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Late Pleistocene mammalian fauna from Prescott Valley, West-Central Arizona.

In 1984, a partial skeleton of a mammoth was discovered by A. Jordan on Fain Ranch near the headwaters of the Agua Fria River in Prescott Valley (SE 1/4 SW 1/4 Section 31 T15N R1E, elevation 1,490 m), Yavapai County, Arizona (Fain Ranch Locality). That year, personnel of the Arizona Archaeological Society, Yavapai College, Sharlot Hall Museum, and Northern Arizona University excavated the skeletal remains of a mammoth (Mammuthus), the partial maxilla and mandible of a small horse (Equus), isolated teeth of a bison (Bison), and a partial mandible of a camelid (Camelops). The mammoth was at the bottom of a ca. 4-m-deep arroyo in the basal portion of a clay layer that was overlain by a layer of sand, indicating a fluvial deposit. Deposits composed of cross-bedded sands, and sand and pebble-filled channels, possibly playa lakes along Lynx Creek and the Agua Fria River, were reported near Fain Ranch Locality (Krieger, 1965). Between the clay and sand was a layer of charcoal ca. 1 m above the mammoth. A sample of this charcoal was [sup.14]C-dated to 11,560 [+ or -] 290 years before present, indicating a late Pleistocene (Rancholabrean land mammal age) occurrence. This date was calibrated to 13,410 + 285 years before present.

Pleistocene fossils have been reported from several localities in Prescott Valley, but none has described specimens. A specimen referred to Equus was near Lynx Creek, Yavapai County, ca. 7 km SW Fain Ranch Locality (Hay, 1927). Several teeth of Mammuthus were in terrace gravels above Granite Creek, Yavapai County, ca. 12 km W Fain Ranch Locality (Krieger, 1965). These specimens and other remains of mammoths, camels, bison, and horses from locations throughout Prescott Valley were mentioned by Morgan (1996). The M&M Mastodon Site, Yavapai County, ca. 40 km SW Fain Ranch Locality has yielded remains of a mastodon (Mammut americanum)andacamel (Morgan, 1996; Pasenko, 2011). Mammoths, mastodons, camels, and bison have been reported from a Pleistocene lake bed 3 km N M&M Mastodon Site (Hook, 1956; Morgan, 1996). The purpose of our research was to describe fossils of large mammals excavated at Fain Ranch Locality.

MATERIALS AND METHODS--Fossils described here are housed at the Sharlot Hall Museum (SHM) in Prescott, Arizona. We use measuring protocols of von den Driesch (1976), Dutrow (1980), Agenbroad (1994), and Lister (1996). We report the average of three measurements taken using calipers. We compared postcranial remains of the mammoth from Fain Ranch Locality to those of other mammoths from Yavapai County (Museum of Northern Arizona MNA P1.1271), a sample from the Hot Springs Mammoth Site, Fall River County, South Dakota, those from Dent, Weld County, Colorado, and to mammoths (Mammuthus) and mastodons from the literature. We follow Skinner (1972) and Downs and Miller (1994) in describing teeth of horses, Frison et al. (1976) in describing teeth of bison, and Loring and Wood (1969) in describing deciduous teeth of camelids. Remains of the horse, camel, and bison were compared to specimens in published literature and to specimens in museum collections for taxonomic purposes and for determining age of individuals.

RESULTS--Mammuthus cf. columbi--Remains of the mammoth consisted of a right femur (SHM 85.126.1I), a left radius (SHM 17716.G), a right inominate (SHM 177160), a left tusk (85.126.1J), the axis (85.126.1b), two thoracic vertebrae (SHM 85.126.1c and SHM 17716c), three partial cervical vertebrae (SHM 17716d, 85.126.1d, and one uncatalogued), a partial sacrum (SHM 17716y), and six ribs (SHM17716u, SHM 17716t, SHM 17716p, SHM 177168, and two uncatalogued). Two ribs were represented only by fragments and were not measured. Only one individual was represented.

The left tusk (Fig. 1e) was fairly well preserved except for the proximal portion, which was damaged slightly around the pulp cavity. Curvilinear length of the tusk is 1,301 mm and greatest diameter is 160 mm. The tusk displayed a small wear pattern on the lateral side near the tip. The axis (Fig. 1d) was fairly complete, missing only the transverse processes. Maximum height of axis is 238 mm and width of cranial articular process is 189 mm. The two thoracic vertebrae were missing the body and transverse processes due to nonfusion of these elements, but the spinous process, superior and inferior articular facets, and pedicle were present. Maximum heights of preserved portions of thoracic vertebrae are 257 and 443 mm. The larger (Fig. 1a) is probably a second or third thoracic vertebrae, and the smaller may be a sixth or seventh. Length of the larger thoracic vertebrae is greater than two described by Dutrow (1980) from Hot Springs Mammoth Site, but Dutrow also provided lengths of other thoracic vertebrae as 550-692 mm. Three other vertebrae consisted of the centrum only and have transverse widths of 116, 120, and 152 mm. Curvilinear lengths and proximal widths, respectively, of the three best-preserved ribs are 633 and 43, 763 and 50, and 840 and 52 mm.

