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Pleistocene peccaries (mammalia: Tayassuidae) from western Oklahoma.

Fossil peccaries occur widely in Pleistocene localities across south-temperate North America. They are common in the eastern United States, especially in cave deposits. However, they are much less common as fossils in the Great Plains and southwestern United States. Peccaries have been discovered or reported rarely in Oklahoma. For this reason, several new records from western Oklahoma in the Oklahoma Museum of Natural History (OMNH) are of interest.

Previous records of peccaries of late Pleistocene age in the North American Southwest all are referable to two extinct species, Platygonus compressus (flat-headed peccary) and Mylohyus nasutus (long-nosed peccary). The extant collared peccary (Pecari tajacu) has no Pleistocene record in the United States, where it is known only from archaeological sites of late Holocene age. Its Pleistocene distribution was apparently much farther south, where the species has a Pleistocene fossil record in Guatemala (Woodburne, 1969). In the southwestern United States (California, Arizona, New Mexico, Texas, and Oklahoma) and northwestern Mexico there are about 26 late Pleistocene localities of occurrence for Platygonus and 2 for Mylohyus. Platygonus compressus is known from four sites in California (Diamond Valley Lake, McKittrick, Potter Creek Cave, Rancho La Brea), from three localities on the Colorado Plateau in northern Arizona and western New Mexico (Pit Stop Quarry, Navajo Lake, Sheep Camp Shelter), in five sites in the Basin and Range physiographic province of northwestern Mexico and southern Arizona and the transition zone between the Basin and Range and Colorado Plateau provinces in southwestern New Mexico (Terapa, Sonora, as cf. Platygonus, Murray Springs Arroyo Clovis site; Papago Springs Cave; Ventana Cave, Shelton Canyon), from one site in the Rio Grande Rift Valley (Dry Cave), in two sites on the High Plains of eastern New Mexico (Perico Creek, Blackwater Draw), in five cave sites in the Edwards Plateau of central Texas (Friesenhahn Cave; Hall's Cave; Laubach Cave 1, 3, and 5), and in five open sites in northwestern Texas (Aubrey, Canyon, Kickapoo, Lubbock Lake, Red Springs; Skinner, 1942; Colbert, 1950; Slaughter, 1966; Jefferson, 1991; Dalquest and Schultz, 1992; Harris, 1993; FAUNMAP, 1994; Lucas and Smartt, 1995; Mead et al., 2005, 2006; Morgan and Lucas, 2005; Murray et al., 2005; Hemmings, 2007; Springer et al., 2009).

Mylohyus nasutus was primarily an eastern North American species whose range extended westward only to the central and southern Great Plains. A single record of Mylohyus from the Chihuahuan Desert in trans-Pecos Texas at Fowlkes Cave was based on two fragments of a tusk that were not compared with Mylohyus (Dalquest and Stangl, 1984:449); these fragments probably actually represent Platygonus not Mylohyus. Mylohyus nasutus has been reported from Ben Franklin site, Clear Creek, Friesenhahn Cave, and South Sulphur River, Texas, and from Kanopolis and Sandahl, Kansas (Lundelius, 1960; Semken and Griggs, 1965; Hibbard et al., 1978; FAUNMAP Working Group, 1994).

[FIGURE 1 OMITTED]

In Oklahoma in the late Pleistocene (Rancholabrean North American Land Mammal Age), there are only two records of peccaries. Strain (1937) listed a thoracic vertebra (formerly catalog number S-470; now OMNH 58500) as Platygonus from the C. E. Smith gravel pit (OMNH locality V527), Chickasha, Grady County, Oklahoma, as part of the Chickasha local fauna (Smith and Cifelli, 2000). The vertebra lacks the neural spine and the centrum and neural arch are abraded badly; comparison with known thoracic vertebrae of P. compressus from the Arkansas Ozark Highland shows the specimen from Chickasha to be far larger and heavier. It is too large to represent a peccary and probably represents some other artiodactyl or a perissodactyl. Johnston (1935) provided the only published report of an ice-age peccary in Oklahoma, a M. nasutus based on a rostrum with upper canines and well-worn cheek teeth. The specimen (formerly catalog number 604, now OMNH 5740) came from a gravel bed of Pleistocene age near Alva, Woods County. Except for the dubious record from trans-Pecos Texas, the occurrence at Alva, Oklahoma, is the westernmost locality for the species.

