· 90**J__il"Ill"Illitia - 942~1«vi-+Wer- %4"-*-151*86.226 T41. : - 6...6 &- of the FLORIDA STATE MUSEUM Biological Sciences Volume 33 1988 Number 5 CORMOHIPPARION AND HIPPARION (MAMMALIA, PERISSODACTYLA, EQUIDAE) FROM THE LATE NEOGENE OF FLORIDA Richard C. Hulbert, Jr. - _ 23 1 1 , 1 ~ *~-2 -3-.-- Ez -_- 9 - : ill 8. 1 =r- = -5 0.33»lYS Ifia d ~TWHk#14»- ffi! liif-;ss]=_cj~~~*I.% UNIVERSITY OF FLORIDA GAINESVILLE Numbers of the BULLETIN OF THE FLORIDA STATE MUSEUM, BIOLOGICAL SCIENCES, are published at irregular intervals. Volumes contain about 300 pages and are not necessarily completed in any one calendar year. S. DAVID WEBB, Editor OLIVER L. AUSTIN, JR., Editor Emerims RHODA J. BRYANr, Managing Editor Communications concerning'purchase or exchange of the publications and all manuscripts should be addressed to: Managing Editor, Bulletin; Florida State Museum; University of Florida; Gainesville FL 32611; U.SA This public document was promulgated at an annual cost of $3626.50 or $3.627 per copy. It makes available to libraries, scholars, and all interested persons the results of researches in the natural sciences, emphasizing the circum- Caribbean region. ISSN: 0071-6154 CODEN: BF 5BAS Publication date: October 20,1988 Price: $3.75 CORMOHIPPARION AND HIPPARION (NLAMMALIA PERISSODACTYLA, EQUIDAE) FROM THE LATE NEOGENE OF FLORIDA Richard C. Hulbert, Jr.* ABSTRACT Five species of Cormohipparion are recognized from the late middle Miocene to late Pliocene (late Barstovian through Blancan) of Florida. A small sample of isolated teeth collected from a late Barstovian horizon of the lower Bone Valley Formation, south-central Florida, represents Cormohipparion sp., cf. C sphenodus. In the Clarendonian, a larger species of Cormohipparion, cf. C occidentale, is represented by three isolated teeth, also from an older horizon of the Bone Valley Formation. The very late Clarendonian Love Site produced numerous dentitions of two species of Com:ohippan'on, one small-sized and one medium-sized, here referred to C ingenuum and C plicatile, respectively. The completeness of the referred material allows for new phylogenetic interpretations concerning these previously poorly known species described by Joseph Leidy over a century ago on isolated teeth. The first appearance of C ingenuum was in the early Clarendonian of Texas and Florida, and it persisted in Florida until the late early Hemphillian, a duration of about five million years. It is also recorded from the early Hemphillian Gracias Fauna of Honduras. C ingenuum is not the senior synonym of Nannippus lenticularis (Cope), as proposed by some authors, as the two share few characters but size. C plicatile is limited to the late Clarendonian-early Hemphillian of Florida, but is usually the more common of the two at sites where both occur. The range 6f C. emsliei is extended to the early Hemphillian (ca. 7 Ma), based on its presence at the Moss Acres Racetrack Site. It persisted through the Blancan (to about 2.0 Ma), and is the youngest known representative of the genus in North America. These three species form a monophyletic gr6up, Cormohippan'on (Notiocradohippation) n subgen., united by their relatively very elongated muzzles. This clade was apparently limited to the Gulf Coastal Plain and Central America. Three major trends observed in Florida Cormohipparion over this ten million year period are increasing hypsodonty, increasing enamel complexity (in fossette plications, pli caballins, styles, and pli caballinids), and decreasing depth of the d6rsal preorbital fossa. Hipparion is rarely recorded from the late Miocene of Florida. Referable samples include those from the lower Bone Valley Formation (?Clarendonian), the Love Site, the Moss Acres Racetrack Site, and the Withlacoochee River 4A local fauna (late early Hemphillian). The latter two are among the youngest records of H*arion from North America. They are all tentatively referred to the medium-sized species, Hipparion tehonense. * The author is a Postdoctoral Research Fellow at the Florida State Museum, Universityof Florida, Gainesville FL 32611. HULBERT, AC.,JR. 1988. Connohippan'on and H*arion (Mammalia, Perissodactyla, Equidae) from the late Neogene of Florida. Bull. Florida State Mus., Biol. Sci. 33(5):229-338. 230 BULLEI'IN FLORIDA SrATE MUSEUM VOL. 33(5) Cladistic analysis based on 58 dental and cranial characters suggests the follawing phylogenetic hypotheses: (1) the Tribe Hipparionini is monophyletic and contains a minimum of six North American genera and several species of uncertain generic affiliation; (2) Hippanon, Me,ychippus s.s., Nannippus, Connohippanon, and Old World hipparionines form one monophyletic clade, the Hippanon-genus group; (3) Neoh (Us~ fs . IP ..f'k 'f - =b <0\ .2-4 i , 1... " 5" 75\ D 4 -'. 1cm r 9/ --i,-/ 3 Imth ts £*.Clk:'J,t#2$*~1 E -<7 A- irlf/ LIZIAW , Ch- ./77. , ' i . i , \'660 'li.A -U --3E3 FIGURE 10. Occlusal views of lower cheekteeth 6f Cormoh*arion ingenuum from the Agricola Road Site, Hookers Prairie Mine (early Clarendonian), Polk County, Florida. A. UF 98126, associated R p2-m3. B. UF 98129, associated L p2-p4. C. UF 98136/98145, probably associated R dp4-ml. D. 98146, R m12. E. UF 98154, R m12. HULBERT: CORMORHIPPARION & HIPPARION FROM FLORIDA 257 gramm without comment, and Lucas (in Leidy and Lucas 1896) and Hay (1902) followed this synonymy. Pseudh*anon gramm is now known to differ greatly from "H." ingenuum in size, facial morpholgy, and enamel pattern (Webb 1969a; Webb and Hulbert 1986). Leidrs (1885) referral of ingenuum to the genus Hippothen'um reflected no particular phylogenetic significance. During this period, all hipparionine and many merychippine species were placed in Hippotherium, a name then considered valid for most if not all Eurasian and North American hipparionines (e.g. Cope 1889). Starting about 1900, Hippanon replaced Hppothedum as the commonly used genus for hipparionine equids (e.g. Hay 1902). Gidley (1907) resurrected Leidys species from Cope's synonymy, and it has subsequently been considered a valid species. Gidleys use of the binomen "Hipparion" ingenuum did have phylogenetic significance, however, as he felt that "H." ingenuum (along with "H." plicatile and 'H." venusmm) "...belong to an American branch of the Hippan'on group of the Old World" (Gidley 1907:906). Gidlers separation of hipparionines into two genera, Neohippan'on (for New World species) and Hippan'on (primarily for Old World species), was not accepted by his contemporaries Matthew and Osborn (see e.g. Osborn 1918:173), who continued to refer all hipparionines to a single, broadly defined genus. Matthew (1924) concluded that the Miocene hipparionines named by Leidy from Florida and South Carolina (including "H." ingenuum) were not especially related to Old World species, a view also endorsed by Simpson (1930). Matthew and Stirton (1930:354) and Simpson (1930:188) placed "H." ingenuum in Matthew's (1926) subgenus Nanni»us. Stirton (1940) raised Nannippus to the status of genus, and included "H." ingenuum in his listing of species of Nannippus. None of these authors stated any specific reasons for this referral, but presumably it was based on the relatively small size of the holotype (Table 1). Most subsequent references to the species have been to "Nannippus" ingenuus (e.g. Quinn 1955; Forst6n 1975; MacFadden 1984). Large samples of Clarendonian and early Hemphiman equids are now known from Florida. Four species of medium- to large-sized hipparionines (UTRL between 115 and 140 mm) are present; as the oldest available name, Connoh*anon ingenuum should apply to one of them. Two, although of similar size as the holotype of C ingenuum, are distinguished from it primarily on qualitative grounds. One, referred to Neohippan'on trampasense by MacFadden (1984) and Hulbert (1987b), differs in its more elongated protocone, narrower fossettes, and stronger metastyle. Hipparion sp., cf. H. tehonense, described below, differs in its rounded lingual protocone margin, simpler fossettes, and weaker pli caballin. The holotype of C ingenuum (and that of C plicatile) falls within the observed range of enamel morphology of the two remaining populations, which are very similar to each other and are distinguished primarily by size. That the two do in fact represent distinct species is emphasized by differences in DPOF morphology, diastema length, p2 258 BULLETIN FLORIDA SrATE MUSEUM VOL. 33(5) A 18.0 19.5 21.0 B 19.5 21.0 22.5 24.0 C u. 111 lilli -r-, ....'',1.'....1 16.5 18.0 19.5 21.0 rfff rn rn r. .. 19.5 21.0 22.5 24.0 E /~f ./ «, 13.5 15.0 16.5 10-F 5- , 16.5 18.0 19.5 FIGURE 11. Histograms of the distribution of occlusal anteroposterior length (APL), transverse width (TRW), and basal crown length (BAPL) for combined Florida samples of upper third and fourth premolars (P34) of Cormohipparion ingenuum (4 C, and E) and C. plicatile (B, D, and F). Analized samples exclude very slightly worn and very heavily worn individuals. A-B. P34 APL; n = 53 for C. ingenuum, n = 121 for C plicatile (similarly, the first value given in the following for sample size is that of C ingenuum, the second C plicatile). C-D. P34 TRW; n = 53, 123. E-F. P34 BAPL; n = 48,121. Distributions are interpretted as being primarily bimodal with varying degrees of overlap between the two species. All measurements in millimeters. HULBERT: CORMORHIPPARION & HIPPARION FROM FLORIDA 259 morphology, and unworn crown height. However, the only significant difference between isolated, worn upper cheekteeth is size. For Connohipparion ingenuum and C plicatile to both be valid, distinct species, it is necessary to show: (1) that two different-sized populations of Cormohipparion exist at Mixson's Bone Bed and Other nearby localities; (2) that the holotype of C ingenuum uniquely matches upper cheekteeth of the smaller-sized population in size and morphology; and (3) that the holotype of C plicatile matches upper cheekteeth of the larger-sized population. If only a single population exists, or if the two match with the same population, then C plicatile must be considered a junior synonym of the older name, C ingenuum. If one (or both) can be shown to be specifically undiagnostic by falling within the observed range of variation of more than a single species, then it should be considered a nomen dubium. Leidy and all subsequent authors have correctly identified the type of C ingenuum (USNM 3306, Fig. 1) as a true molar. However, since its initial description, the holotype tooth of C plicatile (USNM 3292) has also been described as a "molar" (Leidy 1887:310;-Osborn 1918:192; MacFadden 1984:170). Leidy used the term to refer to both true molars and premolars. Leidy and Lucas (1896:50) described USNM 3292 as a molar, "...probably the third...," presumably a reference to a P4. There are several features of the specimen that indicate that it is indeed a premolar, and not a molar. These include the relatively large and widely open parastyle and mesostyle, and the complex pli caballin. Most importantly, the mesostyle angles anteriorly in labial view when the occlusal surface is face down, a distinguishing characteristic of P345 and not M125 (Bode 1931). That the holotype of C plicatile is a premolar is important, because the size difference between the holotypes of C plicatile and C ingenuum (Table 1) might merely reflect the typical significant size differences between P345 and M125 of Connoh*pation. The range in size and CV of the combined sample are greater than that observed in other single-species quarry or faunal populations (Table 6), strongly suggesting the. presence of more than a single species. Frequency distributions of occlusal and basal crown dimensions (Figs. 11, 12) are bimodal, but do indicate an appreciable amount of overlap between the two populations. Much of the apparent overlap in APL is lessened when the data are adjusted for crown height. The Mahalanobis distances between the two populations are significantly different (at p < 0.01) for both of the separately analyzed samples of P34 and M12. The holotype of C ingenuum clearly clusters with the molars of the smaller-sized population both in univariate and multivariate analyses (Tables 1,3; Figs. 11-13), indicating that the smaller sample should be referred to the older.name. The ho16type of C plicatile, while within the range of only the larger-sized population for TRW and BAPL (Tables 1,3; Fig. 11D, 11F), has a relatively small APL (19.7 mm) that is within the observed range of both populations (Table 3; Fig. 11B). Linear discriminint analysis separates tlie two species with very little overlap 260 BULLETIN FLORIDA STATE MUSEUM VOL. 33(5) A r-1 16.5 18.0 19.5 18.0 19.5 21.0 22.5 C r-' 15.0 16.5 18.0 D 18.0 