The partial left inominate (Fig. 1c) included the acetabular fossa, pubis, ischium, obturator foramen, and most of the ilium. The large, fan-shaped portion of the ilium was incomplete, but most of the iliac crest was preserved. A prominent ischial spine was present. The width, representing one-half of the pelvis, is 711 mm. Maximum width of acetabulum is 184 mm and minimum width of shaft of the ilium is 193 mm. Estimated oblique height of pelvic apertures were 270-280 mm. The acetabulum has a fairly deep cotyloid notch that opens into the acetabular fossa. Width of the wing of the ilium from tuber coxae to pelvic aperture is 485 mm. Widths of wings of the ilium for mammoths at Hot Springs Mammoth Site were 520-640 mm and minimum widths of shaft of the ilium were 185-260 mm (Agenbroad, 1994). The sacrum consisted of two preserved sacral vertebrae (fused) containing portions of the spinous tubercles and articular processes for each. The femur (Fig. 1b) consisted of the diaphysis only, representing a total length >892 mm. Diameter at mid-shaft of femur is 124 mm and least circumference is 351 mm. The radius was missing the distal epiphysis and greatest length was > 598 mm.

Several characteristics of postcranial elements are different between mammoths and mastodons (Olsen, 1979). Some of these characteristics are associated with epiphyses, but the proboscidean from Fain Ranch Locality had few of these. However, several notable characteristics of vertebrae in Mammut and Mammuthus are distinguishable. Thoracic vertebrae of Mammuthus are wider and more robust near the centrum and lamina that form the vertebral foramen (neural arch) than they are in Mammut. Accordingly, the vertebral foramen is wider in Mammuthus. The axis in Mammut generally has a greater height, especially the dorsal one-half, than Mammuthus (Olsen, 1979; Miller, 1987). Also, the dorsal one-half of the axis in Mammut is narrower than that of Mammuthus. Similarly, Olsen (1979) showed a proportionately narrower diaphysis of the femur in Mammuthus than Mammut. Circumference of mid-shaft of femur for the proboscidean from Fain Ranch Locality is smaller when compared to those of Mammut (Table 1). Ratios between least circumference and length of femora for a sample of Mammut americanum from Boney Spring, Benton County, Missouri, were 0.40 and 0.44. This ratio for the proboscidean from Fain Ranch Locality is 0.39, but would have been smaller because epiphyses were missing. This corresponds more to the suggestion by Olsen (1979) that femora of mammoths have a narrower diaphysis. From these comparisons, the proboscidean from Fain Ranch Locality resembles a mammoth more than a mastodon.

Determination of species for mammoths relies mostly on morphometrics of crania and teeth. Because the mammoth from Fain Ranch Locality is missing these elements, an assignment to species is not possible. However, Mammuthus primigenius (woolly mammoth) is not known from Arizona (Agenbroad, 1984; Mead et al., 2005) and the earlier M. meridionalis (southern mammoth) has not been found in late Pleistocene localities. Distribution of M. columbi (Columbian mammoth) in North America is throughout western Canada and most of the United States and Mexico, with numerous fossils from west of the Great Lakes, in the Great Plains, and in the Southwest (Agenbroad, 1984). In Arizona, remains of M. columbi have been reported from >70 localities, including one previously recorded from eastern Yavapai County (Saunders, 1970; Mead et al., 2005). Several of these localities have cultural affiliations. Given the late Pleistocene age and these distributional patterns, it is probable that the mammoth from Fain Ranch Locality represents M. columbi, but based on available material, it can only be referred to M. cf. columbi. The late Pleistocene age for the mammoth from Fain Ranch Locality is near the period established for Clovis-age humans, but no artifact was recovered at Fain Ranch Locality. It is believed that one-third of mammoth sites with [C.sup.14]-dates [less than or equal to] 11,500 years before present have cultural affiliations (Agenbroad, 2005), which means Fain Ranch Locality is an example of a Clovis-age site without a known cultural association.

[FIGURE 1 OMITTED]

Epiphyses of the mammoth from Fain Ranch Locality were absent, except for the proximal epiphysis of the radius, signifying that fusion of the diaphyses and epiphyses had not occurred at time of death. According to Lister (1994), the proximal epiphysis of the radius is fused to the diaphysis at the age of 33 African elephant years for a male M. primigenius. Likewise, for a male African elephant (Loxodonta africana), the proximal epiphysis of the radius fuses at 19 African elephant years for a female and >32 for a male (G. Haynes, 1991). The corresponding distal epiphysis is fused at >24 African elephant years for a female and <32 for a male African elephant. According to G. Haynes (1991), the femur of male African elephants can be unfused as late as 22-24 African elephant years for the proximal epiphysis and 28-30 years for the distal epiphysis. For female African elephants, proximal and distal epiphyses of the femur can begin to fuse as early as 12-14 African elephant years. Unfused proximal and distal surfaces of the preserved diaphyses of the mammoth from Fain Ranch Locality correspond to a 1 or 2 in the G. Haynes (1991) grading system of 0-5 for diaphyseal-epiphyseal fusion, based on Loxodonta. A 1-2 indicates increased complexity of conjoining surfaces of diaphysis and epiphysis, i.e., more roughening and detailed sculpting of surfaces, such that they fit together more precisely, but the epiphysis is still unfused (G. Haynes, 1991). Based on this sequence of fusion in extant and extinct proboscideans, age of the mammoth from Fain Ranch Locality at time of death was ca. 12-20 African elephant years if it was a female and 22-32 years if it was a male.

