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Recent fossil discoveries of early human ancestors from paleoanthropological sites in Africa and elsewhere have demonstrated how various phases of human evolutionary history were much more complica...

Fossil discoveries of early Australopithecus species from Woranso-Mille have played a significant role in improving our understanding of mid-Pliocene hominin evolution and diversity. Here, we describe two mandibles with dentitions, recovered from sediments immediately above a tuff radiometrically dated to 3.76 ± 0.02 Ma, and assess their taxonomic affinity. The two mandibles (MSD-VP-5/16 and MSD-VP-5/50) show morphological similarities with both Australopithecus anamensis and Australopithecus afarensis. Some of the unique features that distinguish Au. anamensis from Au. afarensis are present in the mandibles, which also share a few derived features with Au. afarensis. Their retention of more Kanapoi Au. anamensis-like traits, compared to the fewer derived features they share with Au. afarensis, and the presence of Au. anamensis at Woranso-Mille in 3.8-million-year-old deposits, lends support to their assignment to Au. anamensis. However, it is equally arguable that the few derived dentognathic features they share with Au. afarensis could be taxonomically more significant, making it difficult to conclusively assign these specimens to either species. Regardless of which species they are assigned to, the mosaic nature of the dentognathic morphology and geological age of the two mandibles lends further support to the hypothesized ancestor-descendant relationship between Au. anamensis and Au. afarensis. However, there is now limited fossil evidence indicating that these two species may have overlapped in time. Hence, the last appearance of Au. anamensis and first appearance of Au. afarensis are currently unknown. Recovery of Australopithecus fossils from 4.1 to 3.8 Ma is critical to further address the timing of these events.

Many important Pliocene hominin specimens have been recovered from Woranso-Mille, a paleontological research area in the Afar region of Ethiopia, including the complete cranium of Australopithecus anamensis, a partial skeleton of Australopithecus afarensis, mandibular and maxillary elements representing a new species, Australopithecus deyiremeda, and a partial foot of an as-yet-unnamed species. Woranso-Mille is the only site, so far, to have reported the co-existence of more than one early hominin species between 3.8 and 3.3 Ma and the temporal overlap between A. anamensis and A. afarensis. Thus, the site has important implications for our understanding of the paleoecology and taxonomic diversity of early hominins and their ecological niche. This paper explores the paleohabitats of Woranso-Mille through its faunal community ecological structure and taxonomic composition using correspondence analysis and Forbes modified similarity index. The results suggest that Pliocene Woranso-Mille was a mosaic of different habitat types, with riparian woodland and floodplain grassland along rivers draining into a lake, along with less mesic habitats such as woodland, grassland, and shrubland. The apparent high level of vegetation heterogeneity may have promoted dietary specializations and niche differentiation among the different Australopithecus species at Woranso-Mille and allowed for their co-existence at the site.

Suids are among the most common mammalian groups in the Plio-Pleistocene vertebrate fossil record of Africa and the most studied largely due to their significance as biochronological indicators. However, despite their abundance in the fossil record, the remains are mostly isolated teeth and fragmentary crania and mandibles. As a result, disagreements have persisted in terms of their taxonomy and phylogenetic relationships. Here, we present for the first time a detailed description of the cranial anatomy of Nyanzachoerus jaegeri based on two crania recovered from middle Pliocene deposits of the Woranso-Mille paleontological study area, Afar region, Ethiopia. Understanding the cranial morphology of this species is particularly significant given the recent reclassification of Nyanzachoerus jaegeri to the genus Notochoerus based largely on the incisor and symphyseal morphology of specimens from Kanapoi, Kenya. Here, we show that the two genera are clearly distinguished from each other by distinct morphological features of the cranium such as the shape of the braincase, orientation of the zygomatic arches, and premolar/molar ratio, among others. Furthermore, we show that the mandibular and dental morphological features identified by some workers as characteristic of Notochoerus are variable among tetraconodont species and that Nyanzachoerus jaegeri best fits within the genus Nyanzachoerus.

The Woranso–Mille (WORMIL) area in the west-central Afar, Ethiopia, contains several Pliocene basalt flows, tuffs, and fossiliferous volcaniclastic beds. We present whole-rock major- and trace-element data including REE, and Sr–Nd–Pb isotope ratios from these basalts to characterize the geochemistry, constrain petrogenetic processes, and infer mantle sources. Six basalt groups are distinguished stratigraphically and geochemically within the interval from ~ 3.8 to ~ 3 Ma. The elemental and isotopic data show intra- and inter-group variations derived primarily from source heterogeneity and polybaric crystallization ± crustal inputs. The combined Sr–Nd–Pb isotope data indicate the involvement of three main reservoirs: the Afar plume, depleted mantle, and enriched continental lithosphere (mantle ± crust). Trace element patterns and ratios further indicate the basalts were generated from spinel-dominated shallow melting, consistent with significantly thinned Pliocene lithosphere in western Afar. The on-land continuation of the Aden rift into western Afar during the Pliocene is reexamined in the context of the new geochemistry and age constraints of the WORMIL basalts. The new data reinforce previous interpretations that progressive rifting and transformation of the continental lithosphere to oceanic lithosphere allows for increasing asthenospheric inputs through time as the continental lithosphere is thinned.