Given the paucity of material, it is useful to record herein three additional occurrences of late Pleistocene peccaries in Oklahoma based on specimens in the OMNH vertebrate paleontology collection. Two of these are isolated occurrences and one is associated with two other taxa of vertebrates (Fig. 1).

Rush Creek--A single isolated specimen from OMNH locality V1638 bears an old catalog number 20-4-S7 dating from the 1930s and 1940s when J. Willis Stovall, curator and director, amassed many fossils of vertebrates for the University of Oklahoma Museum of Geology. An old catalog card corresponding to that number gives the locality incorrectly as "9 miles East and 2 miles South of Rush Springs, Grady County." However, in the OMNH vertebrate-paleontology archives are two letters of correspondence between the apparent discoverer-collector of the specimen and Stovall. According to the letter of 17 April 1939 from R. H. Bond to Stovall, the "fossil was found in ditch along north side of road, approximately 100 yards [approximately 100 m] west of lane leading to house in south central part of section 24, T.3N., R.6W. This location is 5 miles [8 km] south and 10 V miles [16.8 km] east of Rush Springs." A small amount of yellowish red silt and gravel fills the interior of the broken dentary beneath the teeth, a matrix that is clearly derived from the Permian red beds of the Whitehorse and El Reno groups that predominate as surface bedrock throughout Grady County and the region.

[FIGURE 2 OMITTED]

This specimen, recently recataloged as OMNH 73929, is a piece of the right dentary with the posterior end of the diastema and the p2-p4 (Fig. 2). The dentary clearly is referable to Platygonus based on the nonmolariform premolars that have high anterior cusps and low posterior cusps. Length of the premolar row is 31.1 mm; other measurements (Table 1) indicate the specimen is comparable in size to P. compressus from other localities (e.g., Gidley, 1920; Lucas and Smartt, 1995). No other remains or taxa were found at the site.

South of Yukon--An isolated specimen was found south of Yukon, Canadian County, Oklahoma (OMNH locality V1639). According to a note written with ball-point pen in blue ink on a brown paper bag in which the fossil was stored originally, the specimen is "from bed of creek in situ 20 ft [6 m] below surface South of Youkon [sic]." It was found by J. M. Paden of Oklahoma City. Although undated, the writing probably dates to the 1940s or 1950s. No other data were associated with the specimen and no correspondence pertinent to it could be found in archives of the OMNH vertebrate paleontology collection. Internally, the fragments of jawbone are filled with yellowish-brown silt and clay, stained with limonite as is common in many late-Neogene and Quaternary fossils of vertebrates in western Oklahoma.

The specimen, OMNH 73930, is a right dentary of P. compressus shattered in many pieces; one of the larger pieces includes the posterior end of the diastema, empty alveoli for p2, and two teeth, the p3 and p4. Another piece of the dentary contains the broken m2 and intact m3 (Fig. 3). All of the teeth are well worn, indicating an old but not geriatric individual. Measurements of the teeth are provided in Table 1. No other remains or taxa occurred with the dentary fragments.

[FIGURE 3 OMITTED]

Quartz Mountain Canal--A small collection of fossils including three taxa (prairie dog, mammoth, and peccary) was found ca. 0.4-0.8 km N and 1.5 km W Blair, Jackson County, Oklahoma, by Lawrence LeVick in about 1992. The bones were found during dragline operations on a small drainage canal, about 1 m deep in the canal. This is one of many small canals in the area associated with establishing the system (West Canal, Blair Lateral, and Altus Canal) for irrigation and providing a water supply to the city of Altus and Altus Air Force Base. The author visited Lawrence and Jean LeVick at their home near Blair on 10 August 2007 and on the same day visited the locality with Lawrence LeVick and Don Wyckoff. The site (OMNH locality V1632) as shown to me is in the SW 1/4 section 30, T4N, R20W, at an elevation of ca. 442 m. The locality is southwest of the Quartz Mountains, a small cluster of hills in the western part of the Wichita Mountains Uplift at the confluence of the Elm Fork Red River and North Fork Red River. The Wichita Mountains are a chain of exposed Cambrian-age intrusions of granite and other igneous rocks in southwestern Oklahoma. The North Fork Red River is dammed as Lake Altus-Lugert where the river crosses the Quartz Mountains.