Measurements of postcranial elements were compared to the population of male M. columbi from Hot Springs Mammoth Site, South Dakota (Dutrow, 1980; Agenbroad, 1994), mammoths from Dent, Colorado (Saunders, 1999), a mammoth (MNA P1.1271) from eastern Yavapai County, Arizona (Saunders, 1970), and Mammut americanum from Missouri, Utah, and Indiana (Saunders, 1977; Graham et al., 1983; Miller, 1987; Woodman and Branstrator, 2008; Table 1). In these comparisons, size of skeletal elements of the mammoth from Fain Ranch Locality is at the low end of the range of measurements because of unfused bones. The axis of the mammoth from Fain Ranch Locality also is slightly smaller than that of a previously reported axis (MNA P1.1271) from eastern Yavapai County (Saunders, 1970). The small size of the mammoth from Fain Ranch Locality suggests it was a female or small male, but, based on epiphysis-diaphysis fusion in other mammoths (Lister, 1994), the proximal epiphysis of the radius is early to middle in this sequence, indicating the mammoth from Fain Ranch Locality had considerable ontogenetic growth remaining. In this regards, it could be a male. In addition, the estimated ratio of height of pelvic aperture to shaft of ilium for the mammoth from Fain Ranch Locality is 1.40-1.45, which Lister and Agenbroad (1994) believed represented a male. Females have wider pelvic apertures, because of birthing, and generally have ratios of 2.4-3.0.

Fairly good preservation and condition of much of the remains indicates rapid burial of bones with little weathering. None of the remains displayed gnawing marks or spiral fractures. Absence of cranial material suggests either the cranium was not preserved, or, because of taphonomic differences, was not deposited with postcranial material. The skull and mandible of proboscideans often lag behind other skeletal elements in a depositional environment, mainly because of flotation of the skull due to diploe (Voorhies, 1969; Agenbroad and Mead, 1994). However, tusks can be one of the last elements deposited (Agenbroad and Mead, 1994), and because the mammoth from Fain Ranch Locality had one preserved tusk, the cranium may not have been preserved. Remains were disarticulated and scattered. The femur was ca. 2 m from the inominate, which was lying next to the tusk. Several ribs were near a thoracic vertebrae. The axis and several partial vertebrae were near the inominate and tusk.

Equus cf. conversidens--Remains of the horse consisted of left and right partial dentaries (SHM 96.127.16 and SHM 96.127.18), and a fragmentary maxilla with three associated upper teeth (SHM 96.127.1a). The left dentary included the posterior one-half of the horizontal ramus, coronoid and condylar processes, and p4-m3. The right dentary (Figs. 1f and 1g) included most of the horizontal ramus, coronoid and condylar processes, and p2-m3. No incisor was preserved. All lower teeth exhibited wear, but judging from the small amount of wear and modest amount of eruption of m3s, the individual was probably a young adult. For modern horses, the m3 begins to erupt at 3.5-4.5 years of age (Silver, 1970). Lower molars were fairly simple, displaying little accessory folding of enamel. The m1-m3 displayed ectoflexids that were short with no penetration of the isthmus. Metaconid-metastylid valleys (linguaflexids) of p3-m3 were generally broadly U-shaped, but several approach a more V-shape. Occurrence of both U-shaped and V-shaped linguaflexids in the same jaw of small horses from the Pleistocene have been reported previously (Scott, 1996). Lower molars had no pli-caballinid.

Characteristics of teeth of the horse from Fain Ranch Locality were similar to those referred to Equus conversidens (Mexican ass) by Dalquest and Hughes (1965), Mooser and Dalquest (1975a), and Dalquest (1979). Length of the p2-m3 series of the right mandible is 151.7 mm. This is similar to those for E. conversidens described by Mooser and Dalquest (1975a), which were 147.6-150.0 mm, and those in Dalquest and Hughes (1965), which were 145.0 and 154.0 mm. In contrast, the toothrow series for the similar taxon E. niobrarensis can be >173 mm (Skwara and Walker, 1989). Other measurements of lower molars and mandibles of the horse from Fain Ranch Locality and comparative specimens are listed in Table 2. Upper molars were incomplete, missing portions of occlusal surfaces; therefore, accurate measurements of lengths and widths were impossible to obtain. One of the best preserved upper teeth was a P4; its width is 21.1 mm.

The holotype of E. conversidens has a P4 with a width of 22.3 mm (Hibbard, 1955). It is probable that the upper molars represent the same species as the mandible, but this is not certain. No other fossil of horses was recovered. The holotype of E. conversidens was described briefly in Owen (1869) and was based on maxillae, palate, and upper cheek teeth from the Valley of Mexico. The holotype was redescribed, as well as additional cheek teeth, by Hibbard (1955); he concluded that E. conversidens, E. francisci, and E. tau probably were one species. Lack of an associated mandible and lower teeth for the holotype raises questions as to its validity as a separate species. Upper cheek teeth of Pleistocene horses have more variability in morphology than the more diagnostic lower teeth. Equus achates, E. barcenaei, E. francisi, E. littoralis, and E. tau were listed as synonyms of E. conversidens by Dalquest and Hughes (1965). However, type specimens of E. conversidens, and referred material from Aguascalientes, Mexico, were considered to be two species (E. conversidens and E. tau) by Mooser and Dalquest (1975a). This conclusion was based mostly on measurements of teeth and the tooth row, which they noted were similar in E. conversidens and E. tau. Similarities in morphology of teeth between E. conversidens and closely related E. niobrarensis and E. scotti were discussed by Skwara and Walker (1989), who concluded that, although there was overlap in size and morphology, E. conversidens generally was smaller.