International audience; A fossil hominin cranium was discovered in mid-Pliocene deltaic strata in the Godaya Valley of the northwestern Woranso-Mille study area in Ethiopia. Here we show that analyses of chemically correlated volcanic layers and the palaeomagnetic stratigraphy, combined with Bayesian modelling of dated tuffs, yield an age range of 3.804 ± 0.013 to 3.777 ± 0.014 million years old (mean ± 1σ) for the deltaic strata and the fossils that they contain. We also document deposits of a perennial lake beneath the deltaic sequence. Mammalian fossils associated with the cranium represent taxa that were widespread at the time and data from botanical remains indicate that the vegetation in the lake and delta catchment was predominantly dry shrubland with varying proportions of grassland, wetland and riparian forest. In addition, we report high rates of sediment accumulation and depositional features that are typical of a steep topographic relief and differ from younger Woranso-Mille fossil localities, reflecting the influence of active rift processes on the palaeolandscape. Q1 Until recently, field work in the Woranso-Mille area (Afar, Ethiopia) has concentrated on exposures along the Mille River that include more than 120 m of fossiliferous sedimentary and volcanic strata that are between 3.8 and 3.2 million years (Myr) old. East of Korsi Dora (Fig. 1a), sedimentary sequences that are younger than the extra-regional approximately 3.57-Myr-old Kilaytoli tuff 1 have yielded specimens of at least two hominin species, Australopithecus afarensis and Australopithecus deyiremeda 2-5

A fossil hominin cranium was discovered in mid-Pliocene deltaic strata in the Godaya Valley of the northwestern Woranso-Mille study area in Ethiopia. Here we show that analyses of chemically correlated volcanic layers and the palaeomagnetic stratigraphy, combined with Bayesian modelling of dated tuffs, yield an age range of 3.804 ± 0.013 to 3.777 ± 0.014 million years old (mean ± 1σ) for the deltaic strata and the fossils that they contain. We also document deposits of a perennial lake beneath the deltaic sequence. Mammalian fossils associated with the cranium represent taxa that were widespread at the time and data from botanical remains indicate that the vegetation in the lake and delta catchment was predominantly dry shrubland with varying proportions of grassland, wetland and riparian forest. In addition, we report high rates of sediment accumulation and depositional features that are typical of a steep topographic relief and differ from younger Woranso-Mille fossil localities, reflecting the influence of active rift processes on the palaeolandscape.

Fossiliferous deposits at Woranso-Mille span the period when Australopithecus anamensis gave rise to Australopithecus afarensis (3.8–3.6 Ma) and encompass the core of the A. afarensis range (ca. 3.5–3.2 Ma). Within the latter period, fossils described to date include the intriguing but taxonomically unattributed Burtele foot, dentognathic fossils attributed to Australopithecus deyiremeda, and one specimen securely attributed to A. afarensis (the Nefuraytu mandible). These fossils suggest that at least one additional hominin lineage lived alongside A. afarensis in the Afar Depression. Here we describe a collection of hominin fossils from a new locality in the Leado Dido’a area of Woranso-Mille (LDD-VP-1). The strata in this area are correlated to the same chron as those in the Burtele area (C2An.3n; 3.59–3.33 Ma), and similar in age to the Maka Sands and the Basal through lower Sidi Hakoma Members of the Hadar Formation. We attribute all but one of the LDD hominin specimens to A. afarensis, based on diagnostic morphology of the mandible, maxilla, canines, and premolars. The LDD specimens generally fall within the range of variation previously documented for A. afarensis but increase the frequency of some rare morphological variants. However, one isolated M3 is extremely small, and its taxonomic affinity is currently unknown. The new observations support previous work on temporal trends in A. afarensis and demonstrate that the large range of variation accepted for this species is present even within a limited spatiotemporal range. The value added with this sample lies in its contribution to controlling for spatiotemporal differences among site samples in the A. afarensis hypodigm and its contemporaneity with non-A. afarensis specimens at Woranso-Mille.

Woranso-Mille is a paleoanthropological site in Ethiopia sampling an important and under-represented time period in human evolution (3.8–3.6 million years ago). Specimens of cf. Australopithecus anamensis, Australopithecus afarensis, and the recently named Australopithecus deyiremeda have been recovered from this site. Using multiple habitat proxies, this study provides a paleoecological reconstruction of two fossiliferous collection areas from Woranso-Mille, Aralee Issie (ARI) and Mesgid Dora (MSD). Previous reconstructions based on faunal assemblages have pointed, due to the presence of aepycerotins, alcelaphins, and proboscideans, to the existence of open habitats as well as more closed ones, based on the occurrence of cercopithecids, giraffids, and traglephins. Results from community structure analysis (proportions of locomotor and dietary adaptations) at ARI and MSD indicated a predominance of open habitats, such as shrublands. Mesowear analysis revealed that ungulates of all dietary types (grazers, leaf and fruit browsers, and mixed feeders) were present in nearly equal proportions. Ecomorphological analyses using linear measurements of the astragalus and phalanges indicated that bovids utilizing locomotor behaviors associated with all habitat types were present, though the intermediate-cover habitat bovids were best represented in the sample (Heavy cover at ARI and Light cover at MSD). Together, these results suggest that the ARI and MSD localities were heterogeneous habitats (mosaics), likely with densely vegetated areas along a paleo-river and more open regions (woodlands, grasslands) available away from the river.

