Peabody Museum

of Natural History

Yale University

New Haven, CT 06520

Postilla Number 175

2 May 1979

Diversification and Biogeography of the One-Toed Horses Onohippidium and Hippidion

Bruce J. MacFadden Morris F. Skinner

Received 14 June 1977 Abstract

In this report we describe a large sample of Onohippidium galushai, new species, from the Late Hemphillian (Early Pliocene) of west- ern Arizona, one ramus of Hippidion sp. from the Early Hemphillian (Late Miocene) of the Texas High Plains, and one ramus of cf. Hip- pidion sp. from the Irvingtonian (Late Pliocene to Early Pleistocene) of southern California. These one-toed horses were previously thought to have been confined to Plio- Pleistocene deposits of South America Onohippidium from Arizona POSSESSES a re- tracted nasal notch, deep preorbital facial fossae, and dental pattern diagnostic of South American representatives of this genus. Hip- pidion from Texas and California exhibit the dental characters, especially in the deep ec- toflexids, seen in South American representa- tives of this genus. This is the first report of these horses from North America. The pres- ence of Onohippidium and Hippidion in North America demonstrates that diversifica- tion of these ‘“endemics” occurred prior to dispersal to South America during the Pleis- tocene great faunal interchange

© Copyright 1979 by the Peabody Museum of Natural History, Yale University. All rights reserved No part of this publication, except brief quotations for scholarly purposes, may be reproduced without the written permission of the Director, Peabody Museum of Natural History.

Introduction

One of the most interesting groups of*fossil horses is Known from Plio-Pleistocene de- posits of South America. These horses, which include Hippidion, Onohippidium, and Parahipparion, are, so far as is known, charac- terized by unusual morphological adapta- tions. Both Hippidion and Onohippidium have retracted nasal notches (Fig. 1). Along with this retraction, the nasal bones remain elon- gate and unreduced “splints.” Some other mammals, such as tapirs, have retracted nasal regions; which is an adaptation to the presence of a proboscis. Retracted nasals

A

NASAL NOTCH |

NASAL SPLINTS \

B LACRIMAL FOSSA i\

BUCCINATOR FOSSA \

Fig. 1.

Skulls of Hippidion (A) and Onohippidium (B) show- ing characters discussed in text (adapted from Burmeister, 1875, and Sefve, 1912, respectively). The nasal splints in the Onohippidium specimen are broken anteriorly. Not to scale.

One-toed Horses

Postilla 175

A PREFOSSETTE POSTFOSSETTE

HYPOCONAL GROOVE

PREPROTOCONAL lh held "—HYPOCONE

POSTPROTOCONAL VALLEY

PROTOCONE

B ISTHMUS,

METASTYLID

METACONID ,

ENTOCONID

HYPOCONULID

PARALOPHID————

é HYPOCONID ren ere Ne ECTOFLEXID

Fig. 2.

Left upper (A) and left lower (B) cheek teeth of a one-toed horse showing characters discussed in text. Nomenclature follows Stirton (1941), Skinner and Taylor (1967), and Skinner (MS).

are rare in fossil horses, although this feature has been described for certain “Hipparion” from Eurasia which apparently possessed a | tapir-like proboscis during life (e.g., Sefve, 1927; Forstén, 1968; Sondaar, 1971). The only other reported occurrence of nasal modifica- tion in horses is in Onohippidium and Hippi- dion. The type of nasal modification seen in these two genera has not been observed in other horses. It is possible that these horses did not possess a proboscis because: 1) The nasal bones are not posteriorly retracted as seen in other mammals with a proboscis; 2) The elongate nasal splints would have re- stricted movement of soft anatomical struc- tures. Simpson (1951, p. 147-148) states that: “The nasal bones [in Hippidion]. . .must

have been well supported by a nasal septum of cartilage and other tissues. .. .” It seems that this type of nasal modification is related to olfaction, but a more specific function is not certain.

It is generally stated that Onohippidium

and Hippidion were derived from a group of one-toed horses from North America (e.g., Matthew, 1924; Stirton, 1940). The purpose of this report is to describe the first-known re- cord of these two genera in Hemphillian and Irvingtonian deposits of North America and to discuss their biogeographic significance.