19.5 21.0 E 13.5 15.0 16.5 F to -; 5 -3 r E rTH...., 15.0 16.5 18.0 FIGURE 12. Histograms depicting the distribution of occlusal anteroposterior length (APL), transverse width (TRW), and basal crown length (BAPL) for combined Florida samples of first and second molars (M12) of Connohippmion ingenuum (A, C, and E) and C. plicatite (B, D, and F). Analized samples exclude very slightly worn and very heavily worn individuals. A-B. M12 APL; n = 52,110. C-D. M12 TRW; n = 53, 105. E-F. M12 BAPL; n = 56,120. Distributions are interpretted as being primarily bimodal with varying degrees of overlap between the two species. All measurements in millimeters. HULBERT: CORMORHIPPARION & HIPPARIONFROM FLORIDA 261 20- A - -t y ~11111,111 10- -4 -3 -2 -1 0 1 2 3 4 20- 8 - 10- 17-71 ~in -,/11 111 -5 -4 -3 -2 -1 0 1 2 3 4 FIGURE 13. Results of canonical discriminant analysis between Connoh*anon ingenman (shaded) and C. plicatile (not shaded) using four variables, APL, BAPL, TRW, and PRL. Variables standardized to x = 0 and s = 1 before analysis. Separate analyses performed for a sample of combined upper first and second molars (A) and a sample of upper third and fourth premolars (13). Sample sizes are, for C. ingenuum, 48 premolars and 55 molars; for C. p/icatile, 110 premolars and 107 molars. Histograms show distribution of canonical scores; actual results produced no overlap for the premolars, and only three molars were "misidentified" (i.e. discriminant analysis would assign them to the other species). The holotypes of both species were included in the analysis (their scores indicated by arrows), and each fell within the OR of its respective population. The results are consistent with there being two morphospecies of Cormoh*arion present in the late Miocene of Florida. and places USNM 3292 with the larger sample (Fig. 13). Thus, the link between USNM 3292 and the larger-sized population, while somewhat more uncertain than that of the type of C ingenuum and the smaller-sized population, is reasonable. Also, the best matches for the peculiar fossette morphology of USNM 3292 are found in the larger-sized population (e.g. UF 17211, Fig. 16B). The conditions for validation of both C ingenuum and C plicatile set forth above are thus apparently met, although the holotype of C plicatile is atypical for certain characters relative to the majority of specimens referred to that species. These are judged to reflect individual variation rather than species-level differences. Qualitative analyses of the dental material described above indicate that 'Hippothen'um" ingenuum is more closely related to C sphenodus and C occidentale (excluding other species from Florida) than to species from any other hipparionine genus from North America, and that it can be referred to Connoh*parion sensu MacFadden (1984). C ingenuum shares the following features with C sphenodus and C occidentale which in combination serve to distinguish them from other North American hipparionine genera: high, well developed ectostylids on deciduous lower premolars; lower permanent 262 BULLETIN FLORIDA STATE MUSEUM V0L. 33(5) premolars with moderately reduced ectoftexid depth and moderate pli caballinids in early wear-stages; high frequency of plicated isthmuses and paralophids; strong protostylids on p3-m3 and dp34; p2 and P2 much longer than p34 and P34 with expanded paraconids and anterostyles, respectively; oval or elongate-oval protocones isolated until late wear-stages; and highly complex and intricately plicated fossettes and pli caballins until at least middle wear-stages. While some of these character states evolved independently in other hipparionine genera, in combination they support the referral of ingenuum to Connohipparion (see also phylogenetic section below). This assignment can be tested with the discovery of more complete material, especially well preserved skulls. References to C ingenuum outside of Florida have been relatively infrequent. Quinn (1955:73) listed both Nann*us tehonensis and N. ingenuum [sic] as occurring in the Lapara Creek Fauna of the Texas Gulf Coastal Plain. Forsttn (1975) later referred the entire sample of medium-sized hipparionines from both the Lapara Creek and Clarendon faunas to Nannippus cf. ingenuum. She gave no specific reasons for her referral, and presented no comparisons with material from Florida. MacFadden (1980; 1984) referred the same population from the Clarendon Fauna to H. tehonense, and a portion of the Lapara Creek sample is also referable to this species otHipparion. Most of the other specimens from the Lapara Creek Fauna included in "Nannipmls cf. ingenuum" by Forstdn (1975) are referable instead to Nannippus s.s. (Hulbert 1987a). Only two specimens from Texas appear to represent C ingenuum, TMM 31204-1 and 31081-501. Although other Lapara Creek specimens might belong to this taxon, they are isolated teeth that are best regarded as specifically and generically indeterminant. It is worth noting that this is a rare (perhaps unique) instance of Quinn (1955) underestimating the taxonomic diversity of a group of Gulf Coastal Plain equids. MacFadden (1984:132) synonymized "Nannippus" ingenuus and N. lenticularis (a late Hemphillian species particularly well known from the Coffee Ranch, Edson, and Uptegrove localities of the Great Plains) "...based on overall similarity in size, crown height, and dental pattern... ." This synonymy, if valid, would extend the range of C ingenuum north of the Gulf Coastal Plain to Nebraska. However, examination of large numbers of individuals of both taxa fails to substantiate MacFadden's synonymy. While of similar size ("H." ingenuum is slightly larger on average, but observed ranges of tooth length and width overlap), there are numerous important differences between the two. In C ingenuum, a moderately developed DPOF is located posterior to the infraorbital foramen. In N. lenticularis, the posterior cheek region doe5 not contain a fossa, but according to MacFadden (1984:130) there is a small fossa located anterior to the infraorbital foramen. This observation is based on a single specimen, F:AM 113731. I believe that this feature is an artifact of the preservation and/or preparation of this particular specimen, and that most of HULBERT: CORMORmPPARION & HIPPARION FROM FLORIDA 263 what has been interpreted as a fossa is the exposed inner wall of a maxillary sinus. In any event, the facial regions of the two are different, even with our presently inadequate knowledge of the cranial features of both species. Significant dental differences between the two taxa are: 1. Unworn crown height of upper and lower third and fourth premolars of C. ingenuum is about 45 mm, of first and second molars about 50 mm. In N. lenticularis, unworn P34 crown height is about 50 mm, and 57 mm for M12. 2. In C. ingenuum, the P2 and p2 are much longer than the other premolars, with well developed anterostyles and paraconids, respectively. In N. lenticulmis, they are relatively short, with poorly developed anterostyles and paraconids. The former is characteristic of Connohipparion and Neohippan'on, the latter condition is found in Nannippus and H. tehonense. A reduced anterostyle is a derived character state for hipparionines (see below). 3. In C. ingenuum, pli caballins are well developed, frequently bifurcated or multiple, and persistent throughout wear. in N. lenticularis, pli caballins are at best moderately developed, not branched or multiple, and are often absent or lost with wear (e.g. FAM 111731, MacFadden 1984, Fig. 98D). 4. In C. ingenuum, fossette plications are very complex, often deep and bifurcating. In N. lenticularis, fossette plications are shallow and usually simple, with a much greater tendency for the pli hypostyle to be absent. 5. In C. ingenuum, the lingual border of the protocone is often straight or concave, especially in premolars. In N. lenticularis, it is nearly always convex. The protocone also tends to connect to the protoselene much earlier in the premolars of N. lenticularis, at MSCHs as great as 25 mm. 6. In C. ingenuum, the protostylid is prominent and is isolated from the protoconid only for a short period of wear. In N. lenticularis, the protostylid is small, usually remains isolated from the protoconid until very late wear-stages, and is occasionally absent (see also Dalquest 1983). 7. In C. ingenuum, the metaconid of the p2 and dp2 typically have an anterolabial plication that closes off the metaflexid. In samples of N. lenticularis, such structures are absent or very rare. There are additional, minor, differences between the cheekteeth of the two taxa, but those listed (plus their respective cranial morphologies) suffice to indicate that they belong in separate species (and genera). MacFadden (pers. comm.) has suggested that the differences listed above represent chronoclinal variation, but in my opinion they are of greater magnitude than those observed between early and late populations of any other equid species with a long, well 264 BULLEI'IN FLORIDA STATE MUSEUM VOL. 33(5) documented chronologic range (e.g. Neoh(ppan'on trampasense or C occidentale). Typically in those cases, younger populations show more advanced features than those observed in more rudimentary form in the older population. Samples of intermediate age are of intermediate grade. With regard to C. ingenuum and N. lenticularis, some of the features of the younger taxon (those numbered 3,4,5, and 7) are more primitive than those of the older. Others (2 and 6), which are derived in the younger population, are not observed in any rudimentary or incipient form in the older samples. N. lenticularis is best considered a distinct, valid species. Connoh*anon ingenuum is known from three periods of time: early Clarendonian (ca. 10-11 Ma, Lapara Creek and late Agricola Faunas); latest Clarendonian-early Hemphillian (ca. 8-9 Ma, Love Site, McGehee Farm, Haile 19A, Mixson's, i.e. the Archer Fauna of Webb and Hulbert 1986); and late early Hemphillian (ca. 6-7 Ma, e.g. Port Manatee, Withlacoochee River 4A, 4X, and Dunnellon sites). The Honduran sample (see below) may date from either of the two latter periods. As might be expected in such a long-ranging taxon, some morphological differences exist between populations from different eras. Unlike those described above, the differences are minor, and more primitive states are observed only in older populations. Unfortunately, the youngest samples are the most poorly known; their small sample sizes prevent detailed analysis and permit only specific identification. However, detailed comparisons can be made between the cheekteeth of the oldest and intermediate populations. The sample of Connohippanon ingenuum from the Agricola Fauna (material from Hookers Prairie, Phosphoria, and Silver City mines in referred specimens listing) is clearly intermediate in size between C ingenuum and C plicattle from the Archer Fauna (Tables 1-4, Figs. 9, 10). This is the case for mean P34 and M12 APL, TRW, and BAPL. The sample of P25 from the older fauna is too small for reliable analysis. Mean p2, p34, and m12 apl and bapl are also intermediate between average values for C ingenuum and C plicatile from the Love Site (Table 4), but are all nearer the means for C ingenuum. Mean atw and ptw, however, are not statistically different from the Love Site sample of C ingenuum for all lower cheekteeth. That the Agricola Fauna sample belongs to C ingenuum and not C plicatile nor their common ancestor is suggested by two characters. The shape of the lingual border of the protocone varies, but in 4 of 7 premolars and 8 of 12 molars it is straight or concave (Fig. 9C, 9D, 9F-9H). In the sample of p23,4 of 7 display a plicated metaconid on the occlusal surface (Fig. 1OA), a much greater frequency than that state is observed in C plicatile. Flattened labial protocone borders and plicated p2 metaconids are derived character states uniting C ingenuum and C emsliei (see below). Thus their appearance in the Agricola Fauna sample, although in lesser frequencies than observed in younger populations, suggests phylogenetic affinity with the C ingenuum-C emsliei clade. With their HULBERT: CORMORHIPPARION & HIPPARION FROM FLORIDA 265 toothrow lengths of 112-130 mm, C ingenuum and C emstiei are also smaller than their sister taxa in ConnohOWarion, that have average toothrow lengths greater than BO mm. The Agricola Fauna sample referred to C ingenuum has toothrow lengths of about 120-125 mm (Table 4). This moderate size is most parsimoniously regarded as another synapomorphy uniting it with C ingenuum. In a lineage decreasing in size with time, older populations will not unexpectedly be larger than younger ones. In a more simplistic but less parsimonious