Mandibular morphology of Australopithecus afarensis is well known based on abundant fossil mandibles of adult individuals from multiple sites in Ethiopia (Hadar, Woranso-Mille, and Middle Awash) and Tanzania (Laetoli). However, there are only a few juvenile mandibles of the species known from these sites. Here, we describe two recently discovered Pliocene hominin juvenile mandibles from Woranso-Mille (KSD-VP-1/29 and MKM-VP-1/626), that have been radioisotopically dated to 3.6 million years ago. We assign these mandibles to Australopithecus afarensis based on their possession of mandibular morphological features considered as distinctive of the species. These specimens not only increase the sample size of juvenile mandibles of A. afarensis from poorly known time period, but also add new information on the degree of variability in juvenile mandibular morphology within the species. Their dentition samples the whole spectrum of size and shape variation in A. afarensis. Our analysis further indicates that symphyseal morphology of the Laetoli mandibles of early A. afarensis falls within the range of variation of mandibles of the species from Hadar, Woranso-Mille, and Middle Awash. Moreover, the fact that the new 3.6 million-year-old juvenile mandibles from Woranso-Mille have a more receeding symphysis than any of the juvenile and most adult mandibles of A. afarensis from the younger Hadar sequence lends support to the observed trend in symphyseal morphological change in the A. anamensis–A. afarensis chronospecies lineage.

The incorporation of C4 resources into hominin diet signifies increased dietary breadth within hominins and divergence from the dietary patterns of other great apes. Morphological evidence indicates that hominin diet became increasingly diverse by 4.2 million years ago but may not have included large proportions of C4 foods until 800 thousand years later, given the available isotopic evidence. Here we use carbon isotope data from early to mid Pliocene hominin and cercopithecid fossils from Woranso-Mille (central Afar, Ethiopia) to constrain the timing of this dietary change and its ecological context. We show that both hominins and some papionins expanded their diets to include C4 resources as early as 3.76 Ma. Among hominins, this dietary expansion postdates the major dentognathic morphological changes that distinguish Australopithecus from Ardipithecus, but it occurs amid a continuum of adaptations to diets of tougher, harder foods and to committed terrestrial bipedality. In contrast, carbon isotope data from cercopithecids indicate that C4-dominated diets of the earliest members of the Theropithecus oswaldi lineage preceded the dental specialization for grazing but occurred after they were fully terrestrial. The combined data indicate that the inclusion of C4 foods in hominin diet occurred as part of broader ecological changes in African primate communities.

Middle Pliocene hominin species diversity has been a subject of debate over the past two decades, particularly after the naming of Australopithecus bahrelghazali and Kenyanthropus platyops in addition to the well-known species Australopithecus afarensis. Further analyses continue to support the proposal that several hominin species co-existed during this time period. Here we recognize a new hominin species (Australopithecus deyiremeda sp. nov.) from 3.3-3.5-million-year-old deposits in the Woranso-Mille study area, central Afar, Ethiopia. The new species from Woranso-Mille shows that there were at least two contemporaneous hominin species living in the Afar region of Ethiopia between 3.3 and 3.5 million years ago, and further confirms early hominin taxonomic diversity in eastern Africa during the Middle Pliocene epoch. The morphology of Au. deyiremeda also reinforces concerns related to dentognathic (that is, jaws and teeth) homoplasy in Plio-Pleistocene hominins, and shows that some dentognathic features traditionally associated with Paranthropus and Homo appeared in the fossil record earlier than previously thought.

Eucyon is a paraphyletic genus of jackal-sized canids that predated the appearance of true Canis in North America, Eurasia and Africa. In this study, we describe a new species of Eucyon, E. kuta, from the Woranso-Mille Area, Afar Region, Ethiopia. The new species differs from all other described Eucyon in having the distal cusps of the p4 united into a crest. It may be the youngest Eucyon known from Africa and is larger than described Eucyon from that continent. Other described African Eucyon are briefly reviewed, and we suggest that some material may be referable to foxes (Vulpini), while other material may belong to the small bear-dog Bonisicyon illacabo (Amphicyonidae) or a closely related taxon.

The Woranso-Mille paleontological study area, located in the central Afar region of Ethiopia, is one of the most important Pliocene sites in eastern Africa. Since the Woranso-Mille Research Project began its investigation in 2005, more than 7,500 mammalian fossils, including hominins, have been collected from 80 vertebrate localities. This paper provides a preliminary assessment of the Woranso-Mille carnivoran record, a record that is of great interest given the high level of species richness of African carnivorans during the middle Pliocene. Craniodental and postcranial material of canids, lutrine mustelids, viverrids, herpestids, machairodontine and feline felids, and hyaenids has been recovered. Thus, this diverse fauna includes not only the largest carnivorans from this time period (e.g., Enhydriodon and Homotherium), but also some of the smallest, including mongooses, civets, genets, and felids, some of which represent new species. However, the diversity of small taxa does not yet approach that found in the roughly contemporaneous Upper Laetolil Beds of Tanzania. In contrast, lutrine mustelids are better represented at Woranso-Mille than at Kanapoi (Kenya) or Laetoli (Tanzania), which is to be expected given the diversity of habitats represented at these sites. While more material from these sites and others are necessary to truly understand the increased diversity within the early to middle Pliocene eastern African carnivoran guild, it is clear that the material from Woranso-Mille has the potential to fill many of the gaps in our knowledge of carnivorans during this time period.

Recent discoveries of multiple middle Pliocene hominins have raised the possibility that early hominins were as speciose as later hominins. However, debates continue to arise around the validity of most of these new taxa, largely based on poor preservation of holotype specimens, small sample size, or the lack of evidence for ecological diversity. A closer look at the currently available fossil evidence from Ethiopia, Kenya, and Chad indicate that Australopithecus afarensis was not the only hominin species during the middle Pliocene, and that there were other species clearly distinguishable from it by their locomotor adaptation and diet. Although there is no doubt that the presence of multiple species during the middle Pliocene opens new windows into our evolutionary past, it also complicates our understanding of early hominin taxonomy and phylogenetic relationships.

