlimb elements of Smilodon gracilis from Leisey lk (A) UF 67081, right femur in anterior veir (B) UF 81731, lefttibia in anterior view. Scale bar 50 mm in length. BERTA LEISEY SHELL PIT FOSSIL CARNIVORES 489 Table 16. Measurements of'f*mur, tibia, and libula of Sm,lodon. Prox Prox. ShaR ShaR Distal Distal Weighti I.mgth Width Depth Width Depth Width Depth (kg) Femur S. gracilis Missy) UF 67081 279 65.7 - 23.7 26.0 54.4 50.2 87 UF 82652 274 63.8 - 20.5 23.4 53.5 53.3 67 (Inglis IA) UF 18112 - 67.4 38.0 31.7 28.4 - - 111 S. populator (Arredondo) UF 2562 334 86.0 - 30.5 28.0 71.5 66.5 - (Rancho La Brea) OR (N=10) 317- 82.7- - 30.1- 26.8- 65.2- 63.9- 155 408 108.8 - 40.5 35.4 90.2 80.3 Tibia S. gracilia O.eisey) UF 81731 226 56.6 57.3 20.8 24.2 40.5 28.0 UF 81732 - 54.2 56.0 - - - - (Haile XVA) UF 17496 - - - 22.6 20.2 41.9e 29.5 94(Leod) F:.AM 95329 - - - 20.8 33.0 44.3 32.8 S. populator (Arredondo) UF 2562 252 77.4 - 24.5 - 51.7 - (Rancho La Brea) OR (N=10) 239- 72.5- - 25.1- - 45.0- - 305 90.4 33.0 63.3 Flbula S. gracilis gEEey) UF 67088 - - - - 21.5 13.4 S. populator (Rancho La Brea) OR (N=10) 212.7- 32.5- - 9.4- 9.2- - 17.2- 265.0 45.9 15.3 11.9 26.8 1 Calculations bascd on Shaw and TejadD-Floms (1985: Table 2) BERTA: LEISEY SHELL Prr FOSSILCARNIVORES 491 Table 17. Measurements ofmetacarpal 4 III, and IV of Smilodon. Prox: Prox ShaR ShaR Distal Distal I.ength Width Depth Width Depth Width Depth Metacarpal II S. gracilis (Leisey) UF 86752 67.5 18.7 18.8 10.7 10.8 16.2 15.6 (Haile XVA) 74.9 18.8 22.0 13.3 13.0 21.0 18.2 (Inglis IA) 75.0 18.8 22.3 12.0 11.4 17.9 17.2 (Port Kennedy) 70.9 15.9- 19.3- 11.9 11.8 18.7 13.0 OR 17.5(2) 22.9(2) Metacm,al m S. gracilis (Leisey) UF 87269 82.2 18.6 - - - - - (Port Kennedy) - 20.9 18.9 10.6 10.1 17.5 15.4 - - 20.6 - - - - S. populator (Reddick) UF 2537 100.0 26.0 24.6 17.4 - 25.6 - (Rancho La Brea) OR (N=77) 83.0- 24.2- 22.8- 14.5- - 22.4- - 110.0 30.7 29.2 20.0 29.4 Metacarpal IV S. gracilis G.cisey) UF 86747 71.2 14.7 15.4 9.5 10.9 14.8 15.6 (Inglis IA) 77.5 16.7 19.4 10.7 10.2 16.8 16.8 (McLeod) 83.7 19.4 20.6 11.8 - 20.1 18.0 S. populator (Reddick) UF 2537 93.0 23.2 23.8 14.0 - 22.4 - (Rancho La Brea) OR (N=733) 79.0- 18.9- 20.6- 12.3- - 19.6- - 107.0 26.6 27.4 16.0 24.6 The tibia of this species at Leisey is complete. The proximal end differs from that of S populator in the sharper projection of the posterior border of the lateral condyle (Fig. 9B). In posterior view, the distal end exhibits a stepped posterior m=rgin with well developed grooves for Mm. flexor hallucis longus, flexor digitorum longus, and tibialis caudalis. 492 BULLEIIN FLORIDA MUSEUM NATURAL HISTORY VOL. 37, Fr. II, No. 14 Table 18. Measurements ofmetatarsal IV and V of Smilodon. Prox. Prox ShaR ShaR Distal Distal Ikagth Width Depth Width Depth Width Depth Me¢atarsal IV S. gracilis O.Emey) UF 81712 88.2 15.8 19.9 10.3 12.0 15.1 14.2 UF 87273 88.4 15.0 19.1 9.5 11.0 14.7 15.3 (Inglis IA) 89.5 17.6 21.8 11.2 12.3 16.3 15.5 (McLeod) 91.6 20.3 24.7 11.6- 12.5- 17.9- 16.0- 13.