NEW MEXICO

| Se ‘ 4 ee

Be colle

MEXICO

Fig. 3.

Fossil localities discussed in this report. 7. Unde- scribed local fauna, unnamed rocks of the Ogallala Group, Early Hemphillian (Late Miocene) of the Texas Panhandle. 2. Wikieup local fauna (I.f.), Big Sandy Formation, Late Hemphillian (Early Pliocene) of western Arizona. 3. Sediments probably equiva- lent in age to the Vallecito Creek |.f., Irvingtonian (Late Pliocene-Early Pleistocene) of the Anza- Borrego State Park of southern California.

Abbreviations

The following institutions are referred to in the text:

F:AM Frick: American Mammals, De- partment of Vertebrate Paleontology, The American Museum of Natural History

LACM Section of Vertebrate Paleontol-

ogy, Los Angeles County Museum of Natural History

3 One-toed Horses

Postilla 175

All measurements of specimens are in mil- limeters and are abbreviated as follows:

A=P Greatest anterior-posterior length

il Greatest transverse width (labial-lingual; external- internal)

R Right side of tooth row

L Left side-of tooth row

@ Measurement approximate

The dental nomenclature used in this paper and illustrated in Figure 2 follows Stirton (1941), Skinner and Taylor (1967), and Skin- ner (MS),

Systematic Paleontology

Class Mammalia Linnaeus, 1758 Order Perissodactyla Owen, 1848 Family Equidae Gray, 1821

Genus Onohippidium Moreno, 1891

Revised Distribution. Early Pliocene (Late Hemphillian) of Arizona and Late Pliocene and Pleistocene of South America (Sefve, 1912; Hoffstetter, 1952; Webb, 1976; this re- Port).

Generic Diagnosis. Horse with nasal notch that lies as far anterior as P?-P4 and as far Posterior as the orbit. Nasal bones not re- tracted and remain elongate splints. Well-

developed dorsal lacrimal fossa and no ven-

tral malar fossa. Incisors with cement-filled Cups (infiditula). Protocone oval to rounded with weak preprotoconal groove and well- developed postprotoconal valley. Lower Cheek teeth with either shallow ectoflexids (externomedial valleys) and isthmuses in the Permanent premolars or deep ectoflexids in the deciduous premolars and molars. Very Simple enamel plications. Monodacty! Metapodials.

pnohippidium galushai, new species ig. 4,

Fig. 4.

Onohippidium galushai, new species, from the Late Hemphillian of Arizona. A. F:AM 100077, holotype, left lateral view of partial skull. B. F:AM 100077, holotype, left upper dentition. C. F:AM 100078, right lower dentition, reversed.

Pliohippus sp., Lindsay and Tessman, 1974, p.6.

Holotype. F:AM 100077, partial skull with well-preserved dentition.

Hypodigm. Hundreds of specimens, which consist of skulls, jaws, dentitions, and post- cranials in the F:AM collection. There are specimens possibly referable to this species in other institutions, such as the small collec- tions from the same locality at the University of Arizona Laboratory of Paleontology, but these were not examined during the present re- search.

Locality and Age. Frick field crews collected the F:AM specimens during the 1950s from the Big Sandy Formation of Mohave County, Arizona (Fig. 3). This formation, which con- sists of lacustrine clay to marginal sand facies deposited in an enclosed basin, outcrops along the Big Sandy River from approximately 8.5 km north to 12 km south of the town of Wikieup. Sheppard and Gude (1972) present a detailed stratigraphic description of this formation. Frick field crews collected a few specimens from washes north of Wikieup, but the bulk was collected from a locality approx-