Australopithecus afarensis is the best-known and most dimorphic species in the early hominin fossil record. Here, we present a comparative description of new fossil specimens of Au. afarensis from Nefuraytu, a 3.330-3.207 million-years-old fossil collection area in the Woranso-Mille study area, central Afar, Ethiopia. These specimens include NFR-VP-1/29, one of the most complete mandibles assigned to the species thus far and among the largest mandibles attributed to Au. afarensis, likely representing a male individual. NFR-VP-1/29 retains almost all of the distinctive archaic features documented for Au. afarensis. These features include a posteriorly sloping symphysis, a low and rounded basally set inferior transverse torus, anterosuperiorly opening mental foramen, a lateral corpus hollow bound anteriorly by the C/P3 jugae and posteriorly by the lateral prominence, and the ascending ramus arising high on the corpus. Dental morphology and metrics of the Nefuraytu specimens also falls within the range of Au. afarensis. The presence of this species at Woranso-Mille between 3.330 and 3.207 million years ago confirms the existence of this species in the area in close spatial and temporal proximity to other middle Pliocene hominin taxa such as the one represented by the Burtele foot (BRT-VP-2/73) and the recently named species Australopithecus deyiremeda. This has important implications for our understanding of middle Pliocene hominin diversity.

Kolpochoerus (Mammalia: Suidae) is a suine genus represented by a number of species from Plio-Pleistocene sites in Africa. While the general trends in Kolpochoerus evolution are broadly known, gaps in the fossil record preclude an understanding of the details of its evolutionary tempo and mode. Here, we describe a new species, Kolpochoerus millensis, based on new fossil material from the Woranso-Mille and Gona sites in the Central Afar region of Ethiopia and dated to 3.5–3.8 million years ago (Ma). Third molars of K. millensis are metrically and morphologically intermediate between the early Pliocene K. deheinzelini and earliest late Pliocene K. afarensis. It appears that K. deheinzelini, K. millensis, and K. afarensis are temporally disjunct and phenetically distinguishable parts of a single evolving lineage. The recognition of these chronospecies provides additional evidence for anagenetic evolution. It demonstrates clearly the presence of transitional forms in the fossil record. The extensive and well-dated Kolpochoerus fossil record serves as one of the best documented examples of the occurrence of phyletic evolution. Moreover, K. millensis is one of the best biochronological markers in eastern Africa for the time between 3.5 and 3.8 Ma.

Recent fieldwork at Woranso-Mille, Ethiopia led to the recovery of an abundant, diverse mammalian fauna that includes remains of the early hominid Australopithecus afarensis. Proboscideans are among the taxa well sampled at the site, primarily by gnathodental specimens, dated to the mid-Pliocene interval of 3.8–3.6 Ma. These fossils document traces of the last anancine gomphotheres (Anancus ultimus) in eastern Africa and several elephant taxa. Comparative study of the elephant fossils indicates the presence of cf. Mammuthus sp. “Hadar-type,” cf. Elephas ekorensis, E. recki brumpti, and cf. Loxodonta adaurora adaurora. Proboscidean evolution in the mid-Pliocene is interesting because during this time archaic elephants were completely replaced by basal members of crown elephant lineages, taxonomic diversity was high (multiple elephant species, anancine gomphotheres, stegodonts, and deinotheres), and elephants were undergoing substantial reorganization of the craniodental masticatory apparatus, presumably in response to the spread of more open habitats and greater competition for grazing resources. The Woranso-Mille sample is important because this interval is only represented elsewhere in eastern Africa by a small number of sites, and because adaptive diversification among early crown elephants requires greater clarification. Morphometric contrasts among the fossil dentition from Woranso-Mille presage the differential success of elephant lineages in eastern Africa during the Pleistocene, providing hints about the beginnings of competitive displacement. Differences between E. recki brumpti from Woranso-Mille and the slightly younger Sidi Hakoma Member of the Hadar Formation reveal the beginnings of continuous, directional morphometric change that characterized the lineage. Reconsideration of E. recki subspecies indicates that they are arbitrary lineage divisions tied to geochronological boundaries (with utility for biochronological correlation at well sampled sites) rather than real phylogenetic entities, but does not reject monophyly or anagenetic evolution of the lineage.

The earliest evidence ofAustralopithecusgoes back toca4.2 Ma with the first recorded appearance ofAustralopithecus ‘anamensis’at Kanapoi, Kenya.Australopithecus afarensisis well documented between 3.6 and 3.0 Ma mainly from deposits at Laetoli (Tanzania) and Hadar (Ethiopia). The phylogenetic relationship of these two ‘species’ is hypothesized as ancestor–descendant. However, the lack of fossil evidence from the time between 3.6 and 3.9 Ma has been one of its weakest points. Recent fieldwork in the Woranso-Mille study area in the Afar region of Ethiopia has yielded fossil hominids dated between 3.6 and 3.8 Ma. These new fossils play a significant role in testing the proposed relationship betweenAu. anamensisandAu. afarensis. The Woranso-Mille hominids (3.6–3.8 Ma) show a mosaic of primitive, predominantlyAu. anamensis-like, and some derived (Au. afarensis-like) dentognathic features. Furthermore, they show that, as currently known, there are no discrete and functionally significant anatomical differences betweenAu. anamensisandAu. afarensis. Based on the currently available evidence, it appears that there is no compelling evidence to falsify the hypothesis of ‘chronospecies pair’ or ancestor–descendant relationship betweenAu. anamensisandAu. afarensis. Most importantly, however, the temporally and morphologically intermediate Woranso-Mille hominids indicate that the species namesAu. afarensisandAu. anamensisdo not refer to two real species, but rather to earlier and later representatives of a single phyletically evolving lineage. However, if retaining these two names is necessary for communication purposes, the Woranso-Mille hominids are best referred to asAu. anamensisbased on new dentognathic evidence.