*2) 14.5(2) 18.0(2) 16.8(2) S. populator (Reddick) UF 2537 101.0 - 28.0 15.6 - 21.5 - (Arredondo) UF 2562 96.0 - 26.3 14.1 20.2 - (Rancho La Brea) OR (N=748) 84.0- 16.1- 25.8- 13.0- - 18.5 - 114.0 19.9 17.0 25.2 - 25.2 - Me~arsal V S. gracilia (Leisey) UF 81709 77.4 15.6 13.2 8.1 8.5 8.5 13.0 (McI,cod) 79.4 23.7 18.5 9.9 11.7 15.0 15.9 Spopulator (Arredondo) UF 2562 78.0 - 24.5 12.1 - 17.1 - (Rancho La Brea) OR (N=722) 71.0- - 24.5- 10.5- - 17.1 - 95.0 28.6 13.8 21.6 The fibula, previously unknown for this species is similar to that of S populator but smaller (Table 16). It differs from that of P. onca in having a less prominent projection of the lateral malleolus. The metatarsals, like the metacarpals, are relatively short relative to total limb lengths (Tables 17, 18). The metatarsals are longer and more slenderly proportioned than the metacarpals, as was previously noted for this species (Berta 1987). Metacarpal n differs from Inglis lA and Haile-15A specimens in lacking a prominent projection of the facet for the trapezium above the articular surface; BERTA LEISEY SHELL PIT FOSSIL CARNIVORES 493 however, it shows a Smi/odon-like sharp depression in this surface. Panthera onca differs in having a more gently concave articular surface. Comparison of limb lengths of S gracilis from Leisey indicate that this sample was intermediate in size between Inglis lA and McLeod/Port Kennedy umples (Tables 14-18). The limbs of S gracilis and S. populator were similarly proportioned (Table 19) and support the interpretation of Smi/odon as a short limbed, robust machairodont with especially powerful forelimbs. Body mass calculations (see Shaw and Tejada-Flores 1985) indicate that S gracihs averaged one-third smaller than S. populator. Discussion.- Smilodon gracilis is by far the most abundantly represented carnivore at Leisey. In addition, the Leisey sample is the largest and best known record of the gracile sabercat. Cope's (1880, 1895, 1899) original description of S gradlis was based on fragmentary material from Port Kennedy Cave (middle Irvingtonian fide Lundelius et al. 1987). This species is much better represented from Florida faunas and has been documented from the late Blancan Haile 154 Santa Fe River 14 Bass Point Waterway and El Jobean faunas, and the Irvingtonian Inglis lA and McLeod faunas (Berta 1987; Churcher 1984). The Leisey and Haile 21A occurrences now can be added to these Irvingtonian records. The gracile sabercat from Leisey shows several characters not seen in the very early Ilvingtonian Inglis lA sample-larger size, typically a single-rooted p3, and reduced p3. The later Irvingtonian occurrences of S gracilis from Port Kennedy Cave and McLeod show most significantly further increase in size. Based on the preceding discussion, the stage of evolution of S. gracilis at Leisey is intermediate between that of Inglis lA and Port Kennedy/McLeod specimens, and supports a late early Irvingtonian age for the Leisey fauna. Homotherium sp. Referred Specimens.- Ikisey 1: UF 95764, right I2. Leisey IA: UF 81738, deciduous incisor, 81733, left ml; 84841, distal right humerus; 63662, left femur. Discussion.- Machairodont cats are in need of revision. Because of the uncertainty regarding machairodont taxonomy (e.g., see Berta and Galiano 1983), formal diagnosis of these few specimens from Leisey must await description of a larger, better preserved sample (skull and skeleton) of this same cat from Haile 21A. The deciduous incisor has a compressed crown and a single basal cuspule erupted on the lingual tooth margin. Serrations are present along the mesial and 494 BULLErIN FLORIDA MUSEUM NATURAL HaTORY VOL. 37, PT. Il, No. 14 Table 19. Limb proportions in Sm,lodon (based on mean values for lengths oflimb elements in mm) S. gracilia & populatorl Humerus 244.5(8) 344.9(10) Radius 