The locality was in the drainage of Bitter Creek, a small tributary of the Salt Fork Red River on the outwash plain of the Quartz Mountains, in relatively flat agricultural land crossed by numerous artificial canals. Lawrence and Jean LeVick told me that Lawrence originally found all of the fossils together in a cantaloupe-sized mud ball that was dredged out of the irrigation canal and dumped on the south side of an east-west-trending section of the canal. Lawrence LeVick, an avocational archeologist, interpreted the mud ball as a rodent burrow remnant or something similar. He found no other bones during a search of the dredged spoil dirt along the canal. The canal was flowing full on the day we visited in 2007 and appeared to have been dredged clean not long before the visit. The mud cleaned out of the ditch on the south side appeared as medium-brownish-gray silt. Lawrence LeVick said the ditches were dredged often and the ditch at the locality is ca. 1 m deep, so the fossils had to have come from a level within ca. 1 m of the surface. Cannon (1967) mapped Pleistocene middle-terrace deposits along the North Fork Red River including several ca. 4 km to the east, 7 km to the northeast, and 9 km to the northwest of the locality. The fossils from Quartz Mountain canal were passed to Don Wyckoff by Lawrence and Jean LeVick, and then to the author for deposition in the OMNH in June 2007.

A small, flat fragment of enamel plate ca. 2.5 mm thick and ca. 21 by 17 mm and a fragment of unerupted, developing crown-cap segment represent pieces of cheek tooth from a mammoth, Mammuthus, of indeterminate species. The pieces are cataloged as OMNH 73932. Little can be said of the fragments except that they confirm the presence of this member of the paleofauna at Quartz Mountain Canal.

Also found at the same time in the Quartz Mountain Canal was a complete right humerus of a prairie dog, Cynomys species indeterminate (OMNH 73933). The bone is from a subadult as evidenced by the incompletely fused proximal epiphysis. It is 44.4 mm in length, 9.1 mm in proximal width, and 12.0 mm in distal width. The specimen shows different preservation and discoloration than the peccary bones and mammoth tooth fragments, being more permineralized rather than leached (as the peccary bones are). This suggests the prairie dog might not be contemporaneous with the peccary. The burrowing habit of prairie dogs could have introduced the bone intrusively into the deposit in which the peccary and mammoth were preserved. Radiometric dating of each taxon (if possible; the bones may be leached of their organic content) or rare earth element (REE) analysis could potentially determine contemporaneity of the three specimens.

Several pieces from the Quartz Mountain Canal definitely represent Tayassuidae, although they cannot be unequivocally referred to either Platygonus or Mylohyus. Tayassuidae and related Old World pigs (Suidae) have four-toed feet, usually with separate metapodials, unlike other artiodactyls in which the central metapodials are completely and solidly fused into a cannon bone. The side toes are reduced in tayassuids more than in suids. In suids, the central metapodials (of digits III and IV) are unfused and the side toes (digits II and V) are well developed. In tayassuids, metacarpals III and IV are slender and unfused; metatarsals III and IV usually are fused at the proximal end (Lundelius, 1960:22-23). In the late-Pleistocene long-nosed peccary M. nasutus, the side toes of the manus are present but reduced, while in the pes, the metatarsal and phalanges for digit II are present but digit V is absent. As a result, M. nasutus had medial and lateral dew claws on each front foot, but only one (medial) dew claw on each hind foot. In the late-Pleistocene flat-headed peccary P. compressus, the side toes are reduced completely, and the species did not have dew claws. Tayassuids also have vertically oriented, relatively straight canines, unlike the outwardly and upwardly curved canines of Suidae.