(40)Ar/(39)Ar dating of tuffs and mafic lavas, tephra geochemistry, and paleomagnetic reversal stratigraphy have been used to establish the chronostratigraphy of the Pliocene hominid-bearing fossiliferous succession at Woranso-Mille, a paleontological study area in the western part of the central Afar region of Ethiopia. The succession in the northwestern part of the study area ranges in (40)Ar/(39)Ar age from 3.82-3.570 Ma, encompassed by paleomagnetic subchron C2Ar (4.187-3.596 Ma). One of the major tuff units, locally named the Kilaytoli tuff, is correlative on the basis of age and geochemistry to the Lokochot Tuff of the Turkana Basin. A hominid partial skeleton (KSD-VP-1) was found in strata whose precise stratigraphic position and age is still under investigation, but is believed to correspond to the later part of this interval. Woranso-Mille fills a significant gap in the fossil record of northeastern Africa at the time of the lower to middle Pliocene transition, when many extant species lineages of African fauna were established.

Hominid fossils predating the emergence of Australopithecus have been sparse and fragmentary. The evolution of our lineage after the last common ancestor we shared with chimpanzees has therefore remained unclear. Ardipithecus ramidus , recovered in ecologically and temporally resolved contexts in Ethiopia’s Afar Rift, now illuminates earlier hominid paleobiology and aspects of extant African ape evolution. More than 110 specimens recovered from 4.4-million-year-old sediments include a partial skeleton with much of the skull, hands, feet, limbs, and pelvis. This hominid combined arboreal palmigrade clambering and careful climbing with a form of terrestrial bipedality more primitive than that of Australopithecus . Ar. ramidus had a reduced canine/premolar complex and a little-derived cranial morphology and consumed a predominantly C 3 plant–based diet (plants using the C 3 photosynthetic pathway). Its ecological habitat appears to have been largely woodland-focused. Ar. ramidus lacks any characters typical of suspension, vertical climbing, or knuckle-walking. Ar. ramidus indicates that despite the genetic similarities of living humans and chimpanzees, the ancestor we last shared probably differed substantially from any extant African ape. Hominids and extant African apes have each become highly specialized through very different evolutionary pathways. This evidence also illuminates the origins of orthogrady, bipedality, ecology, diet, and social behavior in earliest Hominidae and helps to define the basal hominid adaptation, thereby accentuating the derived nature of Australopithecus .

The Bodo partial distal humerus with an approximate age of 0.6 million years is one of the very few postcranial remains from the African Middle Pleistocene. Despite its fragmentary status, comparisons of the Bodo humerus with other fossil hominid and modern human samples reveal some insights into African hominids of this time period. The Bodo partial humerus displays distal humeral features very common in the European Middle and Late Pleistocene hominids, such as a relatively wide olecranon fossa and relatively thin lateral and medial pillars adjacent to the fossa. In Africa, the postcranial fossils from the Middle and Late Pleistocene exhibit a surprising amount of morphological diversity. The presence of ‘typically’ Neandertal traits in some, but not all, Homo ergaster, H. Rhodesiensis, and early H. sapiens, together with the greater similarity of some African specimens than others to recent humans, highlights the problem of interpreting the anatomical variation that characterizes African fossil humans. An analysis of frequency–based patterning can help us understand this striking variation. Populations of Middle Pleistocene African hominids, such as the one represented by the Bodo specimen studied here, may have been variable, and their skeletons may have contained a mosaic of modern and non–modern human traits.

New observations on the Late Miocene and Earliest Pliocene mustelids from the Middle Awash of Ethiopia are presented. The Middle Awash study area samples the last six million years of African vertebrate evolutionary history. Its Latest Miocene (Asa Koma Member of the Adu-Asa Formation, 5.54–5.77 Ma) and Earliest Pliocene (Kuseralee and Gawto Members of the Sagantole Formation, 5.2 and 4.85 Ma, respectively) deposits sample a number of large and small carnivore taxa among which mustelids are numerically abundant. Among the known Late Miocene and Early Pliocene mustelid genera, the Middle Awash Late Miocene documents the earliest Mellivora in eastern Africa and its likely first appearance in Africa, a new species of Plesiogulo , and a species of Vishnuonyx . The latter possibly represents the last appearance of this genus in Africa. Torolutra ougandensis is known from both the Late Miocene and Early Pliocene deposits of the Middle Awash. The genus Sivaonyx is represented by at least two species: S. ekecaman and S . aff. S. soriae . Most of the lutrine genera documented in the Middle Awash Late Miocene/Early Pliocene are also documented in contemporaneous sites of eastern Africa. The new observations presented here show that mustelids were more diverse in the Middle Awash Late Miocene and Early Pliocene than previously documented.