Variability in morphology of teeth in small horses from the late Pleistocene, together with different diagnoses for E. conversidens and lack of a mandible and lower teeth for the holotype, have left uncertainties as to validity of the specimen from the Valley of Mexico as a holotype (Scott, 1996, 2004). This and the lack of phalanges hinder assignment of species to the one horse from Fain Ranch Locality. Based on these comparisons and the current status of E. conversidens, specimens from Fain Ranch Locality are referred tentatively to this species. Equus conversidens is to be a small, stout-limbed equid with lower cheek teeth having U-shaped linguflexids, a full isthmus, and short-moderate ectoflexids (Hibbard, 1955; Dalquest and Hughes, 1965; Mooser and Dalquest, 1975a; Dalquest, 1979; Harris and Porter, 1980; Kurten and Anderson, 1980; Scott, 1996).

This is the first report of E. conversidens in Yavapai County, Arizona, but it has been reported from Canada southward into El Salvador (Churcher, 1975; FAUNMAP Working Group, 1994; Cisneros, 2005). Previously, E. conversidens was reported from only two localities in Arizona; Papago Springs (Skinner, 1942) and Stanton's Cave in the Grand Canyon (Harington, 1984). There are other reports of Pleistocene horses or Equus in Prescott Valley, but these were not described (Hay, 1927; Morgan, 1996).

Bison--Remains consisted of a left m2 (SHM 96.127.2f), left dp3 (SHM 96.127.2e), and a right dp4. The dp3 (Fig. 2d) had two widely branched roots, which would accommodate the incoming p3. A small portion of the distal end of dp3 was missing, including a portion of that root. The labial side of the crown was mostly flat except for a slight fold on the hypoconid. There was a deep lingual fold that penetrated one-half of the occlusal surface between the entoconid and metaconid, and a slight fold on the entoconid. The occlusal surface of dp3 had extensive wear. A small oval fossette was on the posterior lophid. The dp3 had a slightly pointed appearance from the sides with the apex being immediately mesial to the center of the tooth. Length and greatest width of dp3 is 19.5 and 8.83 mm, respectively; however, length was probably slightly longer. This tooth resembles a dp3 in morphology, size, and wear, for immature Bison depicted in Skinner and Kaisen (1947: plate 12) and Chandler (1916:figure 12).

The dp4 (Fig. 2c) was composed of two specimens (SHM 96.127.2c and SHM 96.127.2d) that fit together. The dp4 had three preserved roots. The anterior lophid had a semi-U-shaped prefossette with the distal portion oriented slightly more lingually then the mesial one-half. The hypoconulid had a fossette as well, and appeared as a deep pit. The postfossette on the second lophid was not preserved due to breakage. The protoconid was U-shaped. Exostylids were present between the protoconid and hypoconid, and between the hypoconid and hypoconulid. These exostylids had occlusal surfaces about triangular in shape and similar to those depicted for an immature specimen of Bison antiquus in Chandler (1916). Length and greatest width of the dp4 are 36.07 and 12.1 mm, respectively. This tooth also compares closely to a dp4 illustrated in Skinner and Kaisen (1947:plate 12) for an immature Bison. Measurements of deciduous premolars of Bison are provided in Table 3. There is no species-level taxonomic indicator for deciduous premolars of Bison.

The unworn m2 (Fig. 2e) was well preserved missing only portions of the roots. An exostylid was present between the protoconid and hypoconid. Greatest length and width of m2 are 40.8 and 14.5 mm, respectively. Comparative measurements of m2s for B. antiquus from Chandler (1916) are 39 by 19, 36 by 21, 37 by 20, 39 by 19, 37 by 16, and 35 by 15 mm. Preserved height of left m2 (SHM 96.127.2f) is 75.2 mm. Morphology of m2 and deciduous premolars is consistent with those of late Pleistocene Bison, but because dental measurements and morphology are variable among species, and specific determination of Bison is based mostly on postcranial elements, horncores, and cranial material, teeth of bison from Fain Ranch Locality can be assigned only to genus.

Presence of isolated deciduous and permanent teeth enables one to hypothesize about age of the bison. If teeth were from one individual, then they represent death of an immature bison. Previous studies of replacement and succession of teeth in Bison has revealed that dp2, dp3, and dp4 can be shed at different times, and that m1 and m2 come into use by the time dp3 and dp4 are being shed and replaced by premolars. Bison 1.5 years of age from Glenrock Site, Converse County, Wyoming, had dp2s, dp3s, and dp4s that were well worn with roots showing, and m2 usually was unworn or had slight wear (Frison and Reher, 1970). Bison 1.7 years of age from Hawken Site, Crook County, Wyoming, had deciduous premolars in place at this age with m1 fully erupted and in wear and m2 erupted, exhibiting variation from light to moderate wear on facets I and II, and light wear on facets III and IV (Frison, et al., 1976). By 2.7 years of age, Bison at Hawken Site had lost dp2, and dp3 was being replaced by p2 and p3, but dp4 was in place. At this age, m1 and m2 were in full wear. Similar results were reported for Bison from Casper Site, Natrona County, Wyoming, but it was noted that eruption of premolars could vary by <2 years (Reher, 1974). In some instances, dp2, dp3, and dp4 could be in place at the same time. According to Reher (1974), by 2.6 years of age, dp2 and dp3 were at varying stages of being shed and m1 and m2 were in regular wear. Following these comparisons, and given the wear on the three isolated teeth of bison from Fain Ranch Locality, they could represent one individual. If so, the individual would have been 1.5-2.6 years of age at time of death.