[FIGURE 4 OMITTED]

The peccary material from Quartz Mountain Canal consists of a partial canine, a small skull fragment with an occipital condyle, and several associated bones ofa manus or pes (OMNH 73931). The canine is a rather poorly preserved right upper tusk mostly consisting of the root, but a small part of the base of the crown is present (Fig. 4). The preserved portion lacks enamel but still shows a small remnant ca. 1 cm long of the anterior wear surface in the dentine of the base. The surface is caused by wear against the posterior face ofthe lower canine, as is typical of peccary tusks. The tusk is relatively robust, heavier than the canines in the muzzle (OMNH 5740) of M. nasutus from Alva. In the fossil from Quartz Mountain Canal, anteroposterior diameter at base of C1 is 20.1 mm and transverse width is 12.0 mm, contrasting with 15 and 11 mm for these measurements, respectively, for the Mylohyus from Alva mentioned above (Johnston, 1937). The canine from Quartz Mountain Canal is much heavier than a subadult C1 of P. compressus from a cave in the Arkansas Ozark Highland that also was available for comparison. Tusks of peccaries are sexually dimorphic with those of males larger than females, and thus, the specimen from Quartz Mountain Canal potentially came from a male but does not help differentiate between Mylohyus and Platygonus. The occipital condyle is of appropriate size and shape to represent the same peccary as the tusk and foot bones.

The foot skeleton from Quartz Mountain Canal consists of the distal portions of metapodials III and IV, including the distal articular ends and short lengths of the shafts, both associated first and second phalanges, and one of the two ungual phalanges (Fig. 4c). The metapodials are separate (i.e., unfused) at least at the preserved distal ends. Metatarsals normally are fused in Platygonus and Mylohyus, although the fusion is mainly at the proximal ends in Mylohyus. Metacarpals are fused proximally or along their entire length in Platygonus, and unfused in Mylohyus. It is not possible to determine definitively whether the bones represent the manus or pes. However, because the metapodials show no sign of fusion in the preserved distal portions, the bones possibly represent the metacarpals of a manus and tentatively are interpreted as such herein. Additionally, it is not possible to determine definitively the side of the animal from which the bones came. However, when the distal metacarpal bones are manipulated, the best fit between them is felt when the left of the two metacarpals extends distally slightly beyond the right (relative to the axis of symmetry within the foot). This is the normal configuration in a left foot (relative to the axis of symmetry of the animal); in tayassuids and suids, the lateral metacarpal IV and metatarsal IV are slightly longer than the medial metacarpal III and metatarsal III. Thus, the bones from Quartz Mountain Canal tentatively are considered to represent a left manus, and elements are listed and measured accordingly (Table 2). The tip of the only preserved ungual phalanx is broken.

The foot elements, although diagnostic as Tayassuidae, are less easily assigned to a genus due to a paucity of distinctive features. If the foot is indeed a manus as interpreted herein, the ungual phalanges might be of some importance. Lundelius (1960:22) noted that the manual ungual phalanges of Platygonus are flattened dorsoventrally, whereas those of Mylohyus are flattened laterally. The sole available ungual phalanx from Quartz Mountain Canal is relatively flattened dorsoventrally. Thus, the specimen might represent Platygonus, but few comparative specimens were available to the author to match directly and confirm or refute this observation. When central metapodials of peccaries fuse, they usually do so proximally first and might not fuse at the distal end. Because only distal ends of the metapodials from Quartz Mountain Canal are preserved and are completely unfused, they might represent manual elements (metacarpals) from a Mylohyus (rather than the pedal elements or metatarsals of that genus), or they might represent an immature individual of either Mylohyus or Platygonus in which the metapodials had not yet fused distally. Some authors have noted that metapodials III and IV of Platygonus are stouter than those of Mylohyus (Lundelius, 1960), but the small sample and incompleteness of the metapodials from Oklahoma, as well as potential sexual dimorphism in size, precludes confirmation of identity based on comparative size. In summary, morphology of the preserved skeletal elements is inconclusive for assigning a genus to the specimen from Quartz Mountain Canal.