The Woranso-Mille paleontological study area is located in the central Afar region of Ethiopia, about 360 km northeast of the capital, Addis Ababa. Some parts of this area have been paleontologically known since the 1970s. However, most of the fossiliferous areas were discovered by surveys conducted in the region between 2002 and 2004. By the end of the 2006 field season, a total of 17 vertebrate localities had been designated, and more than 1000 vertebrate fossil specimens collected. Among these specimens, there are more than 20 fossil hominid craniodental and postcranial remains, including one partial skeleton, of Pliocene age (3.5–3.8 Ma). Research at this study area has thus far focused on the geology and paleontology of the early Pliocene deposits along the Mille river and adjacent areas located between the towns of Mille and Chifra. Exposed sediments in the new fossiliferous area are mostly silty sand and silty clay horizons interbedded with a number of volcanic tuffs and basaltic flows suitable for 40Ar/39Ar radioisotopic dating. The total stratigraphic section is about 50 m thick with a minimum age of ~3.5 Ma. The study area also has deposits of early to middle Pleistocene age, although no locality has been designated to date. The new Woranso-Mille paleontological study area provides a crucial temporal window into the time during which Australopithecus anamensis (3.9–4.2 Ma) appears to have given rise to Australopithecus afarensis (3.0–3.6 Ma). Radiometric dates have thus far yielded a minimum age of 3.5 Ma for the hominid localities and this conforms well with a biochronological age estimate of 3.6–3.8 Ma. The associated fauna, particularly the abundance of fossil cercopithecids and presence of diverse aquatic fossil vertebrates, indicates a relatively closed, wooded habitat probably associated with a paleo-Mille river.

Ribs are naturally fragile and, as a consequence, are rarely preserved in the fossil record. The costal elements recovered from the KSD-VP-1/1 partial skeleton are important evidence allowing reconstruction of the hominin thoracic cage. The ribs of KSD-VP-1/1 are examined with respect to their implications for the evolution of Australopithecus afarensis thoracic morphology. Angulation and torsion along the rib corpus and rib declination indicate a broad upper thorax and a deeply invaginated thoracic vertebral column. Implications for the early hominin thoracic bauplan are discussed.

KSD-VP-1/1 is a 3.6 million years old (Ma) partial skeleton of Australopithecus afarensis recently discovered from the Woranso-Mille study area in the Afar region of Ethiopia. The recovered elements of this specimen, which include cervical vertebrae, a complete scapula, clavicle, numerous ribs, pelvis, and elements of the fore- and hindlimbs, greatly enhance our understanding of the paleobiology of early Australopithecus afarensis. Detailed analyses of the cervical vertebrae indicate that Australopithecus afarensis had a highly mobile neck, signaling human-like kinematics consistent with habitual upright posture and bipedalism. Elements of the shoulder girdle exhibit some primitive morphology but are overall more similar to humans than has been previously understood. This similarity is inconsistent with the notion that the Australopithecus afarensis shoulder retained primitive morphology from an African ape-like chimpanzee/human last common ancestor. Morphology of the thorax also indicates that while some individual traits may appear to superficially suggest arboreality, Australopithecus afarensis did not have an abundance of functionally significant morphological traits that would suggest high canopy arboreality as found today in large-bodied apes. Most of the inconsistencies in interpretations of early hominin paleobiology appear to stem from methodological differences, incorrect a priori assumptions, or incomplete information derived from fragmentary specimens.

Early hominin partial skeletons are extremely rare in the fossil record, particularly for the time predating 3 Ma (million years ago). As a result, most of our understanding of Pliocene hominin paleobiology and phylogenetic relationships is based on isolated and fragmentary specimens. The Middle Pliocene species Australopithecus afarensis is one of the best-known early hominin species, and yet only three adult partial skeletons of the species have been recovered thus far, including the one described in this volume. The best known of these is the 3.2 Ma partial skeleton of a small female (A.L. 288-1 ) from Hadar, Ethiopia, and much of our understanding of the paleobiology of this species has been influenced by this specimen. The newly recovered 3.6 Ma partial skeleton from the Woranso-Mille, Ethiopia (KSD-VP-1/1, aka Kadanuumuu ) represents a much larger male individual and may provide us with fresh insights into the paleobiology of Au. afarensis . This specimen not only preserves elements of the forelimb and hindlimb, but also includes complete elements, such as the scapula and clavicle, which were previously known only from fragmentary specimens. This edited volume provides the taphonomy and paleoecology of the partial skeleton, as well as detailed comparative descriptions of the preserved elements of KSD-VP-1/1 and their implications for our understanding of early hominin paleobiology. This chapter will present a basic introduction to the discovery of KSD-VP-1/1 and provide a guide to the contents of the volume.

The pelvis and limb bones of KSD-VP-1/1 are analyzed with respect to their implications for the history of morphology, gait pattern, and bauplan of Australopithecus afarensis . The pelvis is essentially of modern aspect, and differs little from that of modern humans save in a few minor respects, such as, for example, size of the auricular surface and acetabulum. Its ischial conformation is the direct evidence of a strong history of dynamic running in its immediate ancestors, and its general shape confirms the overall Australopithecus pattern of platypelloidy. The implications with respect to the history of the hominin bauplan are discussed, as are those impacted by recent developments in our understanding of developmental biology. These are reviewed in light of other recent analyses of early hominin morphology and locomotion.