scenario, the Agricola Fauna sample could represent the common ancestor of C ingenuum and C plicatite. But this would require three reversals in the C plicatile lineage to reacquire its primitive size, protocone shape, and p2 metaconid morphology. A more likely phylogenetic hypothesis suggests instead that the cladogenetic event that resulted in C ingenuum and C plicatile occurred prior to the early Clarendonian, and that there is a chronologic gap in the first appearance of C plicatile of at least 2 million years. The preceding has emphasized the differences between the older and younger populations of C ingenuum, rather than their similarities (Figs. 5-10). As an example of the latter, the Agricola Fauna Sample includes several examples of unworn or slightly worn P34s, M12s, p34s, and m12s. These demonstrate identical values for unworn MSCH and mcch as specimens from the Love Site, about 45 mm for P34 and p34, 50 mm for M12 and m12. Webb and Perrigo (1984) referred a sample of teeth from the Gracias Formation of Honduras to "H*padon" plicatile. As they noted, in many respects this sample does compare favorably with Connoh*arion plicatile as defined in this study. However, the size of the specimens, as listed by Webb and Perrigo (1984:244), clearly indicates a referral instead to the smaller C ingenuum (Tables 1, 2). The north-central Florida and Central American samples are remarkably similar, considering the geographic distance separating them. One difference is a slightly better developed pli caballinid in the Honduran sample. It is more persistent on some of the p2-p4 than is typical for the Florida sample, and even occurs on some molars. It is not nearly so well developed as in C emsliei, however, nor does the Honduran sample have the derived stylar features and extreme fossette complexity observed in the Pliocene species from Florida (Hulbert 1988b). The relatively advanced nature of the Honduran sample of C ingenuum emphasizes Webb and Perrigo's (1984) conclusion that the age of the Gracias Fauna is early Hemphillian rather than late Clarendonian. The other major study of the Gracias Fauna (Olson and McGrew 1940) also recognized a hipparionine, which was referred to Neoh»adon montezuma. The heavily worn molar they figured (plate 2E) may indeed represent Neohipparion, as its PRL (8 mm) falls outside the OR of C ingenuum (Table 3). The more abundant UF sample from this fauna does not contain any specimens referable to Neohippan'on. As noted by Simpson (1930), Connohippanon ingenuum is commonly found in Florida; however, typically it is less abundant than its congener, C 266 BULLETIN FLORIDA STATE MUSEUM VOL. 33(5) plicatile . For example, the Love Site sample consists of a minimum number of 62 individuals of C ingenuum and 83 of C plicatile (MacFadden and Hulbert in press). Its biostratigraphic range in Florida as recognized here extends from the early Clarendonian to the late early Hemphillian. Simpson (1930, fig. 20C) illustrated a molar labelled as C ingenuum that is instead Nannippus minor (UF/FGS V-1426). The specimen is less complete than represented in the figure and slightly smaller. The slightly worn molar he figured as C plicatile (Simpson 1930, fig. 2OB) is referable to C ingenuum instead. Diagnostic features of this specimen (UF/FGS V-1485) are its narrow protocone with a flattened lingual border and its small TRW. C ingenuum has often been recognized from the late Hemphillian Upper Bone Valley Fauna of Florida (Sellards 1916; Simpson 1930; Webb and Tessman 1968; Webb 1969b), either as"Hippation" ingenuum or 'Wannippus" ingenuus. Most of these records are referable to the more advanced species, C emsliei. Others are either generically indeterminant or referable to N. minor. Both C plicatile and C ingenuum are recognized from the Bone Valley Formation (see referred specimens listing and Figs. 4C-4E, 8C, 9, 10), but are interpreted as being derived from older horizons than that which produces C emsliei. Based on the joint occurrence with C plicatile (see below), Pseudhipparion skinneri (Webb and Hulbert 1986), Neohipparion trampasense, and Calippus cerasinus (Hulbert 1988a), the Nichols Mine "Stream Matrix" Horizon records are either late Clarendonian or early Hemphillian. The early Clarendonian age of the late Agricola Fauna specimens discussed above is based on their co-occurrence with Pseudhipparion cumvallum, Nannippus n. sp., Calippus martini, Protohippus supremus, and Hypohippus sp., cf. H. a#inis (Hulbert 1988a). Connohipparion (Notiocradohipparion) plicatile (LEIDY),1881 Figs. 14-19; Tables 1-7 Hippothen'um plicatile LEIDY 1887:310; COPE 1889:444; LEIDY and LUCAS 1896:50; MACFADDEN 1984:170-174 (in part, not fig. 141). Hippanon plicatile (Leidy), HAY 1902:620; GIDLEY 1907:905; HAY 1916:41-42; OSBORN 1918:192; SIMPSON 1930:187-188 (in part); HIRSCHFELD and WEBB 1968:249; JACKSON 1978:53; WEBB and PERRIGO 1984:243-245 (in part). Hipparion sp., SIMPSON 1930:176. ?Hipparion plicatile (Leidy), SrI RTON 1940:184. Neohippan'on cf. /eptode Merriam, HULBERT 1982:159 (in part). Cormohipparionplicatile (Leidy), HULBERT 1988a:274, 280; HULBERT 1988b:454. T*e Specimen.-- USNM 3292, a R upper cheektooth, probably a P4 (MacFadden 1984, fig. 140). Type Locality and Horizon.-- Mixson's Bone Bed, Levy County, Florida. Alachua Formation, early Hemphillian. HULBERT: CORMORHIPPARION & HIPPARION FROM FLORIDA 267 Distribution.-- Very late Clarendonian through early Hemphillian (about 6.0 to 9.0 Ma) Of central Florida. Referred Specimens.-- Coffrin Creek, Alachua Co., FL: UF 17129 RP2; 17178A L p2; 17178B R p34; UF 58552 R m12; 17223 R m3. Gainesville Creek, Alachua Co., FL: UF 7441 LP34; 7437 L m3. Love Site, Alachua Co., FL: UF 32257 partial skull with R and L I2-I3,P3 and R P4; 32260 assoc. R maxilla with P4-M2 and L manila with P4-M3; 32250 R maxilla with P4-M3; 32255 L maxilla with DP2-Ml; 35891 L maxilla with P4-Ml; 32267 assoc. R DP2-DP4; 96622 assoc. R DP3-DP4; 27993, 32262, 32263 3 assoc. R and L P2-M3; 32283 assoc. R P2,P4-Ml and L Ml-M2; 32270 assoc. R P2-Ml,M3 and L Ml-M3; 32265 assoc. R and L P2-M2; 32264 assoc. R P3-P4,M2-M3 and L P4-M3; 27316 assoc. R P2-M3 (MacFadden 1984, fig. 144); 36289 assoc. R Ml-M2 and L P4; 32276, 32295 2 assoc. L DP2-DP4; 96619-96621 3 assoc. L DP3-DP4; 96618 assoc. L DP3-DP4 and R DP3; 32285 assoc. L DP2-Ml; 32266, 96933 2 assoc. L P2-M2; 53284 assoc. L P3-P4; 35902- 35905, 35920-35922, 35926, 35927, 35929, 35940, 35944, 35946, 35948, 35952, 35954, 35957, 35960, 35961, 96623, 96937 21 R Dn; 36078, 36081, 36084, 36087, 36088, 36090, 36096, 36097, 36100, 36103, 36105, 36112, 36114, 36115, 36119, 36121, 36128-36130, 36132, 36137, 96881 22 L DP2; 96624-96649, 96935, 96936 28 R DP34; 69812, 96650-96685 37 L DP34; 35967, 35970, 35976-35978, 35984-35985, 35988-35989, 35994, 36000-36002, 36005-36006, 36008, 36010, 36015, 36031-36035, 36037, 36040, 36044, 36047, 36051, 36063, 36068-36071, 36263, 36266, 36269 37 R P2; 36143, 36147, 36151, 36153, 36156, 36158-36159, 36162, 36165-36166, 36169, 36171, 36177, 36181-36182, 36184-36186, 36188, 36190, 36193-36194, 36201, 36205, 36209, 36211, 36218-36219, 36223, 36225- 36227, 36237, 36243-36244, 36247, 36250, 36253, 36255 40 L P2; 50641, 50643- 50644, 53287-53289, 53291-53294, 53296, 53298-53299, 53301, 53423-53424, 62326-62350, 62438, 62447, 69810 44 R P34; 53302-53328, 53373, 53422, 62306- 62324, 62381, 62384, 69809 50 L P34; 50642, 50645, 50647-50648, 53329-53332, 53334-53340, 53344-53349, 53351-53353, 53416, 53417, 62351-62365, 69811 42 R M12; 53154-53156, 53354-53360, 53362-53367, 53369-53371, 53374, 53377-53379, 53418-53421, 62367-62376, 62378-62380, 62382-62383, 62385-62389, 62408 48 L M12; 96388-96402 15 R M3; 96403-96428 26 L M3; 32119 R mandible with dp2-dp3; 32157 assoc. R and L mandibles with p2-p3,dp4,ml-m2; 32180 assoc. mandibles with R m2-m3 and L p3-m3; 27317 R mandible with p2-m3 (MacFadden 1984, fig. 145); 32104, 32174 2 R mandibles with p2-m2; 32212 R mandible with p2-ml; 32116 R mandible with p2,p4-m3; 32190 R mandible with p3-m3; 32144, 35893 2 R mandibles with p4-m3; 32107, 32199 2 L mandibles with p2-m3; 32111, 32118, 32160 3 L mandibles with dp2-dp4; 32123, 32163 2 L mandibles with p2-m2; 32113, 32151 2 L mandibles with p2-ml; 32198, 32245 2 L mandibles with p3-m3; 32165 L mandible with p4-m3; 32182, 32105, 36282, 32134, 36285, 36280 6 partial R mandibles; 32197, 32195, 32137, 36287, 35892, 32102 6 partial L mandibles; 35895 assoc. R dp2-dp4 and L 268 BULLETIN FLORIDA STATE MUSEUM VOL. 33(5) dp3-dp4; 32286, 90154 2 assoc. R dp2-dp4; 90160 assoc. R dp2-dp3; 90155, 90156 2 assoc. R dp3-dp4; 32196 assoc. R p2-m3 and L p2-m2; 32293 assoc. L p4,m2-m3 and R p3-p4; 32146 assoc. R p2-m3; 32225 assoc. R p2-p4; 32239 assoc. R p3-m3; 32203, 32229 2 assoc. R p4-m3; 32181, 32223 2 assoc. R ml-m3; 32232, 32221 2 assoc. R m2-m3; 90158 assoc. R dp3 and L dp2-dp4; 90159 assoc. L dp2-dp3; 32227 assoc. L ml-m2 and R m2; 65167 assoc. L ml-m3; 90162-90179 18 R dp2; 69813, 69814, 90121-90159 41 R dp34; 90180-90197 18 L dp2; 90071-90120 50 L dp34; 50355, 50356, 50358-50362, 50364, 50365, 50369, 64926-64927, 64929-64931, 64934-64936 19 R p2; 50370, 50372, 50374-50379, 64937, 64940-64943, 64466 14 L p2; 50461-50479, 50544, 50635, 50639, 64981-64986, 64988-65025, 65027 67 R p34; 50447-50455, 50460, 50458, 50640, 32226, 64944-64946, 64948-64980, 65166 50 L p34; 50420-50437, 50439-50445, 65076-65109, 65028-65050, 69816 83 R m12; 50400-54017, 54019, 65113-65131, 65133-65165, 69818, 90262 73 L m12; 50380-50389 10 R m3; 50390-50399 10 L ml "Stream Matrix" Horizon, Nichols Mine, Polk Co., FL: UF 24638 R P2; 23986, 24636 2 R P34; 28837 L M12; 24626, 24655 2 R dp34; 24628, 24631 2 R p34; 24625 R m12; 24642 L m12. Fort Green Mine, Polk and Hardee Cos., FL: UF 47473 R Ml; 102095 R dp3; 53924 R p34. Four Corners Mine, Hillsborough Co., FL: UF 102620 R P34; 102022 R M3. Unknown mine near Mulberry, Polk Co., FL: AMNH 22482 mandible with p3-m3. Peace River, near Gardner, Hardee Co., FL: UF/FGS V-4889, UF 55948 2 R M12; UF/FGS V-1482 L p34; UF 55951 R m12; 55952 L m12. McGehee Farm, Alachua Co., FL: UF 17220 L ma~illa with DP34; 17221 L PD34; 17121 L P2; 17099, 17124, 17208, 19237, 19428, 45616 6 R P34; 17206, 17210, 17212, 45614, 53548 5 L P34; 9605, 9611, 17213, 3 R M12; 17090, 17207, 17214, 17125, 45612 5 L M12; 45615 R M3; 7234 assoc. R mandible with p2-m2 and L mandible with p2-p4; 9541 assoc. R dp2-dp3; 12050 R dp2; 7240 L dp34; 45619, 45620 2 R p2; 17167, 45627, 53454 3 R p34; 17196, 17197, 45623, 53452 4 L p34; 7243,9606, 16837, 171404 17173, 17192A, 18707, 45624, 45625, 53451 10 R m12; 17100, 17139, 1714OB, 17169A, 17169B, 17172, 17175, 17192B, 17193-17195, 45618 12 L m12; 45631 R m3. Pareners Branch Site, Alachua Co., FL: UF 53564 L P34; 55492 L M3. Haile 5B, Alachua Co., FL: UF 17226 L P34. Haile 6A, Alachua Co., FL: UF 102551 R p2. Haile 19A, Alachua Co., FL: UF 47319, 103729 2 R M12; 103730 RM3; 103731 R dp34; 103732 R m12; 58387 L m12; 103733 L m3. Cummer Mine No. 8, Alachua Co., FL: UF/FGS V-1408 L p34. Mixson's Bone Bed, Levy Co., FL: UF/FGS V-1428 R P2; USNM 3292 R P34 (holotype); FAM 111730 assoc. R P2-M2 and L P3-M3 (MacFadden HULBERT: CORMORHIPPARION & HIPPARIONFROM FLORIDA 269 1984, fig. 142); 107876 badly crushed skull with R and L DP2-Ml; 113620 assoc. L P2-M2 and R Ml; 113621 assoc. R P2-P3,M2-M3 and L P3,M2-M3; 113625-113629, 113638 6 upper cheekteeth; 107874 assoc. R and L mandibles with B,cl,p2-p3,dp4,ml-m2 and R i2 (MacFadden 1984, fig. 143); 113630 assoc. R p2-p3 and L p2; 113634 assoc. L m2-m3; USNM 3309 R p2; RAM 113640 L p2; 113636 R p34; 113631, 113632 2 L p34; 113633, 113635 2 L m12. Moss Acres Racetrack Site, Marion Co., FL: UF 93000 assoc. juvenile crushed skull with R and L DP2-Ml, R and L mandibles with dp2-ml, and various post-cranial elements; 96386 assoc. R Ml-M3 and L M3; 69968 R M12; 69967 assoc. male R and L mandibles with il-B,c,p2-m3; 103754 assoc. R and L mandibles with R il, dp2-dp4,ml-m2 and L di2-di3,dp2-dp4,ml-m2. Withlacoochee River Site 4X, Marion Co., FL: UF 53525 L M12. Dunnellon Phosphate Company mine, Marion Co., FL: USNM 8265 R maxilla with P2-P4 (Hay 1916, plate 28). Dunnellon Phosphate Company, Plant No. 5, near Hernando, Citrus Co., FL: UF/FGS V-1400 R M12. Port Manatee Site, Manatee Co., FL: UF 107541 R m3. Revised Diagnosis.-- Medium-sized Connoh»an'on with toothrow lengths of 128-140 mm in middle wear-stages. Unworn MSCH of P2 about 40 mm and M12 about 58 mm. On average, smaller than C occidenmle, generally with less complicated fossettes, less elongate protocones, and relatively more elongated diastema. More hypsodont and more complex enamel plications than C sphenodus. Larger and higher crowned cheek teeth than C ingenuum; larger than C emstiei with simpler fossette plications and weaker pli caballinids. Description.