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Taxonomic and Evolutionary Affinities of Papio Izodi Fossils from Taung and Sterkfontein

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Taxonomic and Evolutionary Affinities of Papio Izodi Fossils from Taung and Sterkfontein Palaeont. afr., 30, 43-49 (1993) TAXONOMIC AND EVOLUTIONARY AFFINITIES OF PAPIO IZODI FOSSILS FROM TAUNG AND STERKFONTEIN Jeffrey K. McKee Department ofAnatomy and Human Biology, University of the Witwatersrand, Medical School, 7 York Rd., Parktown, Johannesburg, 2193. ABSTRACT Papio izodi is an extinct papionin found at Taung and Sterkfontein. The taxonomic status of southern African fossils sometimes referred toP. izodi is clarified here in order to verify the existence of the species at Sterkfontein and define the morphological characteristics distinguishing it from P. angusticeps, a later species of similar size. P. izodi may be the earliest known species of the genus Papio in southern Africa, as the putative contemporary presence of the derived species Papio hamadryas robinsoni cannot be confirmed at Sterkfontein. P. izodi retains some of the primitive features found in Parapapio broomi, suggesting a close evolutionary link between the two species. KEYWORDS: Cercopithecid taxonomy, faunal correlation, Taung, Sterkfontein. INTRODUCTION framework and to verify its assignment with the additional Papio izodi is an extinct species of baboon named by data. Gear (1926) on the basis of fossil skulls from Taung cave If STS 262 can be confirmed as a representative of deposits found near the type site of Australopithecus P. izodi, then an important link would be established africanus (McKee 1993a). This Pliocene baboon species between the fauna from Sterkfontein Member 4 and is among the earliest representatives ofP apio, and perhaps Taung as one of20 species that the sites have in common the earliest member of the genus in southern Africa. It (McKee 1993b). However, Szalay and Delson (1979) was thought to have been unique to Taung, but the referred similar small baboon fossils from the younger taxonomic status of many early Papio fossils from other sites ofKromdraai (KBE Member 3) and Cooper's toP. southern African Plio-Pleistocene fossil sites remains izodi that had previously been assigned toP. angusticeps enigmatic (Eisenhart 1974; Szalay and Delson 1979; (Broom 1940, 1946; Freedman 1957). In more recent McKee 1991 ). Clarification of the taxonomic problems publications, aimed at establishing a cercopithecid and assessment of the phylogenetic issues concerning the biochronology, Delson (1984, 1988) accepts the specific origins ofthe genus P apio are fundamental steps necessary legitimacy of P. angusticeps. It is crucial to clarify the for faunal dating and environmental reconstructions of taxonomic status of the Kromdraai and Cooper's small the australopithecine fossil sites in which the extinct baboons for purposes of faunal dating and phylogenetic baboons are abundant representatives of the prehistoric reconstructions of the baboons. faunal community. Origins of the genus Papio, and faunal correlations of A study of fossil cercopithecids from Sterkfontein by southern African cave sites, are even more problematical Eisenhart (1974) revealed a single cranium of an due to the questionable identification of a much larger immature adult female (STS 262) which he felt was baboon species from Sterkfontein Member 4. This large representative of P. wellsi, a taxon which has since been species was originally known asP. robinsoni (Freedman synonomised with P. izodi (Szalay and Delson 1979). 1957), and was recognized from the later sites of STS 262, from Sterkfontein Member 4, had been referred Kromdraai, Cooper's and Swartkrans. Eisenhart (1974) to Parapapio broomi by Freedman (1957), but Eisenhart argued that some of the Sterkfontein specimens which noted that the specimen possessed the distinct Freedman ( 1957) had referred toParapapio whitei showed morphological features of Papio, within the range of characteristics of Papio robinsoni. Szalay and Delson Papio from Taung. (1979) agreed with Eisenhart that this taxon, which they P apio izodi is well represented in the Hrdlicka deposits saw as a subspecies of P. hamadryas, was represented in at Taung, with additional specimens emanating from the the Sterkfontein material. Thus Sterkfontein Member 4 is recent excavations (McKee and Tobias 1990; McKee thought to have five species of baboon, including three 1993a). As the complete Taung sample was not studied species ofParapapio (P. whitei,P. broomi andP. jonesi) by Eisenhart (1974), and as P. wellsi has been and two species ofP apio (P. izodi andP .h. robinsoni). As synonomised with P. izodi, it becomes necessary to Sterkfontein and Taung are the earliest known African clarify the status of STS 262 within the current taxonomic sites to have yielded specifically identifiable Papio, the 44 baboon fossils from these