4 One-toed Horses

Postilla 175

imately 12 km south of Wikieup, near the southern outcrop margin. This locality is in- cluded in Sheppard and Gude’s (1972, p. 6) reference section R-3 located“... along a wash that parallels Signal Road about 2,600 ft northwest of the road, from NWY%NWY%SW% sec. 29to NEY@SEVANW'% sec. 29, T. 15N.,R. 12 W., Greenwood Peak quad.” The fossil- bearing zone is located in the upper part of reference section R-3 (Sheppard and Gude, 1972: Galusha, MS). Galusha (MS) terms this zone the ‘Clay Bank Quarry locality,” whichis used in a broad sense to include fossils col- lected locally within this zone. One large con- centration within this locality is approximately 125 mwest of Clay Bank Quarry and is termed “Bird Bone Quarry.” This fossil-bearing local- ity and zone can be relocated with the use of Frick field notes (Galusha, MS), Sheppard and Gude (1972), and the U.S. Geological Survey 7¥2'’ Greenwood Peak quad. During the falls of 1976 and 1977 MacFadden and Dr Noye M. Johnson (Dartmouth College) relo- cated the Clay Bank Quarry locality and col- lected samples for paleomagnetic and fission-track analysis.

The mammalian assemblage from the Big Sandy Formation is informally termed the Wikieup local fauna (I.f.) in Tedford et al. (in press). Apreliminary faunal list for specimens collected by the University of Arizona field crews is presented by Lindsay and Tessman (1974). The Wikieup Lf. is largely undescribed although some comparisons with as- semblages of similar age have been made (e.g., MacFadden, 1977; Tedford et al., in press). Tedford et al. (in press) consider the age of the Wikieup |.f. to be Late Hemphillian (Early Pliocene), which corresponds to an ab- solute time interval of approximately 4 to 5 million years ago.

Etymology. Named in honor of members of the Frick Laboratory, Theodore and Marian Galusha.

Specific Diagnosis. Same as for genus with the following distinctions: Nasal notch re- tracted to above P3-P4. Fossa between buc- cinator and lacrimal fossae not as well de- veloped as other Onohippidium (Fig. 4).

Description. The nasal notch is retracted posteriorly to a position which lies over P°-P4 and is intermediate between the conditions seen in North American Pliohippus and Dinohippus and in South American Hippidion and Onohippidium

There is a well-developed complex of fossae on the cheek. The buccinator fossa Is an- teriormost and in front of the cheek-tooth row. Posterodorsal to the buccinator fossa there is a faint preorbital depression; this is better de- veloped in South American Onohippidium. Posterior to this faint preorbital depression there is a deep and well-defined lacrimal fossa that is also seen in South American Onohippidium. The infraorbital foramen lies above P4—M".

The upper dentition of O. galushaiis very simi- lar to South American Onohippidium. The in- cisors have cement-filled cups. P' is reduced in size. The premolars are larger in cross sec- tion than the molars. The protocone is rounded to oval with a weak preprotoconal groove and a deep postprotoconal valley. Occasionally, especially in the molars, the protocone unites with the metaloph and hypocone forming a cement-filled enclosure. The hypocone is oval and usually slightly smaller than the protocone and there is a dis- tinct posterolateral hypoconal groove. The pre- and postfossettes (lakes) are cresent- shaped. The plications on the fossettes are absent except for the posterior border of the prefossette and anterior border of the post- fossette.

The lower incisors have cement-filled cups. The P1 and its alveolus are absent, which is characteristic of advanced horses. The pre- molars are usually larger than the molars. The deciduous premolars show relatively deep ectoflexids, between the protoconid and hypoconid; however, these ectoflexids are not deep enough to divide the isthmus that con- nects the metaconid and metastylid to the protoconid and hypoconid, respectively. The permanent premolars have relatively shallow ectoflexids that do not divide the isthmus. The

5) One-toed Horses

Postilla 175

enamel pattern is simple with few secondary plications.

Discussion. The retracted nasal notch with unreduced nasal splints, configuration of preorbital fossae, and, to a lesser extent, sim- ple dental pattern provide convincing evi- dence for relating O. ga/ushai to the South American representatives of this genus. So far as is known, Onohippidium has not been iden- tified elsewhere in North America. Other con- temporaneous horses for which skulls are known, i.e., Pliohippus and Dinohippus, have characters different from those seen in Onohippidium. Pliohippus has a different con- figuration of preorbital fossae and no re- tracted nasal notch, whereas Dinohippus has

a simple preorbital area and no retracted nasal notch. There are numerous forms of the Pliohippus-Dinohippus complex for which the preorbital and nasal areas are not known. Therefore, it would be most satisfying if the Pliohippus-Dinohippus complex could be dif- ferentiated from North American Onohip- pidium solely on the basis of dentition. If this dental differentiation were possible, then Onohippidium might be identified at other Late Hemphillian North American localities. However, the dentitions of these contem- poraneous forms are so similar, and there is so much variation within “populations,” that generic differentiation on this basis is virtually impossible.