-- All cranial material referable to Connohippanon plicatile lacks uncrushed or complete pre-orbital regions. Four specimens (UF 32257, 32255, 35891, 32250) from the Love Site and UF 93000 from Moss Acres preserve parts of the facial region, but none contains the entire DPOF. The specimen with the most complete DPOF, UF 32250, is an uncrushed, partial maxilla with well worn but diagnostic P3-M2 (Fig. 14B). It contains the ventral base and rim of a deep (at least 15 mm) DPOF located 46 mm dorsal to the toothrow (Fig. 14A). The three other Love Site specimens also preserve small portions of a fossa. These crania differ considerably from Love Site specimens assigned to Neohippanon #ampasense, in which the DPOF is reduced to a slight depression (MacFadden 1984; Hulbert 1987b). The badly crushed skull of UF 93000 preserves only the dorsal portion of both DPOFs. They were evidentially less well-rimmed dorsally than those of C occidentale, and probably shallower. Another extremely crushed skull of C plicatile, F:AM 107876 from Mixson's Bone Bed, was noted by MacFadden (1984:174) as not possessing a deep DPOF. F:AM 107876 is crushed and fragmented to such a 270 BULLETIN FLORIDA STATE MUSEUM VOL. 33(5) 0 3cm1 B / -I FA , in r Sprot, 9 6444 rrijavFA:51#1 34/2 I I --- . 9 2cm FIGURE 14. Lateral (A) and occlusal (B) views of UF 32250, Cormoh*parion plicatile, Love Site (late Clarendonian), Alachua County, Florida. Lateral view shows the well defined ventral margin of the DPOF. Occlusal pattern of R P3-M2 represents the late wear-stage for this species. degree that not even the orbits are discernible. Until discovery of uncrushed, more complete cranial material, the facial region of C plicatile and the exact morphology of its DPOF must remain poorly documented. The Love Site and Moss Acres Racetrack Site specimens demonstrate the presence of a moderate to deep DPOF, but not its complete morphology. The following dental descriptions are based primarily on the abundant Love Site sample, with added comparisons from other localities. Statistics of cheektooth dimensions of C plicatile are presented in Tables 3 and 4. Upper cheekteeth of C plicatile are characterized by oval protocones with convex lingual borders, richly plicated internal fossette borders (especially in early wear), moderately deep hypoconal grooves open to near the base of the crown, and strong styles. The P2 has a well developed anterostyle, and is by far the longest tooth in the toothrow (Fig. 15A, 15B). The oval protocone of the P2 HULBERT: CORMORHIPPARION & HIPPARIONFROM FLORIDA 271 generally remains isolated from the protoselene until mid-wear. Many P25 with MSCHs of 20 to 25 mm retain isolated protocones. Unworn MSCH of the P2 is about 38 to 42 mm. Fossette plications resemble those of the P34, except that the anterior half of the prefossette usually has two or three small folds. The pli caballin is generally single, although often bifurcated. The P34 (Figs. 14B, 15, 16B) are much larger in occlusal area than the M12, with larger and more widely opened parastyles and mesostyles. The parastyles are often slightly grooved, and metastyles are occasionally developed, but not as prominently as in C emsliei or Neohipparion eutystyle. In very early wear, the protocone of the P34 is elongate, Occasionally with a rudimentary spur, but with wear the spur (if present) vanishes and the protocone rapidly becomes more oval. This change is produced by an increase in protocone width, as protocone length is uncorrelated with crown height (r = 0.05, n = 132), while protocone width has a significant negative correlation with crown height (r = -0.64, n = 130, p < 0.001). Protocone length is generally less than twice its width, a feature useful in distinguishing C plicatile from its slightly smaller contemporary, Neohipparion trampasense. The main axis of the protocone runs in a more direct anterior-posterior direction than that of N. trampasense. The protocone of the P34 does not connect to the protoselene until very late wear-stages (MSCH of 15 mm or less). Fossette and pli caballin complexity also vary considerably with crown height. In early wear, the Well developed pIi caballin is usually multiple, branched, or both (Fig. 15A). With wear, it decreases in length and complexity, usually becoming single by late mid-wear (MSCH of 15 to 30 mm), and lost or rudimentary only in very late wear (Fig. 14B). In the Love Site sample, the anterior half of the P34 prefossette most often is not plicated (60% of examined specimens, n = 112), or has a single, small pli protoloph (32%; Fig. 15A, 15B). The latter condition occurs most frequently in the less worn teeth. The posterior half of the prefossette in early to moderate wear is richly plicated, with four to nine folds, of which several are often deep and bifurcating. The anterior half of the postfossette is only slightly less complex, generally with a deep, often branched pli postfossette and two to five accessory plicati(ms. In moderate to late wear-stages, these internal fossette plications become shallower and less numerous (e.g. Fig. 14B), although even very heavily worn teeth usually retain at least two or three folds. The posterior half of the postfossette has a single (59%, n = 110), relatively shallow and small pli hypostyle, or lacks all plications (39% of observed specimens). Early Hemphillian samples of P34s of C plicatile, while of similar proportions to the Love Site sample (Tables 1, 3), are on average more complexly plicated (Fig. 16B, 16D). For example, of 16 observed P34s, seven (44%) had a single pli protoloph, eight (50%) had a pli protoloph plus one or two accessory plications, and only one lacked all plications on the anterior half of the prefossette (the character state found in the majority of the Love Site sample). Unworn P34 MSCH varies from 50 to 55 mm. 272 BULLEUN FLORIDA STATE MUSEUM VOL. 33(5) A 4 \.Z'. 11*-u»511/ 1~56%3~ 1{j~ f <13.... \*5-7 9 2cm B $1 ** - L - e '*'' + 1, I. *//// 'm"' A tt , 96=. FIGURE 15. Occlusal views of representative upper toothrows of Connoh*parion plicatile from the Love Site (late Clarendonian), Alachua County, Florida. A. UF 32262, L P2-M3, an average-sized individual, early wear-stage. B. UF 32270, R P2-M3, a reiatively small individual, early wear-stage. A Bo.j CL C/=0!~1~Ih 9 2pm D E _~ F 4 1 -=?8 Aill ~ .r# 9». ...I-, FIGURE 16. Occlusal views of upper cheekteeth of Comwhipparion plicatile from the early Hemphillian of Florida. A. UF 93000, L Ml, sectioned to show moderate wear-stage, Moss Acres Racetrack Site (late early Hemphillian), Marion County. B-C. McGehee Farm Site (early Hemphillian), Alachua County, Florida. B. UF 17210, sectioned L P34; this individual more closely matches the enamel pattern of the holotype than any other subsequently found specimen. C. UF 9605, R M12. D-F. Mixson's Bone Bed (early Hemphillian), Levy County. D. F:AM 113620, sectioned L Pl E. F:AM 113620, R Ml, same individual as in Figure 16D. F. F:AM 113629, heavily worn L M12. HULBERT: CORMORHIPPARION & HIPPARION FROM FLORIDA 273 In mid-wear, M12 of C ph-catile have nearly square occlusal surfaces, with length slightly exceeding width on average (Table 3). However, since length is positively correlated with crown height (r = 0.77, n = 124), and width is negatively correlated with crown height (r = -0.50, n = 123), the ratio of length to width changes greatly through ontogeny. The M12 protocone is slightly smaller than that of the P34 (Table 3), but similar in morphology and degree of isolation from the protoselene. The pli caballin is only rarely double (6% of observed Love Site specimens, n = 121), but it persists until late wear, when it gradually disappears in extremely worn teeth (almost always after MSCH is less than 20 mm). In the Love Site sample, the anterior half of the prefossette of the M12 generally lacks plications (81%, n = 120; Figs. 14B, 15). A single, relatively shallow pli protoloph is occasionally found in early wear-stages, and it rarely has accessory plications. The posterior half of the prefossette most often has from three to five, rarely bifurcated plications, and a well developed prefossette loop. The anterior half of the postfossette generally has between two and four plications (86% of Love Site sample, n = 121), usually consisting of a relatively deep, unbifurcated pli postfossette and several labial accessory plications. The posterior half of the postfossette in early wear often has a single pli hypostyle; it is usually lost by mid-wear. As with the P34, M12 of C plicatile from the early Hemphillian localities (Mixson's, McGehee, Moss Acres) are on average more complicated than the Love Site population (Fig. 16A, 16C), although of the same general size (Table 3). For example, a third of the pooled sample of M12 from the younger sites have multiple pli caballins (Fig. 16A, 16F). Also, fossette plications are more numerous and tend to persist longer throughout wear. Unworn MSCH of M12 are about 56-59 mm for the Love Site sample. Based on UF 96386 from Moss Acres (Table 1), Hemphillian C plicatile had increased its unworn crown height by about 8%. Deciduous upper premolars of C plicatile have very large parastyles, generally multiple pli caballins, and oval protocones. The degree of fossette complexity is intermediate between that of the P34 and the M12. Protoconal spurs are not uncommon, and the hypoconal groove frequently has a pli dihypostyle. No specimens are available to indicate the relative size of the DPl, but it was apparently lost in some mature individuals (e.g. USNM 8265). The referred Love Site sample includes numerous associated lower dentitions and partial mandibles referred to Connohipparion plicatile (as well as several hundred isolated lower cheekteeth), but none is complete; i.e. there are no mandibles that include symphyseal regions, and only one with an ascending ramus. However, UF 69967, from the Moss Acres Racetrack Site, has a relatively complete symphysis (Fig. 17). In this adult male individual (ml mcch = 35.3 mm), the incisors form a normal equine arcade about 55 mm in width (across the Bs) and are slightly procumbent. The ils have only small traces of the infundibula remaining, and the Bs do not have enclosed infundibula. The symphyseal region is extremely elongated, with an 1dl of 274 B U LLE T IN F LO R ID A S rA T E M U S E U M V O L. 33(5) 0 3cm1 ./ FIGURE 17. Left lateral view of UF 69967, left mandibular ramus and symphysis of Connohipparion plicatile from the Moss Acres Racetrack Site (late early Hemphillian), Marion County, Florida. Occlusal view of the cheekteeth of this specimen shown in Figure 18(. Note the extremely elongated diastema, characteristic of the subgenus Non'ocradoh / ~0. 00. 4 .400 *Q;. f..0 B 3. 