Bison has been reported at other Pleistocene localities in Arizona, although not as widespread as in the Great Plains. Occurrences of Bison in Arizona include Pima County (Hay, 1927), Ventana Cave, Pima County (Colbert, 1950), Stanton's Cave, Coconino County (Harington, 1984), Lehner Mammoth Site, Cochise County (Haury et al., 1986), and Murray Springs, Cochise County (Agenbroad and Haynes, 1975; FAUNMAP Working Group, 1994; Wilson, et al., 2007). Probable ranges of B. antiquus and B. latifrons include Arizona (Skinner and Kaisen, 1947; McDonald, 1981). Presence of one juvenile would indicate a large presence of Bison in Arizona. Social structure of Bison consists of herds of varying sizes with different percentages representing certain age groups. Bison at Yellowstone National Park have herds that generally are <100 (Meagher, 1973), which contrasts to vast herds known historically from the Great Plains. In Yellowstone National Park, calves that are 2.5 years old constituted ca. 6-11% of a population of bison (Meagher, 1973).

Camelops--Remains of camelids (SHM 85.1262) consisted of a partial mandible with left and right dp3-m1, right di1-di3, right dc1, and portions of left and right vertical rami. The mandible (Figs. 2a and 2b) was in poor condition, with portions o horizontal and ascending rami missing. A diastemal crest was visible on the right ramus, although it was not strongly developed. Ascending ramus was crushed partially and unattached to the horizontal ramus. Le t horizontal ramus had been re-attached to the mandible in a position anterior to its natural location. Le t condyle, coronoid process, and angular process were preserved. Coronoid process was airly complete, missing only small portions o dorsal edges. The coronoid process curved posteriorly and slightly overhung the condyle. Anterior border o ascending ramus gently curved to the area where it joined the horizontal ramus. Symphysis was incomplete, missing portions o its anterior and posterior edges. Preserved length o symphysis is 92 mm, but this length probably reflects young age o the individual and its incompleteness. It is probable that length would have been >110 mm when complete. Preserved greatest width of symphysis is 48 mm. Measurements of SHM 85.1262 are in Table 3.

[FIGURE 2 OMITTED]

No premolar was preserved, but one small, deciduous canine was present on the right ramus. This small canine was broken off and was on its side in a crack behind the right i3. Original position of canine was not apparent. The di3 overlaped di2, and di2 overlaped di1. Deciduous incisors were procumbent, but di1 had a slight upward curve. Dorsal-ventral diameter of di1 is 6.2 mm, of di2 is 5.94 mm, and of di3 is 4.32 mm. Sizes of incisors indicate that they were deciduous.

Left cheek teeth were better preserved and are described here. The dp3 was about triangular in shape, having a small anterior part and a larger posterior part. Lingual and buccal sides of anterior lobe of dp3 were mostly straight. Posterior lobe was molariform in shape and had an oval-shaped fossettid. The dp3 had moderate wear and a length of 22.2 mm. The dp4 was molariform and consisted of three lobes. Each lobe increased in size posteriorly and had a crescent-shaped fossetid. The fossetid on the anterior-most lobe had an anterior restriction. Inflection between the second and third lobes was greater than that between the first and second lobes. Wear on dp4 diminished posteriorly, with the third lobe showing little wear. Length of dp4 is 58.4 mm and greatest width is 18.5 mm. The m1 had two lobes, each with a crescent-shaped fossetid. It appeared that m1 had not erupted at time of death, showed no wear, and was lower than dp4. Greatest anterior-posterior length of unworn m1 varies from 47.3 mm near apex of tooth to 50.7 mm, which was taken slightly below the first measurement. Greatest width of m1 is 17.8 mm.

The spatulate-shaped incisors, a small canine, a diastemal crest, and semi-V-shaped fossetids point toward the specimen being in the family Camelidae. The subfamily Camelinae includes the tribes Lamini (extant and extinct lamines Alforjas, Camelops, Hemiauchenia, Llama, and Palaeolama) and Camelini (extant camels, Camelus, and the earlier giant camels Megatylopus and Titantylopus; Harrison, 1979). Both Lamini and Camelini share several characteristics, but Lamini is distinguished from Camelini by configuration of the anterior end of the nasals and presence of an anteroexternal style (llama buttress) on lower molars (Harrison, 1979). This style, which is strongly developed in extant and extinct llamas, is weak to moderately developed in Camelops. Alforjas and Camelops are distinguished from other Lamini by increased hypsodonty of cheek teeth (Harrison, 1979). Alforjas is known only from Miocene-Pliocene deposits.

Both dp4 and m1 of the camelid from Fain Ranch Locality are hypsodont. Lengths of dp4 and m1 indicate it was a large camelid, much larger than Hemiauchenia. For the late Pleistocene, the largest member of Lamini was Camelops. However, the m1 described here from Fain Ranch Locality is longer than those typically reported for Camelops. This may be a result of its unworn condition, or just an atypical specimen. For teeth of camels, there is a correlation between length of molar and amount of wear. Teeth with less wear are longer than the same tooth displaying greater wear (Dalquest, 1992). The closest length of an m1 to the camelid from Fain Ranch Locality is from Webb (1965), who described one for Camelops from Rancho la Brea, Los Angeles County, California, that had a length of 43.7 mm. There are few comparisons of deciduous teeth of Camelops in the literature. The m1 of the camelid from Fain Ranch Locality does not have the llama buttress typical of other lamine camelids, but, as noted, this is weak to moderately developed in Camelops. Presence or absence of the llama buttress in deciduous premolars has not been discussed previously. Size of the camelid from Fain Ranch Locality, absence of llama buttresses, and age of deposits warrant referring it to Camelops.