KSD-VP-1/1, a partial skeleton of Australopithecus afarensis , was excavated from Pliocene strata at Korsi Dora, 3.3 km southeast of the confluence of the Waki and Mille rivers in the northwestern part of the Woranso-Mille paleoanthropological research site. A tuff collected from ~2.7 m below the fossil horizon, at the bottom of a trench dug 25 m to the east of the fossil excavation, yielded an 40Ar/39Ar age of 3.60 ± 0.03 Ma for anorthoclase feldspar. Strata in the trench and the fossil excavation site comprise a single normal magnetozone interpreted as part of the normal subchron C2An.3n, immediately above the Gauss/Gilbert paleomagnetic transition. Geologic mapping and tephrochemical analyses combined with paleomagnetic data place the fossil horizon and the trench section into local and regional stratigraphic context by constraining the partial skeleton to be younger than the Kilaytoli tuff (KT), a ~4 m thick vitric ash with an anorthoclase feldspar age of 3.570 ± 0.014 Ma. This unit is widely recognized at Korsi Dora, in collection areas north of the Waki-Mille confluence and outside the field area. The KT correlates with the Lokochot Tuff of the Omo-Turkana Basin in Kenya. Sedimentological features of the mudstone and sandstone in and near the excavation site are consistent with deposition in a floodplain or floodplain lake proximal to a stream channel.

Tephra geochemistry and (40)Ar/(39)Ar geochronology are reported for the Waki-Mille area in the northwestern part of the Woranso-Mille paleoanthropological project area in the west central Afar region of Ethiopia. Previous studies documented dentognathic fossils that are morphologically intermediate between Australopithecus anamensis and Australopithecus afarensis and some that are attributed to Australopithecus afarensis. Additional dentognathic remains from the study area were assigned to the newly identified species Australopithecus deyiremeda. These fossil hominin taxa were recovered from volcanic and sedimentary strata containing tuffs ranging in age from more than 3.77 million years ago (Ma) to less than 3.469 Ma. One of the tuffs was correlated based on geochemistry, feldspar mineralogy, and age to the Lokochot Tuff of the Omo-Turkana Basin of southern Ethiopia and Kenya. Variations in major and minor element abundances in volcanic glass demarcate ten geochemically distinct tuffs and tuff sequences, including three that are geochemically similar to widespread regional tuffs, specifically the Lomogol, Lokochot, and β- Tulu Bor/Sidi Hakoma tuffs. A new (40)Ar/(39)Ar age for the Waki Tuff, which is geochemically similar to the Lomogol Tuff, is 3.664 ± 0.016 Ma. Other tuffs in the Waki-Mille area are geochemically dissimilar to regional tuffs documented to date. Identification of tuffs based on character, stratigraphic position, and geochemistry refines local stratigraphic correlations and delineates the geographic distributions of precisely dated fossiliferous levels within the Waki-Mille area.

(2006). A new antelope, Zephyreduncinus oundagaisus (Reduncini, Artiodactyla, Bovidae), from the late Miocene of the Middle Awash, Afar Rift, Ethiopia. Journal of Vertebrate Paleontology: Vol. 26, No. 1, pp. 213-218.

The Middle Awash paleontological study area, located in the Afar Rift of Ethiopia, has yielded fossils spanning the last six million years. The geology and geochronology of the Mio-Pliocene sites of the study area have been refined and a reliable chronostratigraphy has been established by 40 Ar/ 39 Ar radiometric dating. The latest Miocene Adu-Asa Formation is divided into four members distinguished from each other by silicic and basaltic tuff marker horizons, most of which are dated basaltic tuffs. Radiometric dating has constrained the age of the Adu-Asa Formation to between 5.2-5.8 Ma. These dates are also supported by paleomagnetic results and biochronology. More than 2,000 fossil specimens were collected from the Adu-Asa Formation between 1992 and 2000. These fossils document 64 mammalian species belonging to 32 genera, 23 families, and 8 orders. This assemblage includes a number of new taxa. Included in the assemblage are First and Last Appearance Datums (FADs and LADs) of some groups, including the earliest record of the hominid genus Ardipithecus . Most of the taxa indicate a predominance of mesic and wooded habitat during the deposition of the Adu-Asa Formation. In these deposits, colobines, viverrids, mustelids, bovines, boselaphines, and tragelaphines are abundant, whereas alcelaphines are absent. Quantitative analyses of biogeographic relationships of the Middle Awash Late Miocene (MALM) mammalian fauna indicate stronger relationships with other African sites than with faunas from Eurasian sites. The MALM deposits have generated a critical dataset for analytic work on past environments, biogeographic relationships, and African vertebrate evolution. Moreover, the geographic position of the Middle Awash, coupled with its precise calibration and chronological span, make it a key section for interpreting latest Miocene faunal interchanges between Africa and Eurasia.

The Upper Pleistocene localities of Aduma and Bouri have yielded hominid fossils and exten- sive Middle Stone Age (MSA) archaeological assemblages. The vertebrate fossils recovered include parts of four hom- inid crania from Aduma and a complete right parietal from Bouri. Archaeological associations and radiometric techniques suggest an Upper Pleistocene age for these hominids. The more complete cranium from Aduma (ADU-VP-1/3) comprises most of the parietals, the occipi- tal, and part of the frontal. This cranium is compared to late Middle and Upper Pleistocene hominid crania from Africa and the Middle East. The Aduma cranium shows a mosaic of cranial features shared with "premodern" and anatomically modern Homo sapiens. However, the poste- rior and lateral cranial dimensions, and most of its anat- omy, are centered among modern humans and resemble specimens from Omo, Skhul, and Qafzeh. As a result, the Aduma and Bouri Upper Pleistocene hominids are as- signed to anatomically modern Homo sapiens. Am J Phys

Molecular studies suggest that the lineages leading to humans and chimpanzees diverged approximately 6.5–5.5 million years (Myr) ago, in the Late Miocene1,2,3. Hominid fossils from this interval, however, are fragmentary and of uncertain phylogenetic status, age, or both4,5,6. Here I report new hominid specimens from the Middle Awash area of Ethiopia that date to 5.2–5.8 Myr and are associated with a wooded palaeoenvironment7. These Late Miocene fossils are assigned to the hominid genus Ardipithecus and represent the earliest definitive evidence of the hominid clade. Derived dental characters are shared exclusively with all younger hominids. This indicates that the fossils probably represent a hominid taxon that postdated the divergence of lineages leading to modern chimpanzees and humans. However, the persistence of primitive dental and postcranial characters in these new fossils indicates that Ardipithecus was phylogenetically close to the common ancestor of chimpanzees and humans. These new findings raise additional questions about the claimed hominid status of Orrorin tugenensis8, recently described from Kenya and dated to ∼6 Myr9.

