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The Partial Skeleton of Ardipithecus Ramidus TOTEM 137 The Partial Skeleton of Ardipithecus brief mention of a recovered partial skeleton Ramidus (White, Suwa, and Asfaw 1995). It could only be assumed that this taxonomic Arthur Klages revision implied that the new material must have been morphologically and adaptively Introduction distinct from Australopithecus. Paleoanthro- In 1994 a new species of pologists eagerly awaited the description of Australopithecus was described that was the partial skeleton, especially with the purported to be the new stem hominin1 and implications arising from the new genus, but the ancestor of the then widely recognized the wait for the publication would be a stem hominin, Australopithecus afarensis protracted one. (White, Suwa, and Asfaw 1994). Mostly a It was not until 2009 that the collection of dental and mandibular aforementioned partial skeleton was fragments, accompanied by a portion of the described in a publication. Bestowed with basicranium, the newly minted the appellation “Ardi”, in perhaps a con- Australopithecus ramidus was dated to 4.4 scious attempt to capture the notoriety of the Ma, and, thus, became the oldest famous Lucy skeleton, the partial skeleton australopithecine on record (White, Suwa, ARA-VP-6/500 contained more skeletal and Asfaw 1994). The hominin status of Au. elements than Lucy, but was plagued by ramidus, or rather its bipedality, was extreme fragmentation and horrendous inferred from its morphological similarity to presservation (White et al. 2009). This was Au. afarensis and the inferior orientation of given as the main reason for the delay in the foramen magnum (White, Suwa, and publication (White et al. 2009). A series of Asfaw 1994). As Au. afarensis was already eleven articles described the new Ar. established as an obligate biped2, it was ramidus material and these descriptions suggested that Au. ramidus was likely a relied heavily on virtual reconstructions of bipedal hominin (White, Suwa, and Asfaw the fragmentary fossils using micro-CT 1994). However, it was clear that the new technology. A critical analysis of the virtual species not only preceded Au. afarensis reconstructions is necessary, as the morpho- chronologically, but that it also displayed a logical inferences found among the more primitive morphology. Au. ramidus descriptions are only as solid as the quality lacked the postcanine megadontia3 of of the reconstructions and one’s confidence Australopithecus, had thinner molar enamel, in them. and had more ape-like canines, which From the virtual reconstructions, it was inferred by the authors that Ar. ramidus resulted in the naming of a new specific 4 taxon to distinguish this material from was a hominin, a facultative biped , and was known australopithecines (White, Suwa, and most likely ancestral to Australopithecus Asfaw 1994). (White et al. 2009). These conclusions were Shortly thereafter, a corrigendum identical to those already postulated in the reassigned this material to a new genus, and original 1994 descriptions based on only a the “root” australopithecine became the few skeletal elements, and perhaps that was “root ground ape” Ardipithecus ramidus to be expected (White, Suwa, and Asfaw (White, Suwa, and Asfaw 1994, 1995). 1994). What was surprising was the There was little justification of this taxo- extremely primitive morphology of Ar. nomic revision in the corrigendum, and no ramidus relative to Australopithecus (White new material was described, other than a et al. 2009). In spite of claims to facultative TOTEM vol 19 2010-2011 TOTEM 138 terrestrial bipedalism, Ar. ramidus retained a behaviourally, Ar. ramidus was unlike the completely divergent hallux5 (White et al. extant hominids, and consequently, the 2009). Its primitive foot and hand discovery of Ar. ramidus questions their morphology suggested that it retained a value as referential models for the LCA substantial arboreal component in its (White et al. 2009). The analysis of Ar. locomotive repertoire (White et al. 2009). ramidus suggested instead that the LCA There were no adaptations for knuckle- was, in fact, a generalized arboreal walking, vertical climbing, or suspension to clambering ape (White et al. 2009). be found in the skeleton, and this suggested In summary, Ar. ramidus was seen to Ar. ramidus was a palmigrade clamberer, a be far more primitive than Au. afarensis and generalized form of arboreal hominid yet derived enough relative to the hominids locomotion (White et al. 2009). for it to be a considered a hominin (White et Their claims for a hominin status al. 2009). It supposedly resembled the rested most notably on the derived morphology of the LCA more than any other characteristics of the pelvis and foot that taxa with known postcranial elements supposedly enabled Ar. ramidus to walk (White et al. 2009). Yet, it was strongly bipedally, and the derived characteristics of inferred that Ar. ramidus was both ancestral a vestigial canine/premolar complex (White to Australopithecus and morphologically et al. 2009). The honing complex6 