0- .4 / lf/ \00 0 bef#'i,44,/04,1#014'**e.'P##~244*94e##5*r#eme' 4 00 .09 00 00 00 00, 13 11 16 18 20 22 26 25 28 30 31 12 24 29 10 27 23 21 19 17 15 9 Hulbert (1987b) erred in the choice of Me,ychippus insignis as an outgroup for all hipparionines. This resulted only in an incorrect determination of polarity for several character states, and did not affect the relative phylogenetic positions of the in-group taxa. The most notable of the former was that the shallow DPOF observed in Neohipparion and Pseudhipparion was thought to have been derived from a primitively much deeper fossa. The computer- generated analyses suggest instead that a relatively shallow DPOF is the shared primitive condition in the Hipparionini and Equini, based on its development in "M." primus and 'Tarahippus" leonensis (Hulbert 19884· Megchippus s.s. HULBERT: CORMORHIPPARION & HIPPARION FROM FLORIDA 291 FIGURE 25. Most parsimonious cladogram depicting phylogenetic relationships among hipparionine equids relative to three outgroups, "Parahippus" teonensis, "Merychippus" pnmus, and the Equini. The cladogram was produced by the computer program PAUP based on an analysis of 58 cranial and dental characters (Tables 8,9). A. Hypothesized relationships of "Merych*pus" sejuhctus, members 6f the Neohippan'on-genus group, and the H*anon-genus group. B. Hypothesized relationships among members of th-e Hippan'on-genus group. The clad6gram is supported by the following list of synapomorphies. Numbers referring to characters and character states are those of Table 8. The character state to the left of the hyphen is the ancestral state, to the right the derived state. Node 1 (Hipparionini): 31 (1-2); 33 (2-3); 48 (0-1); 52 (0-1); 55 (0-1). Node 2 ("M." sejunctus): 12 (0-1); 27 (2-1); 40 (1-0). Node 3: 27 (24); 28 (2- 4); 35 (1-2); 70 (3-4). Node 4 (Neohipparion-genus group): 20 (1-3); 23 (1-2); 31 (2-1); 43 (0-1); 48 (1-2); 52 (1-2); 63 (0-1); 71 (2-4). Node 5 ("M." coloradense): 16 (2-3);70 (4-5). Node 6: 2 (2- 1); 5 (14); 7 (0-1); 10 (0-1); 23 (2-3); 34 (0-1); 62 (1-2); 63 (1-2). Node 7 (Neohipparion): 19 (0- 1); 24 (0-1); 25 (0-1); 26 (0-1); 27 (4-6); 28 (4-6); 29 (1-0); 38 (2-0); 39 (2-0); 40 (1-0); 46 (1-0), 54 (1-2); 60 (0-1); 61 (0-1); 65 (0-1); 71 (4-7). Node 8 (Pseudh 1 or 1-> 0 counts as 1.0 step (the same as with equally weighted characters). For a character with three states (0,1, and 2), single step transformations, e.g. 0-> 1, 1-> 2, or 1-> 0, each count as 0.5 steps. In the case of four states, single step transformations count as 0.333 steps; or in the general case of n states, each single step counts as 1/(n-1), such that the complete transformation from 0-> (n-1) always equals 1.0. The purpose of this Option is to prevent 298 BULLEIIN FLORIDA STATE MUSEUM VOL. 33(5) characters with many states from having undue influence on the cladogram (Swofford 1985). With this option in effect for the data set in Table 9, PAUP produced two equally most parsimonious cladograms. One of these was that produced with the default equally weighting option and shown in Figure 25. The second cladogram differed slightly in that it united C ingenuum and C emsliei with C occidentale as a monophyletic group, whose closest sister taxon was "H." primigenium. There are several reasons for considering this alternative arrangement less likely than the one previously discussed. It unites C ingenuum, C emsliei, C occidentale, and 'H." primigenium by their more complexly plicated fossettes (characters 32 and 33), and increased size (the same three characters that unite Node 24 in Fig. 25B). C occidentale, C ingenuum, and C emsliei are united only by a flattened lingual border of the protocone (24). C ingenuum and C emsliei are united by the same characters as listed for Node 29 in Figure 25B, except that the decrease in size is of greater magnitude. C ingenuum is derived relative to C emsliei by simplifying its fossettes, i.e. C ingenuum does not really possess any of the three characters that are supposed to unite this clade, so it must reverse all three back to the primitive condition. The alternate tree also requires parallel acquisition of the very elongated muzzle in C plicatile and C ingenuum + C emsliei. This combination of parallelisms and reversals seems less likely than that required for the other tree. Another reason for considering the alternate hypothesis less likely is the chronologic distribution of the tan involved. Although biostratigraphic records should not be used to construct phylogenies (Novacek and Norell 1982), they can be used, along with biogeographic distributions, to choose among phylogenies of equal length. This is especially valid when the fossil record is relatively complete, as is the case for late Miocene horses. In the alternative phylogeny, both C emsliei and C ingenuum possess derived character states relative to one another, thus as sister taxa their time of first appearance should be about the same. In the first cladogram (Fig. 25), C ingenuum has no autapomorphies relativeto C emsh-ei. Thus it may have given rise to C emsliei through anagenesis (if their observed ranges do not overlap in time), or through cladogenesis with the parent species not going extinct durmg speciation (if there is overlap in their temporal ranges, as appears to be the case based on the Moss Acres Racetrack Site record). The chronological ranges of these two species, as currently known, favors the latter scenario. C ingenuum is first known in the early Clarendonian, 4 myr prior to the first appearance of C emsliei at Moss Acres. The alternative cladogram also implies a much older (late Barstovian) origination for C plicatile than is reflected in the fossil record (late Clarendonian). Therefore, I consider the cladogram in Figure 25 to represent the most likely hypothesis of relationships among the species of the Hippanon-genus group, based on the distribution of cranial and dental character states. HULBERT: CORMORHIPPARION & HIPPARIONFROMFLORIDA 299 BIOCHRONOLOGY AND BIOGEOGRAPHY The chronological and geographical distribution of the five recognized species of Connohipparion, the three species of Hipparion, and "Megch*pus" goorisi are outlined in Figure 26. The oldest known species of Hipparion, "H." shirleyi, and "Meiych*pus" goonsi are both at present known only from the Barstovian of the Texas Gulf Coastal Plain. However, as noted by MacFadden (1984), additional specimens may exist in other regions, but would be difficult to separate from other merychippines without very complete material. MacFadden (1984:154-155) extended the chronologic range of 'M." goorisi beyond that of the type locality into the late Barstovian, with reference to specimens from the Cold Springs Fauna. He specifically referred TMM 31242-71 to "M." goorisi, and stated that it came from the "Cold Springs L. F." (p. 155). However, this specimen is from the Point Blank local fauna, stratigraphically the lowest among the local faunas that make up the Burkeville Fauna (early Barstovian; Tedford et al. 1988), and is the holotype of Protohippus vems Quinn (1955). If MacFadden's (1984) reference is followed, then "M." goorisi must become a junior synonym of P. vems. However, the two are distinctly different in fossette complexity, degree of protocone attachment, and crown height and are not specifically identical. The status and phylogenetic position of Protoh»us vems was further discussed by Hulbert (1988a). I do not recognize 'M." goorisi from the Cold Springs Fauna; rather that fauna contains a relatively primitive population of C sphenodus. The late Barstovian-early Clarendonian taxon C sphenodus is widely distributed, from California to Florida, and apparently represents the ancestral stock for the remaining members of the genus (Woodburne et al. 1981; MacFadden 1984). Bernor and Hussain (1985) discount C sphenodus as being "directly" ancestral to Old World hipparionines, because it is too primitive, and noted the variation in DPOF morphology in specimens referred to C sphenodus. Further analysis may show that C sphenodus as presently used is a grade, with two separate clades, one closer to Notiocradoh*pa,ion, and one to C occidentale, respectively. The southern and eastern C sphenodus-derived lineage is represented by C plicatile, C ingenuum, and C emsliei, which together comprise the subgenus Notiocradoh*parion. C plicatile as known is restricted to the late Clarendonian and early Hemphillian of peninsular Florida. C ingenuum is also known from Central America; probably this represents an immigration event southward along the Gulf Coastal Plain from Texas. C emsliei is the youngest surviving species of Connoh*panon in North America (Fig. 26) and is as yet known only from localities along the Gulf Coastal Plain in Florida (Hulbert 1988b) and Louisiana (Manning and MacFadden in press). It is possible that all three had much larger ranges in southeastern North America, and that their apparently limited distribution is GULF COASTAL PLAIN §NAL < WEST COAST GREAT PLAINS MA J TEXAS/LOUISIANA FLORIDA ~ T tl .r ' tZ Z -1 Z Z: IL L~ AA N |BL AN CA N| 14 W 5- 1 G- 144 10- 'D 11'-K 11 Ici ' C 1 C JC E B - H B 15- B U LLE TIN FLO R ID A S rA T E M U S E U M V O L. 33(5) A FIGURE 26. Biogeographic and chronologic distributions of North American species of Comtohipparion and H*ation, exclusive of Central America. Triangles point to the approximate chronologic position of the type locality for each species; dashed lines fill stratigraphic gaps. A. "Me,ychippus" goorisi. B. C. sphenodus. C C. occidentale. D. C. plicatile. E. C. ingenuum. F. C. emsliei. G. Connohipparion sp., Rattlesnake Formation, Oregon. H. Hipparion shir*yi. I. H. tehonense. J. Hippanon sp., cf. H tehonense. K H. forcei. Identifications of C sphenodus and C. occidentale in Florida are tentative (see text) Other than the present, study, references include Hulbert (1988b), MacFadden (1984), Manning and MacFadden (in press), and personal observations of the TMM, F:AM and UCMP collections. HULBERT: CORMORHIPPARION & HIPPARION FROM FLORIDA 301 more a renection of a biased fossil record. For example, the absence of Connohippa,ion in the late Clarendonian and Hemphillian of the Texas Gulf Coastal Plain (Fig. 26) probably just reflects the total lack of faunas of the appropriate age. In the Great Plains, Connohipparion is not observed in any late Hemphillian (6.0 to 4.5 Ma) or younger localities. The Upper Bone Valley Formation sample and those from the Florida Blancan mark the last North American appearance of what was once one of the dominant equid genera of the late Miocene. Its absence in western late Hemphillian and Blancan faunas is probably real, as sites of this age are fairly well known. In general Connohippdrion is much poorer in terms of biostratigraphic utility for correlation of Gulf Coastal Plain and Great Plains localities than Pseudhippan'on, Neohippan'on, Calippus, »or Protohippus. This is because of the long temporal durations of species' ranges (Fig. 26, Table 10) and limited geographic distribution. They are of more use for correlation within the circum-Gulf of Mexico region, e.g. C ingenuum in Honduras, and especially within the state of Florida. The genus Hippanon (sensu MacFadden 1984) ranges from the late Barstovian to early Hemphillian in North America. The record of Hippanon east of the Rocky Mountains is scattered, and it is typically rare, the Clarendon Fauna being a notable exception. In faunas from California, however, the genus is more common. Relative to its contemporaries, Hpparion was not a diverse group. Besides H. shirleyi, only two other chronospecies are recognized; both range from the Clarendonian to the late early Hemphillian. The sample of H»an'on, cf. H. tehonense, in Florida ranges from the early Clarendonian to late early Hemphillian. Records from the Moss Acres Ranch Site and the Withlacoochee River 4A 1.f. are two of the youngest for the genus. MacFadden (1984) referred a similarly aged population from Nebraska to H. foirei. Hipparion is also a poor biostratigraphic indicator in Florida, because of its rarity and long chronologic range. Table 10 summarizes the distribution of equid species in Florida from the late Barstovian to the end of the Blancan, the interval for which Connohipparion is known to have been present. The late early Hemphillian (6-7 Ma) was the period of maximum species richness for the Hippanon-genus group in Florida, with six contemporary species. Only two of these lineages persisted into the late Hemphillian, C emsliei and Nann(ppus minor. This mid- Hemphillian extinction event (at about 6.0 Ma) includes the last appearance in North America of four equid genera, Hipparion, Cal»us, Protohippus, and Plioh»us (Hulbert 1987a, 1988a), in what marks a major event in the decline of the Clarendonian Chronofauna (Webb 1977, 1984). 