The Middle Awash study area of Ethiopia's Afar rift has yielded abundant vertebrate fossils (approximately 10,000), including several hominid taxa. The study area contains a long sedimentary record spanning Late Miocene (5.3-11.2 Myr ago) to Holocene times. Exposed in a unique tectonic and volcanic transition zone between the main Ethiopian rift (MER) and the Afar rift, sediments along the western Afar rift margin in the Middle Awash provide a unique window on the Late Miocene of Ethiopia. These deposits have now yielded the earliest hominids, described in an accompanying paper and dated here to between 5.54 and 5.77 Myr. These geological and palaeobiological data from the Middle Awash provide fresh perspectives on hominid origins and early evolution. Here we show that these earliest hominids derive from relatively wet and wooded environments that were modulated by tectonic, volcanic, climatic and geomorphic processes. A similar wooded habitat also has been suggested for the 6.0 Myr hominoid fossils recently recovered from Lukeino, Kenya. These findings require fundamental reassessment of models that invoke a significant role for global climatic change and/or savannah habitat in the origin of hominids.

The Woranso-Mille study area has thus far yielded more than 120 early hominin fossil specimens dated to between 3.4 and 3.8 million years ago. Previous studies indicate that dentognathic fossil remains from the study area show a mosaic of features shared by both Australopithecus anamensis and Australopithecus afarensis. Here, we describe 12 isolated mandibular fourth premolars recovered from the Woranso-Mille study area and compare them with those of other early hominins using both traditional comparative methods and geometric morphometric methods. The results indicate that the Woranso-Mille sample is most similar to Au. afarensis among hominins. However, some specimens show distinctive features of crown shape, namely an extremely bulging distolingual corner associated with a relatively large talonid and a mesiodistally elongated crown. This unique morphology is accompanied by a root form that is different from those identified previously. The existence of distinctive P4 morphology is intriguing given the presence of more than one hominin species at Woranso-Mille, although support for taxonomic heterogeneity in this sample is equivocal. Further, the taxonomic significance of these features is unclear, as they fail to distinguish Pan from Gorilla and known hominin species from one another.

A large series of fossil cercopithecids has been recovered from the hominid-bearing Woranso-Mille site, Afar State, northeastern Ethiopia. Here we report the taxonomy of those specimens from the Am-Ado, Aralee Issie, Korsi Dora, Makah Mera, and Mesgid Dora collection areas, which are all roughly contemporaneous and dated to between 3.6 and 3.8 million years ago. This series includes a minimum of two cercopithecine and three colobine species. Theropithecus oswaldi cf. darti is by far the most common species in the assemblage, making up over 90% of identifiable cercopithecid specimens. There is also at least one other species of papionin, which cannot be currently assigned to a genus. The colobines are here allocated to Cercopithecoides cf. meaveae and two other species, one small and one large, that cannot be currently assigned to genus. The T. oswaldi cf. darti series from Woranso-Mille is both the earliest and largest identified to date. It documents the earliest occurrence of the T. oswaldi lineage and strongly suggests that parallel evolution of molar morphology has occurred within the genus between T. oswaldi and Theropithecus brumpti . Given the dominance of monkeys at Woranso-Mille, and the preponderance of Theropithecus among cercopithecids, T. o. cf. darti is likely to be the most common mammal present at the 3.6–3.8 million-years-old localities of the Woranso-Mille study area. Some explanations for this unusual occurrence are explored, and implications for the paleoenvironment at Woranso-Mille are also discussed.

A newly discovered partial hominin foot skeleton from eastern Africa indicates the presence of more than one hominin locomotor adaptation at the beginning of the Late Pliocene epoch. Here we show that new pedal elements, dated to about 3.4 million years ago, belong to a species that does not match the contemporaneous Australopithecus afarensis in its morphology and inferred locomotor adaptations, but instead are more similar to the earlier Ardipithecus ramidus in possessing an opposable great toe. This not only indicates the presence of more than one hominin species at the beginning of the Late Pliocene of eastern Africa, but also indicates the persistence of a species with Ar. ramidus-like locomotor adaptation into the Late Pliocene.

Only one partial skeleton that includes both forelimb and hindlimb elements has been reported for Australopithecus afarensis . The diminutive size of this specimen (A.L. 288-1 ["Lucy"]) has hampered our understanding of the paleobiology of this species absent the potential impact of allometry. Here we describe a large-bodied (i.e., well within the range of living Homo ) specimen that, at 3.58 Ma, also substantially antedates A.L. 288–1. It provides fundamental evidence of limb proportions, thoracic form, and locomotor heritage in Australopithecus afarensis . Together, these characteristics further establish that bipedality in Australopithecus was highly evolved and that thoracic form differed substantially from that of either extant African ape.