was similar to Sahelanthropus, and that it could significantly reduced from that of extant and be easily accommodated into a linear extinct hominids, yet was primitive relative account of hominin evolution (Lovejoy et al. to Australopithecus (White et al. 2009). As a 2009c; White et al. 2009). However, as corollary to the canine reduction, it was many of these claims question the suggested that Ar. ramidus lacked significant conventional wisdom of hominin evolution body-size dimorphism, with Ardi herself and in light of proposed alternative weighing in at a hefty 50 kg, compared to theoretical scenarios of hominin evolution, the diminutive Lucy’s 30 kg estimate (White the Ar. ramidus material requires a detailed et al. 2009). The lack of sexual dimorphism examination. The level of confidence in the was a surprising claim, as Au. afarensis as virtual reconstructions is crucial to any well as all other extant and extinct hominids interpretations of Ar. ramidus. The in- are, to varying degrees, sexually dimorphic ferences made about Ar. ramidus are largely in both canines and body size (Plavcan based on these reconstructions, and these 2001). interpretations are only valid if the It was also suggested that Ar. reconstructions are sound. Finally, any ramidus differs from the extant hominines in interpretations of Ar. ramidus need to be exhibiting primitive hominid characteristics. placed within the context of what is known This implies that the extant hominines are, about other penecontemporaneous hominin in fact, highly derived in the morphology of and hominine taxa, as well as within their pelvis and limbs, and have acquired competing theoretical approaches to their unique adaptations to vertical climbing hominin evolution. In an attempt to and knuckle-walking convergently and only minimize bias and avoid presupposing any after the Pan and the Gorilla ancestors particular hypothesis in regards to Ar. diverged from the LCA (White et al. 2009). ramidus, the morphology of this taxon will The lack of a honing complex combined be compared to all relevant hominin and with body-size monomorphism and the hominine taxa. Only then will conclusions novel locomotor repertoire suggested that, TOTEM vol 19 2010-2011 TOTEM 139 of its phylogenetic positioning and related primitive relative to Australopithecus, in inferences be offered. which only the largest specimens show Dentition traces of distal edge wear (Kimbel and The dental elements are by far the Delezene 2009; Suwa et al. 2009b). Thus, best preserved and most numerous elements Ar. ramidus is depicted as evidence of a recovered for Ar. ramidus and were linear transition in the reduction of the examined in detail (Suwa et al. 2009b). The honing complex from Ar. kadabba through most significant hominin characteristic of to Australopithecus (Suwa et al. 2009b). the Ar. ramidus dentition is the reduction of Molar enamel thickness in Ar. the canines, particularly in crown height, ramidus was found to be intermediate relative to extant and extinct hominines between the thin enamel of both (Suwa et al. 2009b). There were no Dryopithecus and extant hominines, and the statistically significant differences between thick enamel of Australopithecus (Suwa et the inferred male and female canines in size al. 2009b). Again, it was implied that Ar. or shape in the recovered sample, or in other ramidus fits within a linear progression in words, the canines were monomorphic and the increase of molar enamel thickness from could not be confidently split into two the Miocene hominines to Australopithecus sexually dimorphic clusters (Suwa et al. (Suwa et al. 2009b). The postcanine 2009b). In this regard Ar. ramidus was said dentition is primitive in size, however, and to fit the hominin pattern (Suwa et al. does not display any sign of the megadontia 2009b). The sample size of 21 canines was inherent in Australopithecus (Suwa et al. claimed to be statistically large enough so 2009b). The postcanine dentition was also that it assured that males are represented in found to be monomorphic in size and the Ar. ramidus sample, which is not the supported the claim that there was limited case for either Sahelanthropus or Orrorin sexual dimorphism present in Ar. ramidus (Brunet et al. 2002a; Senut et al. 2001; (Suwa et al. 2009b). In general, however, Suwa et al. 2009b). there seems to be only a limited amount of The overall size of the canines in Ar. sexual dimorphism in hominin and hominine ramidus are similar to that of female postcanine dentition (Plavcan 2001). Molars chimpanzees, but they differ significantly in are less dimorphic in hominins and their shape—they are diamond-like, rather hominines than they are in the other than dagger-like in Pan or other hominines anthropoids, and it is harder to infer the sex (Suwa et al. 2009b). There is no evidence of of a specimen based on the postcanine a functional canine/premolar honing dentition in these taxa (Plavcan 2001). complex in Ar. ramidus, but there are It was, rather, the canines that played vestiges of a diastema7 (Suwa et al.
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