194.0(3) 265.7(10) Metacarpal U 72.1(4) 90.1(676) Formjr 276.5(2) 367.7(10) Tibia 226.0 273.6(10) Metatarsal n 89.4(4) 99.0(766) Humerus X 100/H+R+Mic 47.9 49.2 Radius X 100/H+R+Mtc 38.0 37.9 Mtc X 100/H+R+Mtc 14.1 12.8 Fcmur X 100/F+T+Mtt 46.7 49.7 Tibia X 100/F+T+Mtt 38.2 37.0 Mt X 100/F+T+Mtt 15.1 13.4 1 M ' 6[lowhkg Mezriam and Stock (1932) distal margins. The tip of the crown and serrations on the distal side are worn away. The root is partially formed and the apical root foramen is open. This specimen indicates the presence of a very young juvenile at Leisey. Comparisons with juvenile Dinobastis from Friesenhahn Cave described by Rawn-Schatzinger (1983) suggests that this individual represents Stage II ofthe eruption sequence. An isolated I2 referred to this genus compares favorably in size and morphology with the Haile 21A specimen (Table 20). Prominent basal cusps with serrated medial and lateral margins are positioned on either side of the central cusp. The lower carnassial, larger than Reddick lA or Friesenhahn machairodonts, compares best with the Haile 21A homothere. The tooth enamel is crenulate, and coarse serrations are present along the sides of the paraconid and protoconid blades (Fig. 1OB). Referral of a distal humerus to this genus is suggested by its similarity to the Haile 21A machairodont and is distinct from Smilodon, the only other large felid represented at Leisey. This element differs from Smilodon in being larger and more robust. In addition, the shaft at the level of the entepicondylar foramen is broader and natter mediolaterally, the supracondylar ridge is not as strongly developed, and the distal end is narrower posteromedially (Table 21). A femur also referred to Homotherium from Leisey, although considerably smaller, agrees well with the Haile 21A homothere. It differs from Smilodon in numerous features, including lack of a prominent tuberosity between the head and BERTA. LEISEY SHELL PIT FOSSIL CARNIVORES 495 A A "<4 C D 0 6 6 6 1 45/Y 3 1 2 4 1 7 2 Figure 10. Homothenum and Miracinon}x from the Leisey Shell Pits. (A-B) Homothenum sp., (C-D) Miracinony~ inexpectatus. (A) UF 63662, left femur in interior view, (B) UF 81733, leR ml in lateral view; (C) anterior and (D) posterior views of UF 124172, right metatarsal II. Scale bar 35 mm in length for 4 20 mm for B, and 30 mm for C-D. 496 BULLETIN FLORIDA MUSEUM NATURAL HISrORY VOL. 37, Fr. II, No. 14 Table 20. C „. and related felids Lenglh Width 4cd Lead m Homotherium sp. Leiscy 13.6 12.6 Dinobaatis serus Friesenhahn' 11.5-12.0 ml Homotherium sp. Leisey 33.9 14.4 16.7e 12.4 Haile 21 A 34.1 15.7 15.2 13.5 Dinobastis serus Reddick 1A2 28.5 11.1 15.0 Friesenhahn 25.6-30.6 10.2-12.8 13.6-15.4 l M 'BilowMeade(1961). 2 Measurements foUow Waldrop (1974 Lpcd=length ofpxotoconid. i.pa*length of paracomid- Table 21. r ~ . and related feli(Is. Prox. Prox. ShaR ShaR Distal ~ngth Width Depth Width Depth Width Humcrus Homotherium sp. LANy - - 75.0 28.5 Haile 21A 361 82 106 105.0 Inglis lA 302c - 91 84.0 - Dinobastis serus 340-358 72-76 92-104 83-84 30-31 Friesenhahn (OR. N =4) Femur Homotherium sp Leisey 318 79.0 28.4 26.0 64.5 65.5 Haile 21A 381 104.0 - - 83.0 - Dinobastis semi Friesenhaim 338-353 89.2-95.7 41.0-42.8 30.0-31.3 27.3-28.8 (OR, N = B BERTA LEISEY SHELL Prr FOSSIL CARNIVORES 497 greater trochanter lesser trochanter more medial in position; intertrochanteric fossa larger anteroposteriorly, forming