Fig. 5.

. 0 7 w + MILLIMETERS

Occlusal and lateral photographs of symphysis and left ramus of Hippidion sp., F:AM 101439, from the

Early Hemphillian of the Texas High Plains.

Genus Hippidion Owen, 1869

Revised Distribution. Early Hemphillian (Late Miocene) of Texas, Irvingtonian (Late Pliocene to Early Pleistocene) of California, and Late Pliocene to Pleistocene of South America (Sefve, 1912; Hoffstetter, 1952; Webb, 1976: this report).

Generic Diagnosis. Horses with nasal notch that lies above M'-M?. Nasal bones not re- tracted and they remain elongate splints. Dor- sal lacrimal and ventral malar fossae absent. Incisors with cement-filled cups. Protocone oval with weak preprotoconal groove and well-developed postprotoconal valley. Lower cheek teeth with deep ectoflexids in the pre-

6 One-toed Horses

Postilla 175

molars and molars. Enamel plications usually more complex than in Onohippidium. Monodacty! metapodials. For diagnostic characters not represented in specimens dis- cussed in the present report, see Burmeister (1875), Sefve (1912), and Hoffstetter (1952).

Hippidion sp. Figs. 5, 6, Table 1.

Referred Specimen. F:AM 101439, left ramus and symphysis with R I,-1,, L };-Ms3

Locality and Age. The Frick field crew of 1941 collected this specimen from unnamed rocks of the Ogallala Group, 10 miles north of Hig- gins, Lipscomb County, Panhandle of Texas (Tedford, personal communication, 1976; see Fig. 3). The fauna from this locality is unde- scribed, but itis similar in age to the Arnett |.f. from adjacent Oklahoma described by Kitts (1957). It is stratigraphically below the Hig- gins I.f. described by Hesse (1940). Tedford etal. (in press) consider the age of this locality to be Early Hemphillian (Late Miocene). Description. |, and|,have cement-filled cups exposed at the occlusal surface (Fig. 5). 1, has cups recessed below the occlusal surface. There is no precanine diastema and C is closely appressed to |3. Sexual dimorphism is pronounced in fossil horse populations and the small canine size in F:AM 101439 indi- cates that this individual is a female. Posterior to the symphysial dentition the ramus |s later- ally constricted for the length of the post- canine diastema, which measures 69.6 mm from the posterior base of C to the anterior base of P,. The mental foramen is approxi- mately midway between C and P». The P; and its alveolus are absent.

The molars are narrower transversely than the premolars. The dental pattern of F:AM 101439 (Fig. 6) is similar to that of Hippidion illustrated by Burmeister (1875). Pz is triangular-shaped and similar in dental pattern to the rest of the molars except for the addition of the anteriormost conid, which is characteristic of the P, in horses, whereas P,, through M; have an anterointernal paralophid. The protoconid

and hypoconid are exteriorly convex. The ec- toflexid is relatively deep in the cheek teeth of F:AM 101489; this character is also seen in other Hippidion such as those described by Burmeister (1875), Sefve (1912), and Hoffstet- ter (1952). When the ectoflexid is very deep it subdivides the isthmus into the antroisthmus and postisthmus (Skinner, MS). The metaconids and metastylids are rounded and subequal in size for a given tooth. There is variation in the metaconid-metastylid com- plex within the tooth row.

Fig. 6. Occlusal view of Hippidion sp., F:AM 101439, show- ing details of dental pattern (see Fig. 5).