The genus Camelops has a complex history having numerous species assigned previously, many of which have now become junior synonyms. Earlier studies placed different forms of Camelops into two groups; a larger group that included C. hesternus, C. huerfanensis, and C. kansanus, and a smaller group that included C. minidokae and C. sulcatus (Savage, 1951). However, size of C. sulcatus was similar to, if not larger than C. hesternus (Webb, 1965). The monograph on Camelops from Rancho La Brea (Webb, 1965) has served as the basis for revisions (e.g., Harrison, 1979; Dalquest, 1992). However, deciduous teeth ofCamelops have not been included in most previous studies, typically are not described, or both.

Available fossils of camelids from Fain Ranch Locality do not allow for assignment to species. Cheek teeth of camelids are not good indicators of species. The skull or metapodials of camelids are more reliable for determining species. However, these elements were not recovered from Fain Ranch Locality. Of the two size groupings mentioned above, the camelid from Fain Ranch Locality is closer to the larger size of the C. hesternus-C. huerfanensis-C. kansanus group. However, m1 is longer and narrower than those for Camelops, and does not have a llama buttress. In addition, dp4 is longer than that of a juvenile C. hesternus from Selby-Dutton, Yuma County, Colorado (Graham, 1981). It is indeterminate at this time if these attributes indicate a new species or a larger form of C. hesternus, C. huerfanensis, or C. kansanus. More material or comparisons are needed. Specific distinctions between C. hesternus and C. huerfanensis are based on morphology of cranium and ventral border of ramus, which were not recovered. It is believed that C. huerfanensis was slightly larger than C. hesternus. An even larger Camelops may have existed during the early Pleistocene in southern California (Webb et al., 2006). At present, the camelid from Fain Ranch Locality can only be referred to the genus Camelops.

Presence of deciduous teeth in the mandible of the camelid from Fain Ranch Locality, little wear on dp4, and the unerupted m1 indicates it was a juvenile. For extant camels, dp3 and dp4 erupt at 4-6 months of age, deciduous canines erupt at 10-12 months, permanent canines do not erupt until 6.5 years, and m1 erupts at ca. 1 year of age. The first permanent incisor erupts at ca. 4 years of age (Silver, 1969). Based on these comparisons, the camelid from Fain Ranch Locality would have been 10-12 months of age at time of death.

Camelops has been found throughout the western United States, Mexico, and Canada. The genus has been reported from several locations in Arizona including Murray Springs and Lehner Ranch in Cochise County, Vulture Cave in Mojave County, Papago Springs Cave in Santa Cruz County, the Shonto fauna in Navajo County, and city of Mesa in Maricopa County (Mead and Phillips, 1981; McDonald et al., 2004; Mead, et al., 2005; Hemmings, 2007; R. D. McCord, pers. comm.).

DISCUSSION--Today, the Prescott Valley area is a mix of chaparral communities and semi-desert grasslands. Antilocapra americana (pronghorn) and Callipepla gambelii (Gambel's quail) are common while Pecari tajacu (collared peccary) and Odocoileus (deer) are less common. Large mammals at Fain Ranch Locality are typical of a late Pleistocene (Rancholabrean) fauna. Four taxa were recovered, but none was adult. The camel and bison were juveniles, and the mammoth and horse were subadults. This could indicate that environmental factors contributed to their death. Presence of clay and sand at the site suggests water was present in some quantity. However, the late Pleistocene age for Fain Ranch Locality is close to the Clovis-age drought, discussed by C. V. Haynes (1991), which occurred ca. 11,300-10,900 years before present. Presence of four taxa that were grazers with hypsodont teeth, as well as herd-structured animals indicates grasses were present in some quantity during the late Pleistocene in central Arizona. Hypsodont teeth of extant and extinct horses are an adaptation to eating coarse, gritty plants, such as grasses (Scott, 2006). Modern Bison from Yellowstone National Park have a diet of predominantly grasses or grass-like plants (Meagher, 1973). Mammoths are believed to have been mostly grazers with diets mainly of C4 vegetation. Dung boluses from Bechan Cave, Kane County, Utah, indicated those mammoths had a diet consisting mostly of grasses and sedges (Mead, et al., 1986). Camelops generally is regarded as occupying scrub grasslands and, although it was adapted for grazing, its long neck and limbs may have provided opportunistic browsing (Hibbard and Taylor, 1960; Webb, 1965).

We thank S. Lynch and L. Lorentzen for providing access to fossils, R. Fain for granting access to Fain Ranch, J. Saunders for discussions on mammoths and camels, G.Jefferson and R. White for conversations on teeth of bovids, E. Scott for discussions about teeth of horses, G. Morgan for discussions on teeth of camels, and the Museum of Northern Arizona for granting access to collections. Appreciation goes to S. Swift for assistance with photos and M. Anderson for help with translation of the abstract. We also appreciate S. Lucas and anonymous reviewers for comments on an early draft of the manuscript.