a broad shelf that extends from the lesser trochanter to the fossa; and finally, distal condyles situated closer together (Fig. 10A). Prior to this record, Homotherium was reported from Florida and for the first time from eastern North America from the early Irvingtonian Inglis lA site (Webb 1974a) and from the Rancholabrean Reddick lA site (Waldrop 1974). Identity of the Reddick lA machairodont, of which more material is now available, as Homotherium rather than Dinobastis is questioned here and must await further systematic study. In any case, although comparable elements are not available for the I.eisey sample in size and morphology, this same machairodont from Haile 21A can be ren,lily distinguished from the Reddick lA cat in having longer upper cnnines and P4 with a distinct protocone supported by a root Miracinonyx inexpectatus (Cope 1895) Referred Specimen.- Leisey 3: UF 124172, right metatarsal IL Discussion.- The late Blancan and Irvingtonian cheetah-like cat Mira- cinonyx inexpectatus only very recently has been reported from Florida by Van Valkenburgh et at (1990), who identified a mandible, metatarsal IV, and several additional unspecified postcranial elements from Inglis lA. The Inglis sample also includes a metatarsal n (UF 45471) that compares very closely in size and morphological features with the metatarsal II from Leisey (Fig. 10C-D). Both the Leisey and Inglis metatarsals are referred to M. inexpectatus based on their large size and striking elongation. Measurements of the Leisey specimen are very similar to those of a metatarsal II of M inexpectatus (Table 22) from the middle Irvingtonian Hamilton Cave, West Virginia (Van Valkenburgh et al. 1990). Table 22. Measurements of the metatarsal H of Miracinon,x inapectatus from Lcisey Shell Pit. Inglis 14 and Hamilton Cave. Abbreviations and measurements for Hamilton Cave specimen are from Van Valkenburgh et al. (1990) Leisey Ingliq Hamilton Cave UF 124172 UF 45471 USNM 401092 Total length (L) 120.6 112.0 120.1 Proximal breadth (BP) 15.7 14.8 12.5 Proximal depth (APP) 23.0 22.8 22.3 MidshaR diameter (BS) 11.8 11.7 10.3 Distal breadth (BD) 17.8 17.6 18.4 498 BULLEr[N FLORIDA MUSEUM NATURAL HISrORY VOL. 37, Fr. Il, No. 14 Family PHOCIDAE Gray 1825 Monachus tropicalis (Gray 1850) Referred Specimena- Leisey IA: UF 81734, right upper first postcanine tooth; 80499, p3. Discussion.- The upper postcanine tooth agrees in detail with a specimen of the modern species Monachus tropicalis from a Florida archaeological site, (Malabar, South Indian Field. Brevard Co.). Measurements for UF 81734 are: greatest length, 11.75 mm, greatest width, 8.3 mm. A second tooth, a well worn and poorly preserved right p3 is tentatively referred to this species. The only other well documented Florida Irvingtonian occurrence of this species is a fragmentary left mandible with m2 from Rigby Shell Pit (UF 36456). SIJMMARY The Leisey carnivores are among the most diverse Florida Irvingtonian assemblages known, second only to the Coleman 2A fauna. Thirteen species represent a minimum number of 36 individuals. The largest, most complete sample of the sabercat Smi/odon graci/is is from this locality and adds considerably to our knowledge of the species. The Leisey sample provides evidence for the gradual increase in size of S gracilis during the Irvingtonian. Based on the stage of evolution of this species an age intermediate between Inglis lA