InP,-P,and M2-M,the metaconid is joined to the protolophid by a constricted antroisthmus and the metastylid is joined to the hypoconid by the postisthmus. In M, the isthmus is not as constricted as in the other teeth and the metaconid-metastylid complex is smaller and more poorly defined than the other molars. This variation in dental pattern of M, is proba- bly a result of differential tooth wear. In the premolars, the hypoconid and entoconid are separated by a constriction. These same parts are less distinct in the molars. In the Ms, posterior to the hypoconulid, there is an iso- lated conid that is also seen in other Hippi- dion. This conid connects to the hypoconulid during later stages of wear. F:AM 101439 is similar in size to the smaller South American Hippidion species. For dental and size differ- ences, Figures 4and5, and Table 1 should be compared to the illustrations and measure- ments of Burmeister (1875), Sefve (1912), and Hoffstetter (1952)

Discussion. The dental characters of F:AM 101439, especially in the diagnostic deep ec- toflexids in both premolars and molars, pre- sent aconvincing argument for its assignment

i One-toed Horses

Postilla 175

to Hippidion. The specific identification of the Texas Hippidion is best deferred to a study that incorporates other relevant specimens The significance of this specimen with respect to the biogeography of South American “endemic” hoses will be discussed below.

Cf. Hippidion sp. Fig. 7; Table 1.

Referred Specimen. LACM 3363, incomplete left ramus with fragments of three cheek teeth, probably P3—M, (Fig. 7)

Locality and Age. This specimen was col- lected from LACM locality no. 1245-15 in sed- iments probably equivalent to the Vallecito Creek I.f. (Downs and White, 1965; Downs, personal communication, 1977). This local fauna occurs in the Matuyama reversed polar- ity epoch below a normal event that repre- sents either the Réunion’ or Olduvai. These Constraints indicate an Irvingtonian (Late Pliocene to Early Pleistocene) age for the cf. Hippidion specimen that is bracketed by the Gauss-Matuyama boundary at about 2.4 mil- lion years ago and either the base of the Ré- Union event at about 2.1 million years ago or the base of the Olduvai event at about 1.85 million years ago (Lindsay, Johnson, and Op- dyke, 1975; Opdyke et al., 1977). Description and Discussion. LACM 3363 is very poorly preserved. The teeth, probably P-M,, are less hypsodont than contemporary Equus. The dental pattern shows deep ecto- flexids in both the premolars and molars, and, therefore, we refer this specimen to Hippidion The principal importance of LACM 3363 is in the recognition of cf. Hippidion from Irving- tonian sediments of North America.

Biogeography

The origin of the South American “endemic” horses has been a matter of some interest in the past. Many students of fossil mammals,

Including Matthew (1924) and Stirton (1940), have recognized that this group was of North

pee eee ee

a

Fig. 7.

Occlusal view of cf. Hippidion sp., LACM 3363, ?P4-M,, from the Irvingtonian of southern California. Enamel is blackened for contrast. Broken areas are indicated by dashed lines (modified from an illustra- tion prepared by Ms. F. Runyan of the LACM).

American origin. Horses are not found in South America until the Uquian (Late Pliocene to Early Pleistocene), roughly 2 million years ago (Marshall et al., 1977), and their appear- ance represents dispersal from North America during the opening of the Central American land bridge (Webb, 1976). Reed (1950, p. 76-77), in his discussion of the Milk Creek |.f. of Arizona, states that: “The horses appear to be related to Pliohippus [Dinohip- pus], and it has been suggested by Dr. R. A. Stirton of the University of California that they may be ancestral to the South American Hip- pidium [sic] of the Pleistocene.” The great “morphological distance” between these North and South American horses has given rise to speculation as to structural and phylogenetic intermediates. Stirton (1940, p. 192) states that: “Upper Pliocene representa- tives of this group, when found, should display a moderately restricted narial notch in con- trast to the extreme observed in Hippidium [sic].” As predicted by Stirton, morphologi- cally intermediate forms are now known from North America as a result of the present re- port

‘The existence, duration, and boundaries of the Réunion event are presently uncertain (see, e.g., Opydke, 1972, and Watkins, 1972)

8 One-toed Horses Postilla 175

Table 1.

Tooth measurements (mm) for Hippidion.

Hippidion sp., F:AM 101439

Ri, Rl, Bale sa elk ele LC, lbedeaee) Leelee elie ee ee ESR lois ile Wh A-P L@e qa) 0) 10.6 Hales ilies TAS@-s2@ Cie 2000. 2 es Oe cine. 20.) v ins 14.6 TZ0 12.4 Wh! ilo: <4ee TAO) = OAR Os esos ee elaine Cf. Hippidion sp., LACM 3363 oleae (Bales bE A-P 30.2 28.4. 27.6@ ele abayiay Water SNS)

Abbreviation: @ = approximate.