302 BULLErIN FLORIDA SrATE MUSEUM VOL. 33(5) CONCLUSIONS Of the five recognized North American species of Connoh*parion and three of Hipparion, all appear in Gulf Coastal Plain faunas except H. forcei. "Meiych»us" goonsi and H. shirleyi are both small, primitive species known from the Barstovian Fleming Formation of southeast Texas, but are as yet unknown from Florida. The ubiquitous late Barstovian-early Clarendonian C sphenodus is provisionally recognized for the first time in eastern North America. Limited samples of isolated teeth are found in the lower of two superposed faunas in the Bone Valley Formation of south-central Florida. C occidentale, well known from the latest Barstovian, Clarendonian, and early Hemphillian of the Great Plains and California, is also provisionally recognized for the first time in Florida, from a limited suite of isolated teeth. The nearest well-documented occurrence is in the Lapara Creek Fauna of southern Texas (Forstdn 1975). Two previously enigmatic hipparionine species described a century ago by Joseph Leidy from Mixson's Bone Bed, Levy County, Florida, are both referred to Connoh*parion. This is based on newly recovered and more complete material from the Love, McGehee Farm, and Moss Acres Racetrack sites. Referred material of C ingenuum ranges in age from early Clarendonian to late early Hemphillian (about 11.0 to 6.0 Ma), and demonstrates minor amounts of chronoclinal evolution, most notably in increasing enamel complexity. The range of C plicatile is limited to the late Claredonian-early Hemphillian (9.0-6.0 Ma), but it is typically more abundant than C ingenuum and shows a greater degree of chronoclinal change. C plicatile is not recognized as yet from localities outside of peninsular Florida. The population of medium-sized hipparionines from the Lapara Creek Fauna is for the most part not referable to C ingenuum, as was suggested by Quinn (1955) and Forst6n (1975), but instead to H. tehonense and to an undescribed species of Nann*pus. Only two Texas specimens are referred to C ingenuum. C ingenuum and C plicatile differ principally in size and crown height. · The Connohipparion population from the Gracias Formation of Honduras, referred to C plicatile by Webb and Perrigo (1984), is notably smaller than Florida samples of that species, but well within the observed range of C ingenuum. C ingenuum is not the senior synonym of Nannippus lenticulan's (Cope), as was proposed by MacFadden (1984). Although of similar size, the two tan differ in facial morphology, crown height, and in numerous details of the dentition. Connohippan-on is not observed in late Hemphillian faunas of the west, as it disappears there along with many other members of the Clarendonian Chronofauna. However, the genus persisted in Florida until near the end of the Blancan. The upper Bone Valley Formation and two late Blancan faunas HULBERT: CORMORHIPPARION & EPPARION FROM FLORIDA 303 from Florida contain a small to moderate-sized and distinctively complexly plicated species, C emsliei (Hulbert 1988b). C emdiei apparently is descended from C ingenuum, but their observed ranges overlap in the late early Hemphillian, based on a new record from the Moss Acres Racetrack Site. Many dental features that appeared in the C ingenuum-C emsliei lineage parallel those observed in Neoh*pan'on, but occur about four million years later. In terms of its degree of hypsodonty, C emsliei is much more primitive than contemporary species of Pseudhippan'on, Neohippan'on, Nann*pus, Astroh»us, and Dinoh*pus, and reflects a Clarendonian grade of evolution (Hulbert 1988b). However, its fossettes have on average more plications than any other North American hipparionine. Hippanon tehonense is recognized (at least provisionally) in Florida, from an early Clarendonian horizon in the Bone Valley Formation, from the late Clarendonian Love Site, and from the early Hemphillian Moss Acres and Withlacoochee River 4A sites. The latter two represent the youngest records of the species. These referred specimens are morphologically similar to the topotypic sample from California, but are on average smaller. LITERATURE CITED Becker, J. 1985. Fossil herons (Aves: Ardeidae) of the late Miocene and early Pliocene of Florida. J. Vert. Paleon. 5:24-31. Bernor, AL 1985. Systematic and evolutionary relationships of the hipparionine horses from Maragheh, Iran (late Miocene, Turolian Age). Palaeovertebrata 15(4): 173-269. , and S.T. Hussain. 1985. An assessment of the systematic, phylogenetic and biogeographic relationships of Siwalik hipparionine horses. J. Vert. Paleon. 5:3247. , M.O. Woodburne, and JA Van Couvering. 1980. A contribution to the chronology of some Old World faunas based on hipparionine horses. Geobios 13:705-739. Bode, F.D. 1931. Characters useful in determining the position of individual teeth in the permanent cheektooth series of merychippine horses. J. Mamm. 12:118-129. Christol, J. de. 1832. [untitled]. 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Pp. 153-210 in M.O. Woodburne (ed.). Cenozoic Mammals of North America: Geochronology and Biostratigraphy. Univ. California Press, Berkeley. 306 BULLETIN FLORIDA STATE MUSEUM VOL. 33(5) Webb, S.D. 1969a. The Burge and Minnechaduza Clarendonian mammalian faunas of north-central Nebraska. Univ. California Publ. Geol. Sci. 78:1-191. . 1969b. The Pliocene Canidae of Florida. Bull. Florida State Mus., Biol. Sci. 14(4):273-308. . 1977. A history of savanna vertebrates in the New World. Part 1: North America. Ann. Rev. Ecol. Syst. 8:355-380. . 1984. On two kinds of rapid faunal turnover. Pp. 417436 in WA. Berggren and JA. Van Couvering (eds.). Catastrophes and Earth History. Princeton Univ. Press, Princeton, New Jersey. , and R.C. Hulbert. 1986. Systematics and evolution of Pseudh*parion (Mammalia, Equidae) from the Late Neogene of the Gulf Coastal Plain and the Great Plains. Pp. 237- 285 in KM. Flanagan and JA. Lillegraven (eds.) Vertebrates, Phylogeny, and Philosophy. Univ. 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Florida locality abbreviations: ARS, Agricola Road Site; COC, Coffrin Creek; DUN, Dunnellon Phosphate Mines; FGM, Fort Green Mine; 4CM, Four Corners Mine; GRD, Gardinier Mine; GZP, Gray Zone, Phosphoria Mine; HSB, Haile 5B; H19, Haile 19A; HPM, Hookers Prairie Mine; LOV, Love Site; MAR„ Moss Acres Racetrack Site; MGF, McGehee Farm; MIX, Mixson's Bone Bed; PHM, Phosphoria Mine; PTM, Port Manatee Site; RZP, Red Zone, Phosphoria Mine; SCM, Silver City Mine; SMN, "Stream Matrix" Horizon, Nichols Mine; W4A, Withlacoochee River.Site 4A; W4X, Withlacoochee River Site 4X. Honduran locality abbreviation: GFH, Gracias Fauna. See Table 3 for univariate statistics. Measurements defined in text and in Figure 3. An "a" before a value indicates that it is an approximation from a broken or waterwom specimen. All measurements in millimeters. Loc Tooth Side APL TRW PRL PRW BAPL MSCH Comment H U LB E R T: C O R M O R H IP PA R IO N & H IP PA R IO N FR O M FLO R ID A 307 Cormohipparion sp., cf. C. sphenodus UF 28434 RZP Ml R a19 18.8 8.2 3.5 15.5 29.7 UF 28432 RZP M12 R 19.8 18.4 6.6 3.9 -- a31 UF 28433 RZP M12 R - a20 8.4 - a16 a43 UF 28435 RZP M3 L 18.7 15.4 7.3 3.0 17.6 38.2 Cormohipparion sp. cf. C. occidentale UF 28607 GZP P34 R 23.4 22.1 7.0 4.1 18.7 37.4 UF 102552 HPM P34 L 22.9 21.8 8.8 3.5 17.6 a29 UF 102553 HPM M12 R 22.6 23.5 8.7 a4 18.6 33.8 Connohipparion ingenuum UF 36107 LOV DP2 L 30.8 18.4 5.2 4.0 27.1 9.8 UF 35953 LOV DP2 R 29.3 17.8 4.9 3.2 26.6 11.7 UF 36122 LOV DP2 L 30.5 17.0 5.3 3.4 28.2 12.7 UF 35932 LOV DP2 R 30.9 18.1 5.8 3.4 27.7 14.0 UF 35947 LOV DP2 R 30.9 17.0 5.3 3.5 27.0 14.5 UF 35953 LOV DP3 R 23.8 17.9 4.7 3.6 20.1 12.1 UF 36122 LOV DP3 L 23.7 16.4 5.4 3.0 -- 12.6 Table 1 Continued. 308 B U LLE TIN FLO R ID A S rA T E M U S E U M V O L 33(5) Loc Tooth Side APL TRW PRL PRW BAPL MSCH Comment Connohipparion ingenuum (continued) UF 35947 LOV DP3 R 212 17.2 5.2 3.3 20.1 14.5 UF 36122 LOV DP4 L 24.2 14.8 5.7 2.8 21.1 13.9 UF 35953 LOV DP4 R 24.5 15.8 5.0 2.8 20.2 15.5 UF 102554 HPM DP34 L 22.2 15.0 4.6 2.9 19.1 15.0 UF 98107 ARS DI'34 R 21.6 15.4 3.4 2.9 19.1 12.5 UF 96691 LOV DI'34 L 213 17.5 5.8 3.4 20.0 9.0 UF 96751 LOV DP34 R .22.5 17.2 6.3 3.6 19.3 11.7 UF 96740 LOV DP34 R 22.9 16.3 6.7 2.9 20.2 16.1 UF 98101 ARS P2 L - 19.7 7.0 4.8 - 13.6 UF 98108 ARS P2 R 26.3 18.8 6.2 3.8 21.5 28.1 UF 28683 GZP n L a27 18.6 5.5 3.6 20.9 28.2 UF 36203 LOV P2 L 24.4 18.0 5.5 3.5 19.5 19.1 UF 32300 LOV P2 L 26.0 18.5 5.3 3.8 - 26.1 UF 36161 LOV P2 L 25.5 17.2 6.4 3.4 20.3 28.8 UF 36020 LOV P2 R 24.2 18.2 6.0 3.6 18.0 29.2 UF 36150 LOV P2 L 26.6 15.8 5.9 4.0 19.8 35.7 F:AM 107875 MIX n L 25.5 18.3 6.4 3.7 20.6 20.4 UF 46323 GFH n L 216 16.8 5.3 3.4 - 35.7 UF 98101 ARS P3 L 20.6 21.2 73 4.5 16.4 16.9 UF 98106 ARS P3 L 21.8 19.9 6.3 4.1 16.5 28.2 UF 32300 LOV P3 L 20.4 20.7 5.9 33 - - F:AM 107875 MIX P3 L 19.7 193 6.8 3.8 - 24.2 UF 17204 W4A P3 R 18.7 18.3 6.7 3.7 - UF 98101 ARS P4 L 20.1 21.2 7.7 4.6 15.9 17.4 UF 98102 ARS P4 L 18.2 19.7 6.9 3.7 16.8 15.2 UF 98106 ARS P4 R 21.5 20.2 6.4 3.9 16.4 30.8 UF 32300 LOV P4 L 19.8 193 6.1 3.6 - - F:AM 107875 MIX P4 L 19.5 19.2 7.0 17 16.7 27.4 UF 17204 W4A P4 R 18.0 17.4 6.8 3.6 - - UF 18329 W4X P4 L 19.1 18.6 7.1 3.7 16.3 318 UF 18034 GFH P4 R 19.6 19.0 7.3 3.4 15.6 41.2 UF 65706 SCM P34 L 21.4 - 19 3.1 16.5 a30 UF 28555 GZP P34 L 820.5 19.1 7.3 3.8 - 35.4 UF 28556 GZP P34 R 21.2 19.8 7.0 4.1 16.4 363 UF 107564 ARS P34 R 19.7 18.1 63 3.1 15.4 212 UF 62464 LOV P34 L 18.8 17.8 6.4 3.6 16.6 19.1 UF 53404 LOV P34 R 18.9 18.4 6.8 3.4 14.7 27.3 UF 53401 LOV P34 R 193 18.3 7.1 3.7 15.6 27.5 UF 53400 LOV P34 R 19.6 18.6 8.0 3.8 14.9 353 UF 62434 LOV P34 R 213 17.3 7.9 33 16.0 42.6 UF 7246 MGF P34 L 193 18.6 6.9 33 16.4 23.4 UF 17200 W4A P34 R 19.3 18.4 5.1 3.6 15.8 21.3 UF 107543 I,IM P34 L 19.8 a18 5.3 3.3 a15.5 a34 UF 46321 GFH P34 L 17.8 17.9 5.9 3.7 14.6 15.6 H U LB E R T: C O R M O R H IPPAR IO N & H IP PA R IO N FR O M FLO R ID A UF 98102 ARS Ml R 18.1 18.4 7.2 4.0 16.2 123 UF 98101 ARS Ml R 18.7 19.8 7.2 4.2 15.1 15.2 UF 98106 ARS Ml L 19.4 18.9 5.9 3.7 15.5 28.7 UF 98104 ARS Ml L 21.4 17.8 6.8 12 143 43.8 UF 32300 LOV Ml L 18.1 17.8 53 3.1 - - FAM 107875 MIX Ml L 18.4 17.9 7.0 16 15.9 27.3 UF 17204 W4A Ml R 16.8 16.8 6.7 3.4 - - UF 98102 ARS M2 R 17.3 18.1 6.4 13 16.1 19.8 UF 32300 LOV M2 L 18.6 17.2 5.6 3.3 - 30.9 F:AM 107875 MIX M2 L 18.4 17.6 7.4 12 15.6 31.2 UF 17204 W4A M2 R 17.2 16.6 6.8 33 UF 98117 ARS M12 R 223 18.9 63 3.1 15.3 47.8 UF 98118 ARS M12 R 19.9 18.0 7.3 12 163 38.2 UF 98121 ARS M12 L 18.1 17.4 6.0 3.2 14.6 33.9 UF 62392 LOV M12 L 20.1 17.0 6.4 3.4 14.8 49.1 UF 62414 LOV M12 R 19.1 16.0 7.2 12 15.1 45.8 UF 62424 LOV M12 R 20.4 17.7 6.1 3.1 15.1 38.0 UF 53394 LOV M12 R 18.1 16.8 5.2 3.3 14.6 25.8 UF 62410 LOV M12 L 17.8 18.9 7.0 4.2 14.6 20.1 UF 45613 MGF M12 L 19.1 18.1 63 3.0 - - UF 107894 H19 M12 L 173 a153 5.6 - 14.4 29.1 USNM 3306 MIX M12 L 18.5 15.8 5.1 2.9 143 40.6 Holotype Table 1 Continued. 310 B U LLE T IN FLO R ID A S TATE M U S E U M V O L. 33(5) Loc Tooth Side APL TRW PRL PRW BAPL MSCH Comment Cormohippan'on ingenuum (continued) F:AM 113623 MIX M12 L 19.1 16.6 5.4 3.3 14.1 35.2 F:AM 113624 MIX M12 L 21.1 17.4 6.1 33 14.4 40.1 F:AM 113638 MIX M12 L 20.4 16.3 4.7 3.3 14.3 39.4 UF 45530 W4A M12 R 17.3 17.8 7.0 3.5 15.0 33.6 UF 28557 GZP M3 R 17.8 14.0 5.4 3.1 16.9 29.8 Connohipparion plicatile UF 36119 LOV DP2 L 32.5 19.4 5.4 4.0 29.0 9.6 UF 35929 LOV DP2 R 31.6 18.8 5.9 4.3 - 10.2 UF 35928 LOV DP2 R 31.8 18.4 53 3.2 28.5 12.3 UF 35931 LOV DP2 R 32.2 18.7 4.8 35 29.5 14.0 UF 36102 LOV DP2 L 32.6 17.6 5.6 3.5 28.9 17.9 UF 96621 LOV DP3 L 24.6 19.0 53 4.5 -- a6 UF 69920 LOV DP3 L 2A.4 20.3 5.8 4.5 - a7 UF 96622 LOV DP3 R 24.5 19.7 5.6 4.7 22.5 10.5 UF 35928 LOV DP3 R 25.0 17.9 5.6 3.3 21.4 13.2 UF 35931 LOV DP3 R 25.2 18.9 5.6 3.6 21.5 13.9 UF 96621 LOV DP4 L 24.9 18.9 6.6 4.8 - 9.2 UF 69920 LOV DP4 L 25.8 19.0 6.7 4.4 -- 12.3 UF 96622 LOV DP4 R 24.6 18.4 65 4.2 21.8 13.0 UF 35928 LOV DP4 R 25.9 16.3 5.9 3.0 20.9 16.2 UF 35931 LOV DP4 R 26.6 16.3 6.0 3.3 21.3 17.5 UF 36244 LOV P2 L 25.3 20.4 6.0 4.1 21.3 15.8 UF 32266 LOV n L 263 18.6 5.7 3.6 21.8 24.5 UF 32270 LOV P2 R 25.2 193 5.8 3.8 21.6 26.1 UF 32263 LOV n R 26.3 17.3 6.1 3.4 21.3 27.8 UF 32262 LOV n L 28.4 19.2 6.2 3.7 -- 28.3 UF 27316 LOV P2 R 26.5 19.0 6.2 3.8 24.0 29.9 UF 35988 LOV n R 30.6 19.9 6.6 4.3 21.9 32.0 UF 36159 LOV P2 L 28.3 19.0 5.7 3.7 22.5 38.7 UF 32265 LOV P2 R 28.7 17.5 5.7 3.9 21.4 39.8 UF 35978 LOV n R 27.8 18.7 5.5 3.9 20.9 40.7 UF 17121 MGF P2 L 26.6 20.4 6.7 4.1 22.2 16.2 UF/FGS V-1428 MIX P2 R 27.1 19.7 6.0 17 22.1 35.0 F:AM 111730 MIX n R 26.8 20.7 6.5 19 21.7 - F:AM 113620 MIX n L 25.6 20.0 5.6 3.9 22.4 24.0 F:AM 113621 MIX n R 29.3 19.7 6.3 17 21.4 34.4 UF 32270 LOV P3 R 22.0 21.3 6.3 4.0 17.2 32.2 UF 32263 LOV P3 R 21.4 19.3 6.5 3.5 16.8 32.9 UF 32250 LOV P3 L 20.6 20.0 8.0 43 - - UF 32266 LOV P3 L 213 20.3 6.1 3.6 17.7 23.0 UF 27316 LOV P3 R 22.2 20.8 6.5 3.7 18.5 23.1 H U LB E R T: C O R M O R H IP PA R IO N & H IP PA R IO N F R O M FLORIDA 311 UF 32264 LOV P3 R 22.2 21.2 6.9 4.1 - 19.4 UF 32262 LOV P3 L 23.2 21.7 7.7 3.9 18.0 32.2 F:AM 111730 MIX P3 R 22.6 22.1 6.6 3.9 - F:AM 113620 MIX P3 L 21.8 21.8 6.2 4.0 17.7 32.0 F:AM 113621 MIX P3 R 22.1 20.8 6.2 14 17.3 30.5 USNM 8265 DUN P3 R 22.7 212 7.6 4.1 - - UF 32264 LOV P4 R 20.6 20.5 7.1 3.8 18.2 22.4 UF 32250 LOV P4 L 20.1 19.7 8.3 4.2 - UF 27316 LOV P4 R 21.0 203 7.1 4.0 17.7 28.0 UF 32266 LOV P4 L 20.4 20.4 7.1 3.9 17.1 28.1 UF 32270 LOV P4 R 21.5 20.7 7.0 3.8 16.6 35.7 UF 32263 LOV P4 R 20.9 19.4 6.6 33 16.7 36.7 UF 32262 LOV P4 L 23.0 21.7 8.2 3.7 16.8 39.9 F:AM 111'DO MIX P4 R 223 22.0 7.4 4.0 17.4 - F:AM 113620 MIX P4 L 213 21.2 6.1 4.0 17.0 37.5 F:AM 113621 MIX P4 L 230 21.2 7.0 3.6 17.5 40.4 UNSM 8265 DUN P4 R 21.0 23.4 7.6 4.4 - 26.3 UF 62327 LOV P34 R 20.4 19.6 6.6 3.0 16.1 29.7 UF 53289 LOV P34 R 24.2 20.3 8.3 3.7 19.8 45.1 UF 53302 LOV P34 L 23.6 213 7.1 3.8 18.1 51.2 UF 17206 MGF P34 L 233 22.0 7.1 3.7 18.0 34.1 UF 17208 MGF P34 R 22.0 22.0 6.1 4.4 18.8 20.8 UF 17212 MGF P34 L 213 22.7 8.1 4.9 18.8 18.8 Table l Continued. 