I extend thanks to D. G. Wyckoff for many discussions of Pleistocene matters and visits to vertebrate-bearing localities in Oklahoma and to L. and J. LeVick for kindly showing me the locality at Quartz Mountain Canal and for donating the fossils of vertebrates to the OMNH through D. G. Wyckoff. Likewise I acknowledge the others whom I never knew but who donated their finds to the OMNH for safekeeping over the generations, and the animals of whom knowledge comes through their fossils. Finally, I thank E. Scott and an anonymous reviewer, as well as the editors, for helping to improve the manuscript.

Submitted 17 March 2010. Accepted 8 May 2011. Associate Editor was Stephen G. Mech.

LITERATURE CITED

CANNON, P. J. 1967. Geologic map of Pleistocene deposits in southwestern Oklahoma. M.S. thesis, University of Oklahoma, Norman.

COLBERT, E. H. 1950. The fossil vertebrates. Pages 126-148 in The stratigraphy and archaeology of Ventana Cave (E. W. Haury, editor). University of Arizona Press, Tucson.

DALQUEST, W.W., AND G. E. SCHULTZ, JR. 1992. Ice age mammals of northwestern Texas. Midwestern State University Press, Wichita Falls, Texas.

DALQUEST, W.W., AND F. B. STANGL, JR. 1984. Late Pleistocene and early Recent mammals from Fowlkes Cave, southern Culberson County, Texas. Pages 432-455 in Contributions in Quaternary vertebrate paleontology: a volume in memorial to John E. Guilday (H. H. Genoways and M. R. Dawson, editors). Carnegie Museum of Natural History, Special Publication 8:1-538.

FAUNMAP Working Group. 1994. Faunmap: a database documenting late Quaternary distributions of mammal species in the United States. Illinois State Museum Scientific Papers. 25(1 and 2):1-685.

GIDLEY, J. W. 1920. Pleistocene peccaries from the Cumberland Cave deposit. Proceedings of the United States National Museum 57:651-678.

HARRIS, A. H. 1993. Quaternary vertebrates of New Mexico. New Mexico Museum of Natural History and Science Bulletin 2:179-197.

HEMMINGS, E. T. 2007. Buried animal kills and processing localities, areas 1-5. Pages 83-137 in Murray Springs: a Clovis site with multiple activity areas in the San Pedro Valley, Arizona (C. V. Haynes, Jr. and B. B. Huckell, editors). Anthropological Papers of the University of Arizona 71:1308.

HIBBARD, C. W., R. J. ZAKRZEWSKI, R.E. ESHELMAN, G. EDMUND,C. D. GRIGGS, AND C. GRIGGS. 1978. Mammals from the Kanopolis local fauna, Pleistocene (Yarmouth) of Ellsworth County, Kansas. Contributions from the Museum of Paleontology, University of Michigan 25:11-44.

JEFFERSON, G. T. 1991. A catalogue of late Quaternary vertebrates from California: part two, mammals. Natural History Museum of Los Angeles County, Technical Reports 7:1-129.

JOHNSTON, C. S. 1935. An extension in the range of fossil peccaries. American Midland Naturalist 16:117-119.

LUCAS, S. G., AND R. A. SMARTT. 1995. Late Pleistocene peccary from northwestern New Mexico. Southwestern Naturalist 40:293-296.

LUNDELIUS, E. L., JR. 1960. Mylohyus nasutus long-nosed peccary of the Texas Pleistocene. Bulletin of the Texas Memorial Museum 1:1-40.

MEAD, J. I., N. J. CZAPLEWSKI, AND L. D. AGENBROAD. 2005. Rancholabrean (late Pleistocene) mammals and localities of Arizona. Pages 139-180 in Vertebrate paleontology of Arizona (R. D. McCord, editor). Mesa Southwest Museum Bulletin 11:1-180.

MEAD, J. I., A. BAEZ,S.L. SWIFT ,M.C. CARPENTER, M. HOLLENSHEAD, N. J. CZAPLEWSKI, D. W.STEADMAN, J. BRIGHT, AND J. ARROYOCABRALES. 2006. Tropical marsh and savanna of the late Pleistocene in northeastern Sonora, Mexico. Southwestern Naturalist 51:226-239.