As presently recognized, the one-toed horses include the genera Pliohippus s. 7. (including Astrohippus), Dinohippus, Onohippidium, Hippidion (including Parahipparion), and Equus s. 1. The phylogenetic relationships of these genera have not been dealt with adequately even in the classic studies of Sefve (1912), Osborn (1918), Stirton (1940), and Hoffstetter (1952). In fact, it might be shown that the taxon “one-toed horses’ is not a strictly natural, i.e., monophyletic group. Based on the limited sample discussed inthis report, we feel that a discussion of the interre- lationships of Onohippidium, Hippidion, and other one-toed horses should be deferred to a more comprehensive study.

The major significance of the present study is in the biogeography of Onohippidium and Hippidion in light of the specimens described here. The time of differentiation of Onohip- pidium and Hippidion from their closest rela- tives has not previously been discussed. lt has been implied that diversification obvi- ously occurred either in North America before dispersal southward or in South America after dispersal southward. We conclude that: 1) Clearly differentiated forms of Onohippidium and Hippidion are found in Hemphillian (Late Miocene to Early Pliocene) deposits of North

America. 2) These genera subsequently dis- persed to South America apparently during the Pleistocene faunal interchange. 3) Cf. Hippidion sp. is also found in Irvingtonian sed- iments in North America

Acknowledgments

We thank Dr. Richard H. Tedford and Mrs. S. Marie Skinner of the American Museum of Natural History for their encouragement dur- ing this research. Dr. Tedford brought the specimen of cf. Hippidion sp. to our attention and Dr. Theodore Downs of the Los Angeles County Museum of Natural History kindly loaned it for study. Messrs. Grant E. Meyer and David Frailey offered criticisms that im- proved earlier versions of the manuscript. Messrs. Raymond J. Gooris and William K. Sacco prepared Figures 4 and 5 respectively. Mrs. Brigitte Meshako and Ms. Betsy Dabakis typed portions of the manuscript. MacFadden conducted some of this research during te- nure of a faculty fellowship in the Department of Geological Sciences at Columbia Univer- sity. The University of Florida contributed funds toward the publication of this research.

9 One-toed Horses Postilla 175

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Downs, Theodore and John A. White. 1965. Late Cenozoic vertebrates of the Anza-Borrego Desert Area, Southern California. Amer. Assoc. Adv. Sci., Sect. E, Abs. Pap.: 10-11.

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Opdyke, Neil D. 1972. Paleomagnetism of deep-sea cores. Rev. Geophys. Space Phys. 10: 213-249. Opdyke, Neil D., Everett H. Lindsay, Noye M. Johnson, and Theodore Downs. 1977. The paleomag- netism and magnetic stratigraphy of the mammal-bearing section of Anza-Borrego State Park, California Quat. Res. (New York) 7: 316-329.

Osborn, Henry Fairfield. 1918. Equidae of the Oligocene, Miocene, and Pliocene of North America Iconographic type revision. Mem. Amer. Mus. Nat. Hist., New Ser., 2: 1-326

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One-toed Horses Postilla 175

Stirton, Ruben A. 1940. Phylogeny of North American Equidae. Univ. California Publ. Geol. Sci. 25: 165-198.

—. 1941. Development of characters in horse teeth and the dental nomenclature. J. Mammal. 22 434-446

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Unpublished References

Galusha, Theodore. MS. Unpublished Frick field notes. On file at the Department of Vertebrate Paleontolo- gy, The American Museum of Natural History, New York.

Skinner, Morris F. MS. Unpublished manuscript on equid dental nomenclature. On file at the Department of Vertebrate Paleontology, The American Museum of Natural History, New York.

The Authors

Bruce J. MacFadden. Formerly of: Depart- ment of Geology and Geophysics, Yale University, New Haven, Connecticut 06520. Present address: Fiorida State Museum, Uni- versity of Florida, Gainesville, Florida 32611.

Morris F. Skinner. Department of Vertebrate Paleontology, The American Museum of Natural History, New York, New York 10024.