312 B U LLE I'IN F LO R ID A S TATE M U S E U M V O L. 33(5) Inc Tooth Side APL TRW PRL PRW BAPL MSCH Comment Connohipparion plicatile (continued) UF 17124 MGF P34 R 219 21.1 7.2 3.7 - 37.4 UF 17226 HSB P34 L 223 21.9 7.6 4.2 17.0 29.6 UF 102620 4CM P34 R 21.8 21.9 8.0 3.9 - 31.3 USNM 3292 MIX P34 R 19.7 21.1 6.9 3.8 17.1 29.7 Holotype F:AM 113626 MIX P34 L - 20.9 63 4.0 16.6 29.5 F:AM 113627 MIX P34 R 20.6 20.6 6.4 19 173 25.2 F:AM 113628 MIX P34 L 21.8 213 6.2 4.2 18.8 22.6 UF 32264 LOV Ml R 19.2 193 6.9 19 15-5 21.4 UF 32266 LOV Ml L 19.4 20.4 6.6 3.9 15.8 25.9 UF 27316 LOV Ml R 18.7 20.6 6.1 3.9 15.9 28.2 UF 32263 LOV Ml R 20.4 18.1 6.8 33 15.8 36.0 UF 32270 LOV Ml R 20.0 19.8 6.4 3.6 15.8 36.9 UF 32262 LOV Ml L 21.1 203 7.4 33 15.6 39.2 UF 32265 LOV Ml R 213 18.4 6.6 12 15.1 52.0 F:AM 111730 MIX Ml L 20.3 19.0 7.4 17 - 48.1 F:AM 107876 MIX Ml L 18.8 19.4 5.6 18 - - F:AM 113620 MIX Ml R 20.9 19.6 7.3 3.5 16.0 43.5 UF 96386 MAR Ml R 22.6 21.2 9.2 3.6 15.9 51.8 UF 32264 LOV M2 R 19.4 18.8 6.9 3.8 16.4 24.9 UF 32266 LOV M2 L 18.9 18.7 6.5 3.6 16.2 295 UF 27316 LOV M2 R 183 18.6 6.2 33 16.2 30.4 UF 32263 LOV M2 R 19.7 173 6.9 3.5 16.0 38.1 UF 32270 LOV M2 L 19.3 185 6.4 3.5 15.8 41.1 UF 32262 LOV M2 L 213 193 73 3.5 16.9 43.9 F:AM 111730 MIX M2 R 20.2 19.1 7.0 3.8 - - F:AM 113620 MIX M2 L 19.6 19.0 6.1 3.4 163 - F:AM 113621 MIX M2 L 22.0 17.1 8.4 2.8 16.4 50.0 F:AM 113637 MIX M2 R 183 18.9 6.4 3.4 15.8 32.0 UF 96386 MAR M2 R 214 20.1 9.9 4.2 183 61.4 UF 53349 LOV M12 R 19.9 21.4 7.3 43 16.9 17.6 UF 53338 LOV M12 R 20.8 19.8 7.7 3.4 18.2 38.0 UF 53334 LOV M12 R 20.3 19.1 7.4 3.6 16.7 45.0 UF 53332 LOV M12 R 23.0 18.8 6.3 3.2 17.2 51.2 UF 50647 LOV M12 R 22.4 16.9 6.4 2.6 15.9 56.5 UF 50648 LOV M12 R 213 153 6.9 2.8 16.2 59.3 UF 9605 MGF M12 R 215 20.8 7.1 3.6 17.1 49.3 UF 9611 MGF M12 R 21.0 19.9 6.1 3.6 16.8 45.6 UF 17214 MGF M12 R 21.4 20.4 7.1 3.5 17.3 38.1 UF 17090 MGF M12 L 203 20.2 5.9 3.6 - - UF 47319 H19 M12 R 20.6 a20 6.6 3.7 15.7 31.1 F:AM 113629 MIX M12 L 20.0 20.1 6.4 4.0 17.7 25.3 UF/FGS 1400 DUN M12 R 20.3 19.9 6.4 3.9 - 30.2 H U LB E R T: C O R M O R H IP PA R IO N & H IP PA R IO N FR O M F LO R ID A 313 Cormohipparion emstiei UF 102022 4CM M3 R 23.5 19.3 7.9 4.1 21.4 30.5 UF 95410 MAR M2 L 19.2 19.1 73 3.6 15.8 a38 UF 102600 4CM M12 R 17.1 16.6 7.0 3.1 15.2 263 UF 102601 4CM M12 L 20.8 16.3 7.1 2.9 16.0 52.9 UF 95410 MAR M3 L 20.2 15.1 8.9 3.1 - a42 Hipparion sp; d. H. tehonense UF 17198 W4A n L 22.0 16.7 5.3 3.0 16.6 22.6 UF 17205 W4A n R 223 18.6 4.8 19 17.0 24.2 UF 17205 W4A P3 R 183 19.1 5.3 3.8 15.4 253 UF 17205 W4A P4 R 18.6 19.8 5.8 3.9 13.9 32.0 UF 102599 4CM P34 R 21.3 19.2 5.6 3.3 - 40.0 UF 62165 LOV P34 L 19.1 17.3 5.0 33 153 17.1 UF 53375 LOV P34 L 18.8 18.2 5.5 3.3 15.4 30.5 UF 17205 W4A Ml R 17.1 18.3 5.1 33 14.3 34.8 UF 102622 4CM M2 L 173 16.1 5.2 3.4 13.2 a42 UF 17205 W4A M2 R 17.0 17.8 5.0 3.4 14.3 38.5 314 B U LLE I"IN FLO R ID A S rA T E M U S E U M V O L. 33(5) TABLE 2. Measurements of selected lower cheekteeth of Connohippan'on and Hipppan'on from Florida, Texas, and Honduras. See Table 4 for univariate statistics. Format and abbreviations as in Table 1. Additional locality abbreviations: BNV, Bone Valley Region, specific mine unknown; H64 Haile dA; LlC, I.2isey Shell Pit 1C; LCF, Lapara Creek Fauna, Texas; Measurements defined in text and in Figure 3. I.nc Tooth Side apl atw ptw mmI entl bapl mcch Comment Cormohipparion sp., d. C. sphenodus UF 28459 RZP p2 L 20.7 73 9.3 7.7 9.5 18.0 25.9 slight wear UF 28458 RZP p34 R 19.3 83 - - 9.4 - 31.9 UF 28460 RZP p34 L 18.7 - 10.0 10.5 9.3 16.2 28.2 UF 28446 RZP m3 R 22.8 9.0 7.8 103 6.8 - 29.9 UF 28447 RZP m3 R 22.8 8.7 8.2 10.1 7.1 - a29 Connohipparion ingenuum UF 90209 LOV dp2 R 28.2 7.1 9.3 9.8 11.5 24.0 9.0 UF 32294 LOV dp2 L 26.4 8.1 9.3 10.2 8.6 22.7 9.7 UP 32133 LOV dp2 R 273 6.6 8.1 10.1 11.3 23.6 11.5 UF 90200 LOV dp2 R 26.6 7.3 9.1 10.9 11.3 23.0 11.8 UF 98130 ARS dp3 L 24.9 93 9.4 13.0 10.8 22.3 13.9 UF 32294 LOV dp3 L 23.2 93 9.4 11.7 8.8 20.2 9.4 UF 90200 LOV dp3 R 24.2 8.9 8.8 11.7 10.1 20.6 13.7 UF 32133 LOV dp3 R 22.7 7.3 8.2 10.6 10.5 20.9 14.5 UF 98136 ARS dp4 R 22.4 9.7 9.1 12.3 6.2 20.3 a8 UF 98130 ARS dp4 L 27.4 8.2 7.7 123 9.8 22.1 18.7 UF 32294 LOV d'p4 L 24.2 9.2 8.7 12.0 8.0 20. 6 12.4 UF 90223 LOV dp4 L 26.8 8.4 7.9 11.8 9.8 19.7 19.1 UF 18126 GFH dp4 L 24.0 7.9 8.4 11.2 9.4 19.1 a18 UF 98138 ARS dp34 L 24.7 8.6 85 11.8 7.3 20.4 12.9 UF 90249 LOV dp34 R 22.7 10.8 10.1 12.2 7.4 21.4 5.7 UF 90222 LOV dp34 L 22.2 9.1 9.2 115 93 19.1 11.2 UF 90237 LOV dp34 R 233 8.9 8.7 11.6 9.3 21.7 13.5 TMM 31081-501 LCF p2 R 22.1 8.5 10.4 8.2 8.7 -- UF 98125 ARS p2 R 21.4 95 11.0 10.0 8.3 18.5 16.7 UF 98126 ARS p2 L 21.0 7.7 11.1 7.2 10.2 18.4 24.6 UF 98128 ARS p2 R 22.1 8.0 10.4 8.5 9.9 18.3 24.1 UF 98129 ARS p2 L 223 8.4 10.3 7.6 9.4 19.1 17.9 UF 24632 SMN p2 L 22.9 8.8 10.8 8.4 10.1 18.8 21.0 UF 64925 LOV p2 R 24.1 9.4 10.8 10.1 10.2 17.7 29.9 UF 64813 LOV p2 L 21.0 9.2 11.9 8.6 8.6 16.9 24.8 UF 64801 LOV p2 L 22.6 8.3 9.9 8.4 9.3 18.5 24.4 UF 64800 LOV p2 L 20.2 7.8 10.3 7.5 8.5 17.0 24.3 UF 64808 LOV p2 R 20.8 7.9 10.1 8.8 9.9 18.3 20.0 UF 64811 LOV p2 R 20.7 7.9 10.4 8.5 83 17.8 14.9 UF 47323 H19 p2 R 22.0 8.6 9.6 8.2 10.1 -- a25 UF 18036 GFH p2 L 19.9 8.3 10.1 9.2 8.4 16.8 24.3 H U LB E R T: C O R M O R H IP PA R IO N & m P PA R IO N F R O M F LO R ID A 315 TMM 31081-501 LCF p3 R 20.3 10.9 11.1 11.8 9.2 - UF 98125 ARS p3 R 20.8 12.1 11.6 13.5 10.0 18.0 21.5 UF 98126 ARS p3 R 22.8 10.5 11.2 13.1 10.8 18.1 34.8 UF 98128 ARS p3 R 22.4 10.8 11.2 13.3 9.9 17.6 31.3 UF 98129 ARS p3 L 20.9 11.0 10.9 12.1 10.1 17.4 23.3 'IMM 31081-501 LCF p¢ R 20.7 11.0 10.4 10.4 8.1 - a32 UF 98125 ARS p4 R 20.4 12.2 10.7 13.1 9.8 17.2 23.3 UF 98126 ARS p4 R 21.9 10.6 103 12.1 10.0 17.6 40.0 UF 98128 ARS p4 R 21.4 10.1 10.1 11.4 9.8 16.5 36.8 UF 98129 ARS p4 L 203 11.0 10.0 11.9 9.7 17.6 28.7 UF 28567 GZP p34 L 20.0 7.9 8.4 9.2 9.7 16.8 40.7 UF 28617 GZP p34 L 19.9 10.3 - 11.0 9.8 - a35 UF 107620 ARS p34 L 20.1 10.0 10.7 11.0 9.9 16.5 32.6 UF 24631 SMN p34 R - 11.4 11.4 13.0 9.9 - a36 UF 24646 SMN p34 L 17.7 10.5 9.6 10.6 7.9 15.0 13.8 UF 64752 LOV p34 R 21.3 10.0 10.4 11.2 10.2 16.3 38.4 UF 64774 LOV p34 R 19.0 93 8.9 10.0 8.5 14.4 37.6 UF 64780 LOV p34 L 21.0 10.5 11.0 11.8 9.7 15.8 33.0 UF 50457 LOV p34 L 20.3 10.4 10.9 12.1 9.8 17.6 29.7 UF 64791 LOV p34 L 18.4 11.6 10.5 113 9.1 15.7 20.6 UF 64768 LOV p34 R 18.7 11.9 10.9 11.4 8.2 16.2 13.6 UF 107529 LlC p34 L 18.0 10.6 10.5 10.7 8.7 - a33 UF 98126 ARS ml L 20.8 9.3 8.2 12.1 7.2 - a34 Table 2 Continued 316 B U LLE nN FLO R ID A S rA IE M U S E U M V O L. 33(5) Loc Tooth Side apt atw Ptw mmi entl bapl nnech Comment Cormohipparion ingenuum (continued) UF 98131 ARS ml L 19.3 9.2 83 11.8 6.9 16.2 25.6 UF 18036 GFH ml L 18.3 93 9.3 11.2 6.8 - a35 UF 98126 ARS m2 L 21.0 8.7 8.1 11.3 7.9 - 40.1 UF 98131 ARS m2 L 19.4 8.8 - 10.7 7.7 16.3 33.3 UF 18036 GFH m2 L 18.2 9.0 8.9 10.9 7.0 15.1 a39 UF 28568 GZP m12 R 22.1 73 6.6 10. 1 9.3 16.7 42.6 UF 28569 GZP m12 R 19.3 8.9 83 11.6 6.7 15 .4 a30 UF 28618 GZP m12 L 22.6 8.3 8.2 10.2 8.9 16.3 40.9 UF 98164 ARS m12 L 21.0 7.8 7.2 9.9 7.7 16.6 a42 UF 98162 ARS m12 L 20.8 9.2 9.0 11.2 7.9 15.3 37.1 UF 107621 ARS m12 L 21.9 6.8 7.2 9.9 7.9 15.7 39.9 UF 98150 ARS m12 R 18.9 9.7 83 11.0 8.3 16.2 25.5 UF 28857 SMN m12 L 18.6 9.7 9.0 10.9 6.1 15.7 25.1 UF.64820 LOV m12 R 21.6 7.7 11 9.6 7.9 15.7 46.6 UF 64818 LOV m12 R 21.0 8.0 7.2 10.0 7.7 15.5 41.5 UF 64878 LOV m12 L 17.8 8.8 8.2 10.6 7.7 14.5 36.2 UF 64899 LOV m12 L 19.1 8.6 7.7 10.4 6.6 16.3 28.9 UF 64876 LOV m12 L 18.1 10.0 9.1 10.9 6.1 15.3 23.0 UF 64901 LOV m12 L 17.3 10.0 83 9.7 43 15:2 13.3 UF 102621 4CM m12 R 18.9 8.9 8.9 10.8 7.7 - 43.0 UF 18035 GFH m12 L 193 9.2 8.9 10.9 8.1 15.1 a39 UF 103791 LOV m3 L 21.2 9.0 8.2 9.3 5.4 20.9 12.6 UF 69819 LOV n,3 L 22.1 83 8.1 10.3 6.6 22.4 a25 UF 103790 LOV m3 L 21.6 9.1 8.2 10.4 7.0 21.2 29.6 UF 103772 LOV m3 R 22.0 8.4 7.7 10.3 6.0 20.3 33.4 UF 107893 H19 m3 R 21.7 83 7.8 10.0 7.2 21.3 a42 UF 107542 FIM na R 20.4 8.4 7.6 10.3 5.6 - a35 Cormohipparion plicatile UF 35895 LOV dp2 R 273 9.1 10.3 12.6 11.0 24.1 8.7 UF 90178 LOV dp2 R 28.2 8.3 10.0 113 11.0 24.3 8.7 UF 90157 LOV dp2 R 28.7 8.7 9.8 12.0 11.4 24.3 12.0 UF 32286 LOV dp2 L 28.4 8.7 10.6 12.0 11.2 25.3 12.1 UF 90154 LOV dp2 R 29.1 8.0 9.9 11.6 11.6 255 13.1 UF 103754 MAR dp2 R 25.8 93 103 13.1 8.7 24.7 5.6 UF 93000 MAR dp2 L 27.6 8.4 8.9 12:9 113 24.3 10.4 UF 90158 LOV dp3 L 23.2 11.7 11.0 13.4 7.7 22.0 a4 UF 35895 LOV dp3 R 24.3 11.2 103 13.3 9.0 213 10.5 UF 90155 LOV dp3 R 25.8 10.9 103 14.7 9.9 23.0 10.5 UF 90156 LOV dp3 R 25.6 9.8 93 11.9 103 22.9 13.6 UF 32286 LOV dp3 L 25.6 9.9 10.2 13.1 10.3 22.7 14.4 UP 90154 LOV dp3 R 26.1 9.8 10.0 12.8 11.6 23.3 14.5 H U LB E R T: C O R M O R H IPPAR IO N & H IP PA R IO N FR O M FLO R ID A 317 UF 24655 SMN dp3 R 25.4 93 9.0 12.6 10.5 22.7 15.7 UF 103754 MAR dp3 R 22.7 11.8 10.2 14.3 8.1 21.9 a6 UF 93000 MAR dp3 L 213 9.3 8.4 13.6 11.7 20.9 13.7 UF 90158 LOV dp4 L 24.6 10.9 10.0 13.8 6.7 22.6 8.6 UF 90155 LOV dp4 R 293 10.2 9.8 15.1 9.1 235 13.9 UF 35895 LOV dp4 R 27.7 10.0 8.7 13.8 9.4 22.1 15.2 UF 90156 LOV dp4 R 28.1 8.6 8.3 11.7 9.9 22.6 16.8 UF 32286 LOV dp4 L 283 8.8 9.1 13.0 93 23.4 18.6 UF 90154 LOV dp4 R 28.6 8.7 9.0 12.7 10.6 23.2 20.0 UF 102095 FGM *4 R 26.9 8.9 83 12.5 8.9 23.4 18.7 UF 103754 MAR dp4 R 217 10.2 9.7 13.7 83 21.8 a10 UF 93000 MAR dp4 L 26.0 83 8.1 14.2 10.1 - 17.0 UF 24650 SMN dp34 R 23.1 9.7 9.3 12.8 93 205 11.6 UF 24626 SMN dp34 R 21.7 12.9 11.7 13.1 6.7 - aS UF 50379 LOV p2 L 22.9 9.4 11.4 10.1 6.1 20.9 11.9 UF 64942 LOV p2 L 212 9.5 11.6 103 10.1 20.8 19.8 UF 50366 LOV p2 R 213 8.1 10.0 9.8 9.6 19.8 21.4 UF 50373 LOV p2 L 23.1 8.0 9.6 8.6 103 19.3 28.0 UF 32196 LOV p2 L 27.1 9.9 12.1 11.1 113 21.0 31.0 UF 50374 LOV p2 L 24.6 8.9 10.8 10.7 10.9 20.0 34.7 UF 102551 H6A p2 R 22.2 10.1 123 11.0 6.8 19.7 13.7 UF 7234 MGF p2 L 22.7 9.6 12.2 10.8 7.6 - 17.0 Table 2 Continued 318 B U LLE nN F LO R ID A S rA T E M U S E U M V O L. 33(5) Loc Tooth Side apl atw ptw mml entl bapl rncch Comment Cormohippan'on plicatile (continued) USNM 3309 MIX p2 R 24.6 83 11.0 9.2 10.3 193 32.7 F:AM 113630 MIX p2 R 210 8.6 12.5 10.7 9.8 19.5 23.9 F:AM 113640 MIX p2 L 24.9 8.6 10.3 10.1 11.5 18.8 31.8 UF 6997 MAR p2 R 23.6 10.4 11.9 12.6 10.6 20.8 a22.5 UF 32225 LOV p3 R 243 12.0 12.8 13.9 10.6 19.0 43.3 UF 35892 LOV p3 L 22.2 12.2 12.2 13.5 10.1 - 28.7 UF 7234 MGF p3 L 21.0 13.4 133 13·7 10.6 - - F:AM 113630 MIX p3 R 21.4 12.5 12.4 13.1 10.1 18.2 26.1 UF 69967 MAR p3 R 21.8 12.6 11.7 143 10.5 18.7 26.3 UF 32225 LOV p4 R 23.9 11.1 10.8 12.5 9.9 18.8 52.9 UF 32133 LOV p4 L 22.8 11.7 12.0 13.0 10.4 17.7 41.6 UF32244 LOV p4 L 22.1 13.3 12.1 13.6 10.1 18.1 22.7 UF 7234 MGF 'p4 L 20.4 13.0 12.4 12.7 10.0 - - UF 69967 MAR p4 R 21.8 12.4 10.9 13.6 10.8 18.6 34.6 UF 50458 LOV p34 L 22.2 13.8 13.0 14.1 9.2 19.0 20.9 UF 50460 LOV p34 L 21.7 11.3 12.1 13.3 10.5 19.2 28.0 UF 50455 LOV p34 . L 21.8 12.3 12.9 13.5 10.7 18.3 34.5 UF 50469 LOV p34 R 24.0 12.3 12.4 14.6 10.6 18.7 40.2 UF 50448 LOV p34 L 24.8 11.3 11.4 12.6 12.0 19.2 47.4 UF 50463 LOV p34 R 233 10.8 10.2 11.9 10.6 18.1 51.0 UF 53924 FGM p34 R 20.5 13.6 - 13.3 8.6 16.4 22.5 UF 53507 BNV p34 L 22.1 11.1 10.1 11.5 10.3 - a35 UF 45623 MGF p34 L 22.1 11.6 11.6 11.9 10.8 - - F:AM 113631 MIX p34 L 24.1 11.1 10.9 12.1 10.8 17.3 41.6 F:AM 113632 MIX p34 L 21.2 12.6 11.7 13.1 9.5 17.7 30.0 F:AM 113636 MIX p34 R 19.4 12.3 11.7 12.7 8.0 17.8 18.9 UF 32219 LOV ml L 219 11.0 9.8 14.3 9.1 18.6 41.8 UF 32223 LOV ml R 26.3 9.1 8.6 12.1 9.9 16.9 55.6 UF 7234 MGF ml R 18.4 11.8 10.0 11.1 8.0 -- a27 F:AM 107874 MIX ml R 22.5 9.1 8.6 11.5 7.0 - 48.8 UF 69967 MAR ml R 20.2 10.9 9.6 12.1 8.2 17.6 32.9 UF 103754 MAR ml R 23.5 9.1 75 11.1 10.1 - 59.9 UF 32203 LOV m2 R 21.2 11.2 10.0 12.9 8.1 17.9 41.9 UF 32146 LOV m2 R 22.9 10.2 9.2 12.6 8.0 17.6 45.8 UF 32219 LOV m2 L 25.1 8.9 8.3 12.8 9.3 19.5 50.1 UF 32221 LOV m2 R 217 93 8.0 11.5 9.2 17.9 53.4 F:AM 113634 MIX m2 L 21.8 9.1 73 10.1 7.2 - 48.3 F:AM 107874 MIX m2 R 21.1 8.1 7.1 10.1 7.8 16.9 54.9 UF 69967 MAR m2 R 21.1 10.7 9.4 12.0 8.4 18.2 35.8 UF 103754 MAR m2 L - 7.0 6.3 - - - 63.0 very slight wear UF 24625 SMN m12 R 18.9 10.7 9.5 11.6 6.3 16.1 30.1 UF 24642 SMN m12 L 19.2 10.2 9.8 12.5 6.9 15.4 38.4 UF 50435 LOV m12 R 19.1 11.2 10.0 11.7 6.3 17.4 19.1 H U LB E R T: C O R M O R H P PA R IO N & H IP PA R IO N FR O M FLO R ID A 319 UF 50578 LOV m12 R 19.7 11.2 93 12.0 6.4 17.6 26.0 UF 50415 LOV m12 L 20.9 11.1 10.4 12.1 7.8 16.9 37.0 UF 50406 LOV m12 L 23.4 10.9 10.5 13.5 8.3 18.3 46.4 UF 50424 LOV m12 R 24.3 8.9 73 11.9 10.1 17.6 57.7 UF 58387 H19 m12 L 21.0 a10.2 - 12.4 9.1 - UF 103732 H19 m12 R 20.6 11.6 10.5 12.8 7.2 17.1 28.9 UF 50386 LOV m3 R 23.1 10.1 8.5 11.0 6.1 22.7 23.5 UF 50394 LOV m3 L 25.3 103 9.8 11.4 7.4 25.1 29.2 UF 50382 LOV m3 R 25.6 10.3 9.0 12.7 6.4 25.2 35.7 UF 50384 LOV m3 R 22.1 9.4 8.4 10.2 6.3 22.7 35.8 UF 50381 LOV m3 R 245 9.2 8.1 10.7 8.2 24.9 42.3 UF 103733 H19 m3 L 211 9.4 8.4 10.6 6.8 - 48.6 UF 69967 MAR m3 R 26.3 10.1 8.7 11.3 8.8 25.8 39.3 Cormohipparion emsliei UF 97259 MAR p2 L 20.9 9.3 11.0 11.8 9.7 - a8 -'UF·101964 GRD p2 L 19.5 7.4 