Submitted 18 August 2009. Accepted 12 June 2011. Associate Editor was Stephen G. Mech.

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Michael R. Pasenko * and Larry D. Agenbroad

Environmental Planning Group, 4141 North 32nd Street, Suite 102, Phoenix, AZ 85018 (MRP) Mammoth Site of Hot Springs, P.O. Box 692, Hot Springs, SD 57747 (LDA)

* Correspondent: mpasenko@epgaz. com
TABLE 1--Measurements (mm) of a subadult mammoth from Fain Ranch
Locality, Yavapai County, Arizona, and those of Columbian mammoths
(Mammuthus columbi) and mastadons (Mammut americanum) from elsewhere
in the United States.

                                           M. cf.        M. columbi,
                                       columbi, Fain         Hot
Measurements                           Ranch Locality    Springs (a)

Curvilinear length of tusk                 1,301          970-2,750
Straight-line length of tusk               1,059          950-1,770
Greatest diameter of tusk                   160            104-227
Greatest length of femur                    >892         1,147-1,317
Posterior width of femur                    >306           295-373
Distal width of femur                       >236           159-315
Diameter of femur at mid-shaft              124            137-199
Least circumference of femur                351              --
Greatest length of radius                   >598           410-690
Proximal width of radius                    110            111-162
Distal width of radius                      >158           168-220
Greatest width of pelvis               ca. 1,422 (e)     1,460-1,820
Width of illium                             485            520-640
Minimum width of shaft of illium            193            185-255
Maximum diameter of acetabulum              172            155-215
Length of obturator foramen                 176            168-208
Height of obturator foramen                 102            79-106
Maximum height of axis                      238            202-328
Width of cranial articular process          189            152-234
Height of cranial articular process         103            126-137
Superior width of axis                      100            109-136
Width across posterior process              125            144-136
  of axis
Superior height of axis                      75            109-136
Width of neural canal of axis                61             60-96
Height of posterior centrum of axis         123            119-164
Height of neural canal of axis               69             53-88
Maximum height of thoracic vertebrae      257, 443        234, 341
Maximum width of thoracic vertebrae       176, 249        298, 322
Maximum width of neural canal of           54, 70          57, 74
  thoracic vertebrae
Maximum height of neural canal of          45, 55          45, 50
  thoracic vertebrae
Width of centrum of thoracic                133           133, 149
  vertebrae
Total length of sacrum                      >206           144-380
Anterior width of sacrum                    167            151-168
Posterior height of sacrum                   47             52-56
Anterior height of sacrum                    85            105-142

                                                               M.
                                         M. columbi,       americanum,
Measurements                               Dent (b)       Overmeyer (c)

Curvilinear length of tusk                    --               --
Straight-line length of tusk                  --               --
Greatest diameter of tusk                     --               --
Greatest length of femur                 1,061, 1,125          --
Posterior width of femur                     328               --
Distal width of femur                      216-268             --
Diameter of femur at mid-shaft              84-158             --
Least circumference of femur                  --               --
Greatest length of radius                     --               --
Proximal width of radius               ca. 115, ca. 131        --
Distal width of radius                        --               --
Greatest width of pelvis                      --               --
Width of illium                               --               --
Minimum width of shaft of illium              --               189
Maximum diameter of acetabulum                --               163
Length of obturator foramen                   --               --
Height of obturator foramen                   --               --
Maximum height of axis                     249, 255            --
Width of cranial articular process         175, 226            --
Height of cranial articular process           --               --
Superior width of axis                        --               --
Width across posterior process                --               --
  of axis
Superior height of axis                       --               --
Width of neural canal of axis                 --               --
Height of posterior centrum of axis           --               --
Height of neural canal of axis                --               --
Maximum height of thoracic vertebrae          --               --
Maximum width of thoracic vertebrae          281               --
Maximum width of neural canal of              --               --
  thoracic vertebrae
Maximum height of neural canal of             --               --
  thoracic vertebrae
Width of centrum of thoracic                  --               --
  vertebrae
Total length of sacrum                       280               --
Anterior width of sacrum                      --               --
Posterior height of sacrum                    --               --
Anterior height of sacrum                     --               --

                                       M. americanum,
                                        Christensen     M. americanum,
Measurements                              Bog (d)       Boney Springse

Curvilinear length of tusk                   --               --
Straight-line length of tusk                 --               --
Greatest diameter of tusk                    --               --
Greatest length of femur                     --            940-1,216
Posterior width of femur                     --               --
Distal width of femur                        --             231-265
Diameter of femur at mid-shaft               --               --
Least circumference of femur                 --             379-507
Greatest length of radius                    --               --
Proximal width of radius                    117               --
Distal width of radius                       --               --
Greatest width of pelvis                     --               --
Width of illium                              --               --
Minimum width of shaft of illium             --               --
Maximum diameter of acetabulum               --               --
Length of obturator foramen                  --               --
Height of obturator foramen                  --               --
Maximum height of axis                    242, 257            --
Width of cranial articular process           --               --
Height of cranial articular process          --               --
Superior width of axis                       --               --
Width across posterior process               --               --
  of axis
Superior height of axis                      --               --
Width of neural canal of axis                --               --
Height of posterior centrum of axis       111, 122            --
Height of neural canal of axis               --               --
Maximum height of thoracic vertebrae         --               --
Maximum width of thoracic vertebrae          --               --
Maximum width of neural canal of             --               --
  thoracic vertebrae
Maximum height of neural canal of            --               --
  thoracic vertebrae
Width of centrum of thoracic                 --               --
  vertebrae
Total length of sacrum                       --               --
Anterior width of sacrum                     --               --
Posterior height of sacrum                   --               --
Anterior height of sacrum                    --               --

(a) From Hot Springs, Fall River County, South Dakota (Dutrow. 1980;
Agenbroad, 1994).