and McLeod is supported for the Leisey fauna. Another machairodont cat, Homotherium, less well known from Florida, is also recorded from this fauna. A large and a small wolf, Canis armbrusteri and C edwardii respectively, as well as the tremarctine bear Arctodus pristinus are additional large carnivores represented at Leisey. Canis edwardii, the second most abundant carnivore, is represented by skeletal material as well as fragmentary maxillae and mandibles. In addition to the above mentioned large terrestrial carnivores, a single pinniped is recorded, the Caribbean Monk Seal Monachus tropicalis. Among the small carnivores recorded from Leisey, the occurrence of the otter Lutra canadensis is significant as it represents the first Irvingtonian record from Florida of this species. Other small carnivores recorded include the fox Urocyon cineroargenteus, 1he raccoon Procyon lotor, the long-tailed weasel Mustela frenata, the spoted *urk Spilogale putorius, andthebobcat Lynx rufus. BERTA: LEISEY SHELL PIT FOSSIL CARNIVORES 499 LITERATURE CITED Be,14 A 1981. Fossil wolves, coyotes and do/ of Florids Plaster Jacket 36:8-24. 1985. The status of Smilodon in North and South Ameria Contrib. Sci. Nat. HiaL Mus. Los Angeles Co. 370:1-15. 1987. ne sabercat Smilodon gracm, Som Florida and a discussion of its relationships (Mammalia, Felidae, Smilodontini). Bull. Florida State Mus., Biol. Sci. 31(1):1-63. . and H. Galiano. 1983. Megantereon hespenu from the late Hemphillian of Florida with remarks on the phylogenetic relationships of machairodoets (Mammalia. Felidae, Machairodontinae). J. Palma 57(5):892-899. Churcher, C.S. 1984. The stalus of Smitodon,opsis (Bro~ 1908) and Ischyrosmilus (Merriam 1918) Life Sci. Contrib. 140, Royal Ontario Museum. Toronto. 59 p. Cope, E. 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Cenozoic Mammals of North America. Geochronology and Biostratigraphy. Univ. Califbrnia Press, Berkeley. Martin. R. A 1974. Fossil mammals from the Coleman IIA Fauna, Sumter County. Pp. 35-99 in S.D. Webb, ed. Pleistocene mammals of Florida. Univ. Presses Florida, Gainesville. Merriam, J. C., and C. Stock. 1932. The Felidae of Rancho La Brea. Carnegie Inst Washington Publ. 422:1-231. Rawn-Schaizinger, V. 1983. Development and cruption sequence of deciduous and pennanent teeth in the saber-tooth cat Homotherium,erum Cope. J. Vert. Paleon. 3(1):49-57. Schaub, S. 1925. Ucber die osteologie von Machaerodus cultridens Cuvier. Ecologae Geol. Helveliae 19:255-266. Shaw, C. A, and A E. Tejada-Flores. 1985. Biomechanical implications of the variation in Smilodon ectocunciforms from Rancho La Brea. Contrib. Sci. Nat. Hist Mus. Las Angeles Co. 359:1-8. Van Valkenburgh, B., F. Grady, and B. Kurth 1990. The Plio-Pleistocene cheetah-like cat Miracinonyx inapectatus of Nodh Americt J. Vert Paleon. 10:434-454. Waldrop, J.S. 1974. The scimitarcal. Homotherium serum, from the Florida late Pleislocene. Pp. 154-157 in S.D. Webb, ed. Pleistocene Mammals ofFlorida. Univ. Presses Florida, Gainesville. Webb, S.D. 19741 Chronology of Horida Pleistocene mammals. Pp. 5-34 m S.D. Webb, ed. Pleistocene Mammals of Florida. Univ. Presses Florida, Gainesville. 1974b. The stallis of Smitodon in the Florida Pleistocene. Pp. 149-133 in S.D. Webb, ed. Pleistocene Mammals of Florida. Univ. Presses Florida, Gainesville. Werdertin, L 1985. Small Pleigtocene felines ofNorth America J. Vert Paleon. 5(3):194-210.