8.8 7.0 8.8 16.0 27.6 UF 97259 MAR p3 R 18.8 13.0 12.0 12.3 93 UF 97259 MAR p4 L 19.4 12.4 10.9 11.9 10.4 -- - UF 102607 4CM p34 L 17.9 11.2 11.4 11.2 9.6 16.8 a17 UF 102641 FGM p34 L 20.2 10.9 11.0 12.0 9.8 - a35 UF 100226 GRD p34 R 18.3 9.6 8.9 10.9 93 14.2 a36 Table 2 Continued 320 B U LLE rIN FLO R ID A S TATE M U S E U M V O L. 33(5) Loc Tooth Side apl atw ptw mml entl bapl mcch Comment Connohipparion emstiei (continued) UF 102588 PHM p34 L 19.7 9.7 10.6 10.4 9.8 16.0 40.0 UF, 97259 MAR ml L 17.6 10.7 10.0 10.8 6.8 - - UF 97259 MAR m2 L 18.7 10.7 8.8 10.8 7.1 - - UF 102589 PHM m12 L 19.0 8.6 9.1 11.1 7.0 16.0 40.3 UF 97259 MAR m3 L 23.2 9.6 8.1 9.6 6.0 - - Hipparion sp., et. H. tehonense UF 17300 WU p2 R 19.0 7.3 8.7 8.0 83 - - UF 21027 W4A p2 L 20.3 7.9 10.0 7.4 8.1 - - UF 17300 W4A p3 R 19.0 9.3 9.1 9.2 7.6 - 39.2 UF 21027 W4A p3 L 18.0 103 11.0 10.6 9.2 - - UF 93203 HPM p34 R 19.9 9.3 9.6 9.6 9.9 15.7 30.0 UF 50753 PHM p34 R 19.7 8.8 9.1 9.6 8.2 14.0 31.7 UF 96526 LOV p34 R 19.1 9.8 10.0 10.1 8.2 - 29.5 UP 17300 Wa ml R 19.9 9.1 8.1 10.4 7.0 - 43.9 UF 21027 W4A ml L 17.2 9.6 9.0 10.7 6.0 - - UF 20868 W4A m2 L 21.7 8.0 7.1 10.1 7.8 - - UF 64917 LOV m12 R 20.3 7.4 7.1 9.7 6.9 - 34.9 UF 64918 LOV m12 R 18.3 8.0 7.1 9.3 6.6 14.1 29.9 UF 95388 MAS m12 L 20.6 7.9 73 9.3 7.2 - a50 TABLE 3. Univariate statistics for upper cheekteeth of Comwhipparion plicatite, C. ingenuum, and C emsliei from Florida. Measurements and their abbreviations defined in text and Figure 3. All measurements in millimeters. The first line in each entry gives x, s, and n; the second line, OR and CK Faunal abbreviations: LOVE, Love Site, Alachua County, late Clarendonian; HEMP, combined sample from various early Hemphillian localities including McGehee Farm, Mixson's Bone Bed, Moss Acres Racetrack Site, and Withlacoochee River 4A and 4X; ARS, Agricola Road Site, Polk County, early Clarendonian; and UBV, combined sample from the Upper Bone Valley Fauna, Polk and adjacent counties, latest Hemphillian Dental abbreviations: P2, sample of upper second premolars; P34, combined sample of upper third and fourthpremolars; M12, combined sample of upper first and second molars. TAXON C. plicatile C. plicatile C ingenuum C ingenuum C. ingenuum C. enisliei FAUNA LOVE HEMP ARS LOVE HEMP UBV H U LB E R T: C O R M O R H IP PA R /O N & H JP PA R /O N FR O M F LO R ID A 321 P2 APL 26.7, 1 .26, 55 27.1 , 1 .36, 4 26.7, 4 .95, 2 24.5, 1 . 24 , 43 25 .2, 0.42, 2 23.7, 2. 14, 5 23.5-30.6,4.73 25.6-29.3,5.03 26.3-27.0,1.86 22.1-26.9,5.06 24.9-25.5,1.68 21.8-27.3, 9.00 BAPL 21.8, 0.79, 54 22.0,0.40,5 21.2, 4.24, 2 19.8, 0.91, 42 20.6,-,1 20.2, 2.21, 4 20.4-24.0,339 21.4-22.4, 1.84 20.9-21.5,ZOO 18.0-21.2,4.60 17.6-22.7,10.95 TRW 19.6, 0.94, 57 20.6, 1.17, 6 19.0, 0.59, 3 17.3, 0.74, 44 17.8, 0.44, 3 18.1, 1.39, 8 17.3-213,4.79 19.7-22.8, 5.70 18.6-19.7,3.08 14.9-18.8,4.27 17.5-18.3,2.45 16.5-19.8, 7.70 PRL 6.1, 0.53, 58 6.5,0.68,6 6.4,0.68,4 5.9,035,43 6.9, 0.61, 3 5.7, 0.55, 11 5.0-7.2,8.66 5.6-7.6, 10.60 5.5-7.0, 10.59 4.6-73,9.37 6.4-7.6, 8.81 4.9-6.6, 9.71 PRW 4.1, 0.52, 57 4.0,0.30,6 3.9,0.60,4 3.6,0.32,42 3.4,0.23,3 3.3, 0.32, 10 3.4-5.6, 12.76 3.74.5,7.59 3.5-4.8, 15.30 3.0-4.8,8.87 3.3-3.7,6.73 2.8-3.7,9.83 P34 APL 21.8 , 1 . 19, 112 22. 1 , 1 .08, 18 20.7, 1 .21 , 9 19.2, 0.73, 49 18.8, 1 . 58, 9 18.9, 1 .40, 23 19.4-25.3, 5.46 19.7-23.9, 4.88 18.2-21.8, 5.83 17.8-21.5, 3.82 15.0-20.5, 8.44 17.0-22.1, 7.39 Table 3 continued 322 B U LLE I'IN F LO R ID A S rA T E M U S E U M V O L. 33(5) TAXON C. plicatile C. plicatile C. ingenuum C ingenuum C ingenuum C. emsliei FAUNA LOVE HEMP ARS LOVE .HEMP. UBV BAPL 17.6, 0.98, 107 17.6,0.72,14 16.3, 0.44, 8 15.5, 0.70, 46 15.8,1.12,4 15.5,1.01,15 15.9-20.0,536 16.6-18.8,4.09 15.4-16.8,2.68 13.9-16.6, 4.52 14.2-16.7, 7.07 14.2-17.9,6.53 TRW 21.0, 0.99, 112 213,0.96,19 19.9, 1.00, 8 183, 0.85, 49 17.4, 2.02, 9 18.2,1.29,24 18.6-24.1,4.70 19.8-23.4,4.46 18.1-21.2, 5.02 16.9-20.7, 438 14.0-19.5,11.6 16.2-20.9,7.06 PRL 7.4, 0.80, 116 6.9, 0.70,19 6.9,0.71,9 6.7,0.69,50 6.4,0.92,9 6.0,0.80,24 5.9-9.2, 10.80 6.1-83,10.04 5.8-7.9, 10.25 5.0-8.7, 10.24 4.8-73,14.49 4.8-8.8, 13.33 PRW 4.1, 0.47, 114 3.9, 0.36, 19 3.8,038,9 3.6,0.37,49 3.5,0.30,9 3.3,0.29,22 3.0-5.2, 11.41 3.44.9,9.23 3.1-4.6,15.20 3.1-4.7, 10.17 2.9-3.8,8.53 2,8-3.9,8.73 M12 APL 20.5 , 1 .36, 115 203 , 1 .51 , 18 19.3, 134, 11 18 .3, 1 .28, 54 18.6, 1 .27, 12 19.2, 1 .09, 24 18.2-25.4, 6.61 17.6-23.9,7.38 17.3-223,7.96 15.6-20.8,7.00 16.8-21.1,6.85 17.5-21.5, 5.66 BAPL 16.4, 0.69, 108 163, 0.69, 11 15.6,0.70,10 14.6,0.57,47 14.9,0.70, 8 15.7, 1.01, 17 15.1-18.2,4.20 15.4-17.7,4.20 143-165,4.48 13.5-15.8, 3.90 14.1-15.9, 4.67 13.9-17.1, 6.41 TRW 19.4, 1.22, 113 19.4,0.94,18 18.1, 0.89, 11 16.7,0.94, 54 17.0, 0.86, 12 17.2, 1.18, 22 15.4-22.0,6.29 17.1-20.8, 4.82 16.4-19.8,4.93 13.8-18.9, 5.65 15.6-18.1,5.03 15.3-20.1, 6.87 PRL 6.9, 0.62, 121 6.7, 0.68, 18 63,038,10 63,0.58,55 6.3, 0.83, 12 6.4,0.64,23 5.7-8.3,8.99 5.6-8.4,6.26 5.8-7.3,8.85 5.2-73,8.84 4.7-7.4, 13.33 5.2-7.5,9.96 PRW 3.8, 036, 120 3.6, 0.41, 18 3.4,0.44,11 3.4,0.39,54 3.3, 0.27, 12 3.2, 0.31, 23 2.6-5.2,14.76 2.84.9,11.37 2.8-4.2,13.00 2.3-4.3,11.45 2.9-3.7,8.11 2.4-3.7,9.72 TABLE 4. Univariate statistics for lower cheekteeth and mandibles of Cormoh*arion plicatile, C. ingenuum, and C entsliei from Florida. Measurements and their abbreviations defined in text and Figure 2. Format as in Table 3, except that the mandible of C emsliei is from the MossAcres Racetrack Site, not the Bone Valley. Dental abbreviations: p2, sample of lower second premolars; p34, combined sample of lower third and fourth premolars; m12, combined sample of lower first and second molars. TAXON C. plicatile C..plicatile C. ingenuum C. ingenuum C ingenuum C. emsliei FAUNA LOVE HEMP ARS LOVE HEMP UBV P2 apl 23.7,1.14,39 23.8, 1.11, 4 21.9, 0.84, 6 215,0.96,19 22.0,-, 1 19.4, 1.46, 3 H U LB E R T: C O R M O R H IP PA R IO N & H IP PA R IO N FR O M F LO R ID A 323 22.0-27.1,4.83 22.7-24.9,4.67 21.0-23.2, 3.82 20.2-24.1,4.48 18.1-21.0, 7.55 bapl 19.9, 0.92, 37 19.3,-,1 18.6, 0.33, 5 18.0, 0.65, 19 16.2, 0.57, 2 18.0-21.3,4.62 18.3-19.1, 1.76 16.9-19.6, 3.64 15.8-16.6, 3.49 atw 9.3,0.74,39 8.8,0.52,4 8.3,0.69,6 8.6, 0.66, 21 8.4,-,1 7.7,0.78,27.5-10.4, 7.97 8.5-9.6,5.88 7.7-9.5,8.24 7.7-9.9,7.70 7.1-8.2, 10.10 ptW 113, 0.91 , 41 11 .5 , 1 .03, 4 105, 0.55 , 6 10.6, 0 .86, 21 9.6,-, 1 9 . 2, 0.57, 29.6-13.3, 7.91 10.3-12.5,8.95 9.6-11.1,5.22 9.0-12.6,8.13 8.8-9.6, 6.15 mmi 10.2,1.13, 41 10.2,0.74,4 8.3,0.98,6 9.1, 0.99, 21 8.0,-,1 8.6,0.32.37.0-12.5, 11.03 9.2-10.8, 7.20 7.2-10.0, 11.76 7.5-11.7, 10.87 8.4-9.0,3.72 entl 9.8, 1.34, 41 9.8,1.63,4 9.5,0.66,6 8.7, 1.37, 21 10.1,-,1 8.8, 1.54, 36.1-12.0,13.62 7.6-113,16.64 8.3-10.2,6.95 53-10.6, 15.85 7.0-9.8, 17.54 p34 apl 22.8,1.34,48 21.4, 1.34, 9 20.9, 0.98, 21 19.5, 1.13, 35 19.7, 0.76, 3 19.3, 1.36, 919.2-25.6, 5.87 19.4-24.1,6.28 18.8-22.8, 4.63 17.6-22.1,5.81 19.2-20.6, 3.84 17.5-21.4, 7.08 Table 4 Continued. 324 B U LLE TIN FLO R ID A S TATE M U S E U M V O L. 33(5) TAXON C. plicatile C. plicatik C ingenuum C ingenuum C ingenuum C. emsliei FAUNA LOVE HEMP ARS LOVE HEMP UBV bapl 18.7, 0.71, 49 17.8,0.37,4 17.4,032,14 16.1, 0.74, 32 16.2,0.71, 2 16.1,1.07,5 17.0-20.6,3.82 17.3-18.2,2.08 163-18.1,2.99 14.4-17.8,4.60 15.7-16.7, 4.36 14.9-17.3, 6.69 atw 12.0, 0.98, 48 12.3,0.74,8 10.6,1.03,20 10.9,0.82,34 11.1, 0.21, 3 9.9,0.74,9 8.4-14.0,8.14 11.1-13.4,5.97 7.9-12.3,9.66 93-123,730 10.9-11.3, 1.88 8.6-11.1,7.50 ptw 12.0, 1 .01 , 48 12.0, 0.78, 8 10.6, 0.76, 18 10.6, 0.61 , 35 11 .2, 0.46, 3 9.5 , 0.76, 10 9.1-14.2,8.45 10.9-133,653 8.4-11.6, 7.21 8.9-11.8,5.77 10.8-11.7, 4.09 8.6-10.9, 8.00 mmi 13.1, 1.07, 48 12.9, 0.68, 9 11.9,1.03,22 11.3, 039, 35 12.1, 0.85, 3 10.7, 0.95, 10 9.9-15.3,8.14 11.9-14.0,5.24 9.2-135,8.67 10.0-12.6, 5.23 11.5-13.1, 7.03 9.2-11.5,8.92 entl 10.4, 1.02, 48 10.1, 0.95, 9 10.0,0.66, 22 9.1,1.03,35 9.1, 130, 3 9.3, 0.77,10 6.1-12.1, 9.88 8.0-11.2,9.35 7.6-11.1,6.67 6.1-10.8, 11.35 7.6-10.6, 1650 8.1-10.6, 8.23 m12 apl 22.2,2.03,68 21.3, 1.90, 15 20.2,1.47,26 19.5,1.63,43 18.2,1.82,4 19.1,1.62,18 18.6-26.3,9.15 18.4-24.4,8.91 17.9-23.3,7.25 17.0-233,8.39 16.2-20.4,10.02 15.9-22.0,830 bapl 17.4, 0.90, 64 17.8, 0.88, 8 16.1, 036, 18 15.3,0.89,41 15.6,-,1 15.5,1.04,8 15.4-20.2,5.15 16.9-19.4,4.94 15.1-17.0, 330 13.2-17.1, 5.79 14.1-16.5, 6.71 atw 10.2,1.08,68 10.7, 1.72, 16 8.9,0.89,24 9.1,1.07,44 9.9,0.70,5 8.3,0.65,19 8.142.6, 1032 8.1-13.4,16.14 6.8-103,10.00 6.241.9, 11.81 9.4-11.1,7.03 7.3-9.5,7.80 ptw 9.3,1.07,68 93,130,16 8.2,0.72,22 8.4,0.84,44 9.0,0.42,5 8.1, 0.81, 18 6.4-11.4,11.49 7.1-12.1, 15.78 6.6-9.1,8.72 63-10.0, 9.96 8.6-9.7,4.65 6.7-93, 10.01 mml 12.2,0.83,68 12.1, 1.34, 16 11.0,0.83,28 10.7, 0.65,44 11.1,0.69,5 10.4, 1.04, 19 103-14.3, 6.78 10.1-14.2,11.10 93-123,733 9.0-12.1, 6.03 10.3-11.8, 6.18 8.7-12.0, 9.92 entl 8.0, 1.21, 68 7.4, 1.21, 16 7.6,1.02,28 6.8,1.08,44 6.5,1.62,5 7.4, 0.89, 18 53-10.1, 14.08 5.5-9.8, 16.29 5.1-9.4, 13.39 43-9.0, 15.80 4.3-7.2,24.97 5.8-8.4, 12.Ott MANDIBLE p2-p4 L 69.4,2.81,9 663,-,1 63.3,1.75,4 59.7,2.26,6 -- 58.3, --,1 64.9-74.2,4.05 62.0-65.7,2.77 56.3-62.9,3.78 ml-m3 L 64.2, 1.91, 6 66.4,-,1 60.7,1.98,2 57.3,1.43,8 - 60.1,-,1 62.1-66.8,2.97 59.3-62.1,3.26 55.0-59.7,2.50 p2-m3 L 133.0,4.79, 3 132.6, -,1 124.2, 3.75, 2 116.1, 0.61, 4 - 117.7, -- , 1 H U LB E R T: C O R M O R H IPPAR IO N & H IP PA R IO N FR O M FLO R ID A 325 129.6-1383, 3.60 1213-126.8, 3.02 115.3-116.7.0.53 dp2-dp4 L 81.2, 1.43, 4 76.1,;1 - 74.7,1.43,3 --- - 79.3-823,1.76 73.1-75.9, 1.91 idl - 106.9,-,1 - 82.2,-,1 - 112.2,-,1 326 B U LLE nN FLO R ID A S TATE M U S E U M V O L. 33(5) TABLE 5. Frequency of occurrence (in %) of different enamel plications in the lower cheekteeth of various populations of Cormohipparion. PARA, posterior plication on paralophid; ANTI, plication on antisthmus or anterior side of isthmus; PSI'I, plication on postisthmus or posterior half of isthmus; PLCB, pli caballinid . The "Hemphillian" fauna for C ingenuum and C. plicatik refers to a combined sample from Mixson's Bone Bed, McGehee Farm, Haile 19A, Coffrin Creek, and the Lower Bone Valley. Species Fauna n PARA ANTI psn PLCB p3 and p4 combined C. occidentale Lapara Creek 24 49 33 50 58 C. occidentale Clarendon 17 26 35 65 59 C. occidentate Xmas-Kat 29 66 62 59 62 C. ingenuum Agricola 23 61 22 35 56 C. ingenuum Love Site 89 60 55 33 39 C ingenuum Hemphillian 6 50 50 33 50 C. plicatile Love Site 130 41 50 20 28 C. plicatile Hemphillian 15 53 73 40 47 C. enisliei Bone Valley 20 85 75 25 100 ml and m2 combined C. occidentale Lapara Creek 16 30 63 38 12 C. occidentale Clarendon 18 0 28 17 0 C. occidentale Xmas-Kat 30 27 63 30 3 C ingenuum Agricola 29 24 21 7 10 C. ingenuum Love Site 115 12 45 9 3 C. ingenuum Hemphillian 12 9 18 0 9 C. plicatile Love Site 165 12 52 2 7 C. plicatile Hemphillian 30 17 67 13 3 C. emstiei Bone Valley 27 33 78 22 56 TABLE 6. Comparison of variation in tooth length measured at the base of the crown (BAPL)of both single quarry and combined guam, (faunal) samples of hypsodont equid species. BAPL is statistically independent of crown height, and thus a better indicator of sample variation when comparing populations that may have different age structures than characters measured on the occlusal surface. Two measures of variation, CV (in %) and OR (in mm) are shown for combined samples of the upper third and fourth premolars (P34), and the upper first and second molars (M12). Numbers in parentheses following each species name are the sample sizes for the P34 and M12, respectively. Only populations with large sample sizes were chosen, except for Cormohipparion occidentale. It was included as it is the best available for a species congeneric with the two species of interest, C. plicatile and C. ingenuum. If the samples from the Love site of C. plicatile and C. ingenuum are combined (to create a single species), the observed variation is much greater than that observed in single quarry or even faunal samples of other species. This along with the bimodal distributions of measured characters (Figs . 11 and 12), and qualitative differences is justification for recognizing two species of Cormohipparion at the Love Site and other Florida localities. When treated separately, the variation each exhibits is equivalent to that observed in other quarry samples. References are Webb and Hulbert (1986), and Hulbert (1987a, 1987b, 1988a). P34 M12 H U LB E R T: C O R M O R H IPPAR IO N & H IP PA R IO N FR O M FLO R ID A 327 Quarry Taxon or Fauna CV OR CV OR Cormohipparion plicatile and C. ingenuum combined (153,155) Love Site 7.91 61 6.65 4.7 Single quarry samples Cormohipparion plicatile (107,108) Love Site 536 4.1 4.20 3.1 Connohipparion ingenuum (46,47) Love Site 4.52 2.7 3.90 2.3 Neoh