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Mammuthus Meridionalis (Proboscidea: Elephantidae) From George E. McDaniel, Jr. & George T. Jefferson California Department of Parks and Recreation, Borrego Springs Mammuthus meridionalis (Proboscidea: Elephantidae) from the Borrego Badlands of Anza-Borrego Desert State Park®, California: phylogenetic and biochronologic implications McDaniel, G.E., Jr. & Jefferson, G.T., 2003 - Mammuthus meridionalis (Proboscidea: Elephantidae) from the Borrego Badlands of Anza-Borrego Desert State Park®, California: phylo- genetic and biochronologic implications - in: Reumer, J.W.F., De Vos, J. & Mol, D. (eds.) - ADVANCES IN MAMMOTH RESEARCH (Proceedings of the Second International Mammoth Conference, Rotterdam, May 16-20 1999) - DEINSEA 9: 239-252 [ISSN 0923-9308] Published 24 May 2003 A nearly complete skeleton of a mature female southern mammoth was recovered from the Borrego Badlands in Anza-Borrego Desert State Park®, California, USA. The remains represent a late evolutionary stage of the Mammuthus meridionalis (NESTI, 1825) chronocline, and constitute the most complete skeleton of this taxon in North America. Magnetostratigraphy and tephrochro- nology date the specimen at approximately 1.1 Ma BP. The osteology and osteometrics are pre- sented herein. This late stage of Mammuthus meridionalis co-existed with an early evolutionary stage of M. columbi (= M. imperator of some authors). Significant morphological differences between the two taxa from the Borrego Badlands suggest that M. armeniacus rather than M. meri- dionalis was immediately ancestral to M.columbi. A dispersal of M. armeniacus to North America approximately 1.2 Ma BP is implied. At least two mammoth lineages (excluding Mammuthus pri- migenius) are seen in North America. An early M. meridionalis lineage and a later M. armeniacus - M. columbi lineage. M. meridionalis is recognized as the earliest species of mammoth in North America. Correspondence: California Department of Parks and Recreation, Colorado Desert District Stout Research Center, Anza-Borrego Desert State Park, 200 Palm Canyon Drive, Borrego Springs, California 92004, U.S.A; e-mail: [email protected] Keywords: Mammuthus meridionalis, taphonomy, osteology, phylogeny INTRODUCTION most complete skeleton of this taxon from In December of 1986, the skull, mandible and North America. about 70% of the skeleton of an adult female The site, ABDSP(IVCM) 1277, lies east of southern mammoth, Mammuthus meridionalis Borrego Springs and south of San Diego (NESTI, 1825) was discovered in the Borrego County road S22 (Fig. 1). Here, the fluvial- Badlands of northern Anza-Borrego Desert floodplain deposits of the Ocotillo Conglo- State Park® (ABDSP), California, USA by G. merate (Dibblee 1954, Remeika & Pettinga Miller of Imperial Valley College Museum 1991, Remeika 1992) have yielded a diverse (El Centro, California) (Miller et al. 1988; Irvingtonian North American Land Mammal Miller 1989; Parks et al. 1989; Miller et al. Age terrestrial vertebrate assemblage, describ- 1991; McDaniel & Jefferson 1997). The spe- ed as the Borrego local fauna by Remeika & cimen, ABDSP(IVCM) V5126, represents the Jefferson (1993), and Remeika et al. (1995). 239 ADVANCES IN MAMMOTH RESEARCH DEINSEA 9, 2003 Table 1 Dental measurements of ABDSP(IVCM) V5126. Explanation: TP = tooth position; L = length in mm; W = width in mm; CH = crown height in mm; PN = number of plates, + indicates minimum number, ( ) estimated number based on stage of wear; LF = lamellar frequency, number of plates / 10 cm, ( ) number of averaged measurements; ET = enamel thickness in mm, ( ) number of averaged measurements; M3/L = upper M3, left; M3/R = upper M3, right; M/3L = lower M3, left; M/3R = lower M3, right. A thorough taphonomic study reveals that and gently curving with an Index of Curva- carnivores and scavengers prior to burial had ture significantly less than in M. imperator heavily damaged the carcass. Magneto- and M. columbi (Fig. 2). These characters stratigraphic and tephrochronologic methods distinguish M. meridionalis from the later date the specimen at approximately 1.1 Ma species (Agenbroad, pers. comm. 1996). The BP (Remeika & Beske-Diehl 1996). corpus of the mandible in ABDSP(IVCM) ABDSP(IVCM) V5126 represents a very V5126 is gracile rather than robust, and has a late evolutionary stage of Mammuthus meri- forward projecting mandibular symphyseal dionalis. The molar dental parameters fall process plus chin that is not turned down as well within the reported range of M. meridio- in M. imperator. The ascending ramus of nalis (see Maglio 1973) (Table 1). The skull ABDSP(IVCM) V5126 is low and inclined and mandible are shorter antero-posteriorly slightly posteriorly. In contrast, the ascending and wider than that in the imperial mammoth, ramus of M. imperator has rotated anteriorly. M. imperator (LEIDY, 1858). The incisor in Significant morphologic differences between ABDSP(IVCM) V5126 is thin, relatively long ABDSP(IVCM) V5126 and Mammuthus Figure 1 Northern Anza-Borrego Desert State Park®, and the Borrego Badlands of eastern San Diego County, California. Explanation: SRC = Colorado Desert District Stout Research Center; S22 = county road 22; X = locality ABDSP(IVCM) 1277; dashed outlined area = Borrego Badlands. 