(b) From Dent, Weld County, Colorado (Saunders, 1999).

(c) From Overmeyer, Fulton County, Indiana (Woodman and Branstrator,
2008).

(d) From Christensen Bog, Hancock County, Indiana (Graham et al.,
1983).

(e) From Boney Spring, Benton County, Missouri (Saunders, 1977).

TABLE 2--Measurements (mm) of mandible and teeth of Equus cf.
conversidens (Mexican ass) from Fain Ranch Locality, Yavapai County,
Arizona, and other specimens of E. conversidens.

                                   Fain Ranch     Dalquest et al.
      Equus conversidens            locality          (1979)

Aboral height of vertical ramus        182              --
Oral height of vertical ramus          217              --
Middle height of vertical ramus        189              --
Height of mandible behind m3           106              --
Length of gonion caudale-aboral
border of alveolus of m3               112              --
Length of m3                      25.60, 26.43      29.8, 25.4
Width of m3                       11.90, 12.06      13.9, 11.9
Length of m2                      22.80, 22.56      22.4, 21.4
Width of m2                       12.90, 13.95      16.0, 14.4
Length of m1                      20.60, 21.67      22.2, 23.0
Width of m1                       13.40, 14.06      15.8, 14.4
Length of p4                      24.30, 24.70,     23.2, 25.7
Width of p4                       14.60, 14.43      17.8, 15.5
Length of p3                          25.84             --
Width of p3                           15.37             --
Length of p2                          29.05             --
Width of p2                           14.71             --

                                  Harris and Porter
      Equus conversidens               (1980)

Aboral height of vertical ramus          --
Oral height of vertical ramus            --
Middle height of vertical ramus          --
Height of mandible behind m3             --
Length of gonion caudale-aboral
border of alveolus of m3                 --
Length of m3                         24.0, 28.1
Width of m3                          11.5, 12.0
Length of m2                         24.1, 22.5
Width of m2                          13.4, 13.0
Length of m1                         23.6, 22.4
Width of m1                          13.8, 14.8
Length of p4                         26.4, 24.5
Width of p4                          16.3, 15.4
Length of p3                         26.6, 25.3
Width of p3                          16.3, 14.3
Length of p2                         30.8, 26.1
Width of p2                          12.1, 10.4

TABLE 3--Measurements (mm) of lower teeth and mandible of Camelops
from Fain Ranch Locality, Yavapai County, Arizona, and other specimens
of Camelops.

                         Fain Ranch
                          locality        Camelops
     Measurement        (SHM 85.1262)   hesternus (a)   Camelops (b)

Length of di1                33              --              --
Width of di1                 18              --              --
Length of di2                32              --              --
Width of di3                 19              --              --
Length of i3                 34              --              --
Width of di3                 12              --              --
Length of dp3               22.2            17.5             --
Length of dp4               58.4            50.5             --
Greatest width of dp4       18.5             14              --
Length of m1              47.3-50.7         36.7            39.5

Greatest width of m1        17.8            27.7            18.9

Length of right              267             --              --
  horizontal ramus
Length of symphysis          >92             --              --

                        Camelops cf.     Camelops
     Measurement        hesternus (c)   sulcatus (d)   Camelops (e)

Length of di1                --             --             --
Width of di1                 --             --             --
Length of di2                --             --             --
Width of di3                 --             --             --
Length of i3                 --             --             --
Width of di3                 --             --             --
Length of dp3                --             --             --
Length of dp4                --             --             --
Greatest width of dp4        --             --             --
Length of m1             30.1, 41.5    32.4, 32.6,    28.2, 38.7,
                                           31.8           43.7
Greatest width of m1     19.7, 18.6    18.3, 21.2,    19.9, 21.5,
                                           20.2           22.3
Length of right              --             --             --
  horizontal ramus
Length of symphysis          --             --         117, 121,
                                                          113

                              Camelops
                            traviswhite (f)
     Measurement        (permanent incisors)

Length of di1                   53.5
Width of di1                    19.2
Length of di2                    47
Width of di3                    16.3
Length of i3                     45
Width of di3                    19.4
Length of dp3                    --
Length of dp4                    --
Greatest width of dp4            --
Length of m1                     --

Greatest width of m1             --

Length of right                  --
  horizontal ramus
Length of symphysis              --

(a) From Selby-Dutton, Yuma County, Colorado (Graham, 1981).

(b) From Roosevelt County, New Mexico (Lundelius, 1972).

(c) From Power County, Idaho (McDonald and Anderson, 1975).

(d) From Lubbock County, Texas (Dalquest, 1967).

(e) From Rancho la Brea, Los Angeles County, California (Webb, 1965).

(f) From Aquascalientes, Mexico (Mooser and Dalquest, 1975b).
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Author:Pasenko, Michael R.; Agenbroad, Larry D.
Publication:Southwestern Naturalist
Article Type:Report
Geographic Code:1USA
Date:Mar 1, 2012
Words:8964
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