MORGAN, G. S., AND S. G. LUCAS. 2005. Pleistocene vertebrate faunas in New Mexico from alluvial, fluvial, and lacustrine deposits. New Mexico Museum of Natural History and Science Bulletin 28:185-248.

MURRAY, L. K., C. J. BELL, T. DOLAN, AND J. I. MEAD. 2005. A late Pleistocene fauna from the southern Colorado Plateau, Navajo County, Arizona. Southwestern Naturalist 50:363-374.

SEMKEN, H. A., AND C. D. GRIGGS. 1965. The long-nosed peccary, Mylohyus nasutus from McPherson County, Kansas. Michigan Academy of Science, Arts, and Letters 50:267-274.

SKINNER, M. F. 1942. The fauna of Papago Springs Cave, Arizona, and a study of Stockoceros; with three new antilocaprines from Nebraska and Arizona. Bulletin of the American Museum of Natural History 80:143-220.

SLAUGHTER, B. H. 1966. Platygonus compressus and associated fauna from the Laubach Cave of Texas. American Midland Naturalist 75:475-494.

SMITH, K. S., AND R. L. CIFELLI. 2000. A synopsis of the Pleistocene vertebrates of Oklahoma. Oklahoma Geological Survey Bulletin 147:1-36.

SPRINGER, K., E. SCOTT, J. C. SAGEBIEL, AND L. K. MURRAY. 2009. The Diamond Valley Lake local fauna: late Pleistocene vertebrates from inland southern California. Pages 217-235 in Papers on geology, vertebrate paleontology, and biostratigraphy in honor of Michael O. Woodburne (L. B. Albright, III, editor). Museum of Northern Arizona Bulletin 65:1-681.

STRAIN, W. S. 1937. The Pleistocene geology of part of the Washita River Valley, Grady County, Oklahoma. M.S. thesis, University of Oklahoma, Norman.

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Nicholas J. Czaplewski

Oklahoma Museum of Natural History, University of Oklahoma, Norman, OK 73072

Correspondent: nczaplewski@ou.edu
TABLE 1--Measurements (in mm) of teeth of two Pleistocene
peccaries, Platygonus compressus, from sites near Rush Springs,
Grady County (73929), and south of Yukon, Canadian County
(73930), Oklahoma.

                           OMNH catalog number

Tooth and measurements      73929      73930

p2
  Anteroposterior length     9.1         --
  Anterior width             6.1         --
  Posterior width            7.2         --
p3
  Anteroposterior length     10.3       9.9
  Anterior width             7.4        8.2
  Posterior width            8.6        9.5
p4
  Anteroposterior length     11.6       11.1
  Anterior width             9.2        9.2
  Posterior width            10.9       10.5
m2
  Anteroposterior length      --        16.7
  Anterior width              --        11.8
  Posterior width             --        13.0
m3
  Anteroposterior length      --        23.1
  Anterior width              --        13.0
  Posterior width             --        11.7

Table 2--Measurements (mm) of foot bones assumed to
represent a left manus from a Late Pleistocene peccary (OMNH
73931) from Quartz Mountain Canal, Jackson County, Oklahoma.

                                Proximal   Proximal   Distal    Distal
      Element         Length     width      depth      width     depth

Metacarpal III          --         --         --       16.3      18.7
Metacarpal IV           --         --         --       15.8      18.5
Phalanx 1 digit III    40.1       16.9       18.5      16.7      12.0
Phalanx 1 digit IV     40.5       15.1       19.1      15.1      12.4
Phalanx 2 digit III    29.5       15.9       17.7      13.8      15.3
Phalanx 2 digit IV     30.0       16.0       17.7      14.0      14.5
Ungual phalanx        35 (a)      13.5       19.1       --        --
  digit III

(a) Estimated measurement of broken bone.
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Author:Czaplewski, Nicholas J.
Publication:Southwestern Naturalist
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Geographic Code:1U7OK
Date:Mar 1, 2012
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