sites may give clues to the (A and B) and Cooper's. These were used for evolutionary origin of the genus, provided that the comparisons with STS 262. taxonomic referrals are beyond a reasonable doubt. Limitations of the data set and the fossil record led to Ecological constraints on the number of similarly the exclusion of many possible measurements. The sized and similarly adapted species has led a number of excluded variables were not found to be useful in authors to doubt the probability of five contemporaneous distinguishing the species in question, or were of an species having existed at Sterkfontein (Freedman 1976; insufficient sample size for analysis. Data on mandibles Eisenhart 1974; McKee 1991). Either the environmental were excluded as few could be reliably associated with parameters are unique or the taxonomic referrals are the skulls that show taxonomically diagnostic features. incorrect. Part of this problem may be resolved with an Variables used in this analysis are listed and briefly examination of the fossils referred toPapio, demarcation described in Table 1; complete descriptions of the of the diagnostic characteristics of taxa, and an assess­ methods used in measuring can be found in Freedman ment of the evolutionary origins of the genus. (1957) and Eisenhart (1974). The data presented on Verification of the taxonomic and evolutionary bilateral traits are the means of the left and right sides affinities of fossils referred to P. izodi requires an where both sides were present, or are derived from assessment of three key issues: 1) Does P. izodi exist in unilaterally preserved structures. Some measurements the Sterkfontein Member 4 fossil deposits, as represented are estimated for cases in which slight damage (e.g. a by STS 262; 2) what are the definitive morphological missing flake of enamel or bone) still allows for a differences between P. izodi fossils and the similarly reasonable approximation to be made. sized papionins found at Kromdraai and Cooper's Evaluations of the similarities and differences among sometimes referred toP. angusticeps; and 3) are there species measurements were done on the basis of known two species of the genus Papio represented in ranges. Difference of sample ranges beyond that which Sterkfontein Member 4? one would expect on the basis of intraspecific variations within modem baboons were deemed to be biologically MATERIALS AND METHODS significant. As sample sizes were very small, no Odontometric and craniometric variables were statistical tests for significant differences were possible. compiled into a data base of measurements from Definitive non-metric morphological traits were also published descriptions of fossil baboons (Freedman observed in the fossils to supplement the data base, as 1957, 1961, 1962 1976; Freedman and Brain 1972; described in the text below. The reference sample for Eisenhart 1974), as well as from the author's own these observations included crania as cited above, as well measurements of the material. The reference sample for as crania referred to Parapapio whitei, Parapapio this assessment includes 12 Papio izodi crania from broomi, and P apio robinsoni by Freedman ( 1957, 1960, Taung and 28 Papio angusticeps crania from Kromdraai 1961, 1962, 197 6) and Eisenhart ( 197 4) for comparisons with the Sterkfontein fossils SWP 31 and SWP 3076. TABLEl Cranial Measurements COMPARATIVE ANALYSIS OF THE FOSSIL GL-IDS Glabella to Alveolare (infradentale superius) DATA IDS-NA Alveolare to Nasion IDS-PNS Alveolare to Post-nasal Spine Taxonomic Affinities of STS 262 MAXTEMP Maximum Temporal Breadth Observations of STS 262 show it to be a well­ INTORB Inter-orbital Breadth (minimum) INTFRBR Inter-frontal Breadth (minimum breadth posterior to preserved partial cranium of an immature adult female supraorbital tori) (Figure 1A).It consists of most of the calvarium, missing NAPL Nasal Aperture Length only the occipital region, and a fairly complete face with NAPB Nasal Aperture Breadth damage to the zygomatic arches and lateral orbits. The MBM2M Maximum Breadth of Maxilla at Mesialloph of W left P3 is preserved; the M1 and M2 on each side are PBM2M Palatal Breadth at Mesial Loph of W present, although some enamel has flaked off. Neither M3 PBM2D Palatal Breadth at Distal Loph of M2 BA-BREG Cranial Height from Basion to Bregma had erupted before death. ORBHT Orbit Height (maximum) Nearly all of the features of STS 262 fit within range of ORBBR Orbit Breadth (maximum) size and morphology of Parapapio broomi, hence its MHM2M Maxillary Height at Mesial Loph of W referral to that taxon by Freedman (1957). Eisenhart (1974: 529) listed four convincing characteristics Dental Measurements (Maxillary) distinguishing the specimen from Parapapio: "1) the P3B P3 breadth MlL M' length large size of the glabella region 2) the steep interorbital MlMB M' breadth, mesialloph drop on the
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