240 MCDANIEL & JEFFERSON: Mammuthus meridionalis from Borrego Badlands 1 (N /2 of section 36, T10S R7E, Fonts Point 7.5 minute USGS topographic quadrangle) is transected on the southern side from south- west to northeast by the Valle Escondido anti- cline (Remeika, pers. comm. 1996). This structure plunges gently to the northeast, exhibits axial faulting, and has a very steeply dipping north limb which approaches vertical. The south limb steepens to over 300 near the axis of the anticline. Locality ABDSP(IVCM) 1277 lies south of the axis, where the strata strike approximately N260 E and dip gently 50 to 60 to the southeast. Stratigraphically, ABDSP(IVCM) V5126 was recovered from a predominately fluvial Figure 2 Scattergram of greatest length of incisor in cm / horizon that complexly interfingers with the Index of Curvature. Explanation: X = Mammuthus meridiona- lis ABDSP(IVCM) V5126; + = M. imperator ABDSP(IVCM) lacustrine Brawley Formation (Dibblee V4056; black dots = M. columbi (Agenbroad 1994); open cir- 1954). Individual beds pinch out to the east cles = M. primigenius (Haynes 1991; Kunz 1916). within the Brawley Formation (Remeika, pers. comm. 1996). The paleozoic metamor- imperator, as represented by ABDSP(IVCM) phic and mesozoic crystalline rocks in the V4214 and V4260, recovered from the same Santa Rosa Mountains to the north and north- stratigraphic horizon in the Ocotillo Conglo- west provide a source for the sandy fossilife- merate, make an ancestor-descendant rela- rous sediments that crop out on both limbs of tionship (Maglio 1973; Madden 1981) the Valle Escondido anticline. between these species highly unlikely. This implies that the steppe mammoth, M. arme- DEPOSITIONAL ENVIRONMENT niacus (FALCONER, 1857), rather than M. AND TAPHONOMY meridionalis, was immediately ancestral to Apparently, the skeleton was deposited within M. imperator. Mammuthus meridionalis is a 1m deep, ephemoral, braided stream chan- recognized as the earliest species of mam- nel system. The texture, sedimentary structu- moth in North America. As first suggested by res and bedding of these deposits are typical Harington (1984), it follows that M. meridio- nalis represents the end of one lineage, and M. imperator the beginning of a second lineage that arrived in North America approximately 1.2 Ma BP. GEOLOGIC SETTING AND STRATIGRAPHY ABDSP(IVCM) V5126 was collected in a geologically complex area south of the east end of the seismically active San Jacinto Fault zone. This setting has made stratigrap- hic correlations with dated horizons within the Ocotillo Conglomerate difficult (Remeika & Beske-Diehl 1996). The general location Figure 3 Bone bed plan view, ABDSP(IVCM) 1277/V5126 (after Miller et al. 1991) (see element numbers,Table 2). 241 ADVANCES IN MAMMOTH RESEARCH DEINSEA 9, 2003 Table 2 Represented skeletal elements of ABDSP(IVCM) V5126. Materials are listed by skeletal element number (see Fig. 3). 242 MCDANIEL & JEFFERSON: Mammuthus meridionalis from Borrego Badlands of distal bajada, fluvial floodplain deposition- ments (Fig. 3, Table 2) indicate that the car- al environments. Local crossbedding and cass had largely decomposed and the remains channel direction at the excavation site sug- had been moved prior to burial. The scatter of gests a north/northeast to south/southwest skeletal elements, separation of once associa- current. Oxidization of iron-bearing minerals ted elements, the lack of many distal foot directly below most of the skeletal elements bones, and inferred depositional conditions, suggests that the fluvial channel may have suggests a short duration transport within the been dry for an extended period before depo- fluvial system after carcass decomposition. sition of the fossil-bearing layer. Also, some ABDSP(IVCM) 1277 was buried in fine- scouring or erosion occurred on the surface at grained siltstones, indicating low energy the time ABDSP(IVCM) V5126, was deposi- transport conditions that are consistent with ted. Overlying the specimen are distal fan Behrensmeyer's (1988) small scale, braided sheet flood deposits. stream channel-fill deposits. The lack of evi- There is no readily identifiable cause of dence of sediment abrasion (Olsen & Ship- death. Based on the extensive wear on the man 1988) on bone surfaces (Miller et al. anterior plates of the last molars (third 1991) is consistent with
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