Friday 25 December 2020

Fossils from the Pliocene Mille-Logya Site in the Afar Region of Ethiopia suggest that the origin of the genus Homo was associated with environmental change.

For many decades, a disparity between the resolution of long and continuous marine paleoclimate records versus fragmentary and time-averaged terrestrial records has hampered our ability to establish precise links between human evolution and major environmental changes. However, a recent proliferation of fieldwork, new drilling campaigns targeting highly detailed and relatively continuous paleolake records, and novel geochemical approaches are helping to assess terrestrial environmental dynamics at finer resolutions. Despite this progress, it remains the case that fauna, particularly hominins, are poorly sampled from the crucial time range between 3 and 2.5 million years ago because fossiliferous sediments dating to this interval are rare. Although prolific deposits of the Omo-Turkana Basin in Ethiopia and Kenya do contain sediments from this interval, the Hominin fossils are fragmentary and their taxonomic identities are uncertain. Sedimentary basins of the Afar in Ethiopia are highly fossiliferous, containing the most complete Hominin record of the past 6 million years, alongside diverse faunas and well-established chronologies, but the 3–2.5 million years ago interval is very poorly represented.

The Mille-Logya Project is a new palaeoanthropological site, dated from roughly 2.9 to 2.4 million years ago, at the northeast end of the well-known Plio-Pleistocene sites in the Awash Valley of the Afar Regional State, Ethiopia. Research at Mille-Logya started in 2012 and systematic geological and palaeontological surveys were conducted in 2014, 2015, 2016, 2018, and 2019.

In a paper published in the journal Nature Communications on 19 May 2020, Zeresenay Alemseged of the Department of Organismal Biology and Anatomy at the University of Chicago, Jonathan Wynn of the Division of Earth Sciences at the National Science Foundation, Denis Geraads of the Centre de Recherche en paléontologie at Sorbonne Universités, Denne Reed of the Department of Anthropology at the University of Texas at Austin, René Bobe of the Primate Models for Behavioural Evolution Lab at the University of Oxford, the Gorongosa National Park, and the Interdisciplinary Center for Archaeology and Evolution of Human Behavior at the Universidade do Algarve, Shannon McPherron of the Department of Human Evolution at the Max Planck Institute for Evolutionary Anthropology, Alan Deino of the Berkeley Geochronology Center, Mulugeta Alene of the School of Earth Sciences at Addis Ababa University, Mark Sier of the Consorcio del Centro Nacional de Investigación de La Evolución Humana, and the Department of Earth Sciences at the University of Oxford, Diana Roman of the Earth and Planets Laboratory at the Carnegie Institution for Science, and Joseph Mohan of the Climate Change Institute at the University of Maine, provide the first report on the geological and palaeontological content of this site.

Location of the Mille-Logya Project area within the greater Afar sedimentary basin, Afar, Ethiopia. Panel (a) shows the location of the Mille-Logya Project within Ethiopia; (b) the localtion of the Mille-Logya Project relative to other major Plio-Pleistocene Hominin sites in the Afar triangle; (c) the distribution of individual fossils and the three main collection zones (Gafura, Seraitu, and Uraitele). Alemseged et al. (2020).

The fossiliferous sediments exposed at Mille-Logya are generally younger than most of other known research areas in the region. Thus, they offer a unique opportunity to elucidate major events in Human evolution including the transition from Australopithecus to Homo, the emergence of Paranthropus, and the advent of manufactured stone tools. Furthermore, although Australopithecus afarensis is the most abundantly preserved Hominin from the region between 3.8 and 2.9 million years ago, its fate is largely unknown because of a regional hiatus in the sedimentary record of the Hadar Basin between 2.9 and 2.7 million years ago. Above this unconformity, the sediments of the Busidima Formation fill a local half graben near the western escarpment of the Ethiopian Rift. Compared to the largely lacustrine and peri-lacustrine Hadar Formation, the Busidima Formation exhibits a very different style of deposition, characterised by low sedimentation rates in almost exclusively episodic, high-energy fluvial settings, resulting in relatively poor fossil preservation. Meanwhile, during this same interval (2.9–2.7 million years ago), lacustrine and peri-lacustrine sediments were deposited at Mille-Logya and continue into younger horizons.

Alemseged et al.'s geological work offers new evidence for the northeast migration of the Hadar Basin, expanding our knowledge of the history of the basin substantially. Three new fossiliferous horizons with differing faunal composition have been identified suggesting in situ faunal change. While the fauna in the older unit is comparable to that at Hadar and Dikika, the younger units contain species that indicate more open conditions along with remains of Homo. New data from Mille-Logya reveal how Hominins and other fauna responded to environmental changes during this key period. Alemseged et al.'s results show a connection between geotectonics, sedimentary basin migration and an in situ faunal change. Alemseged et al. also provide new evidence that could potentially explain what happened to Australopithecus afarensis after 2.9 million years ago and what caused the dispersal to or the emergence of Homo in the region.

Stratigraphic surveys of the Mille-Logya area were geared toward broadening our understanding of the geological history of the region. To achieve this, it is crucial not only to establish chronological relationships between fossiliferous sites, but to investigate stratigraphic, structural and facies relationships between discontinuous exposures of sedimentary basins. Early geological maps of the region showed isolated Plio-Pleistocene sediments within the new site, amidst basalt flows attributed to the Afar Stratoid Series. In these broad-scale maps, the sediments were attributed to undifferentiated Quaternary strata or the White Series (Enkafala Beds; both mapping units, were also used to indicate outcrops of the Hadar Formation, the latter having been much more thoroughly scrutinised since initial fossil discoveries at the Hadar site). Sediments in areas nearby broadly bracket and partly overlap with the strata of the Hadar Formation. Alemseged et al.'s work has identified three new fossiliferous stratigraphic units expanding our knowledge of the geological history of the region and providing context for our faunal and basin analysis.

Sedimentary exposures in the Mille-Logya area provide access to generally disconnected sections of up to about 60m in total thickness. Between these discontinuous exposures, extensive colluvial cover of volcanic, boulder- to cobble-sized material obscures most outcrop. Furthermore, a number of postdepositional faults divide the exposures into disconnected fault-bounded blocks. Hence, Alemseged et al.'s stratigraphic interpretations of relationships between sections are presently based on widespread marker beds, chemical groupings of interfingered basalts and tephras, nine new Argon⁴⁰/Argon³⁹ dates, and several magnetostratigraphic sections. These observations are sufficient to describe the overall stratigraphy, and to divide the sedimentary strata into three main fossiliferous intervals each exposed at one of the three main areas: Gafura, Seraitu, and Uraitele. Alemseged et al. designate these as informal stratigraphic units, with the aim of formalising a regional lithostratigraphic terminology in future work, building on these presently informal terminologies.

Stratigraphic sections, sedimentary content volcanic markers, and relationships among the three fossiliferous zones (Gafura, Seraitu, and Uraitele) at the Mille-Logya Research area. Alemseged et al. (2020).

The lowest stratigraphic unit, Gafura, begins with a sequence of thick, columnar-jointed basalt flows with intra-flow residual paleosols developed on the basalts. The Gafura sediments are poorly exposed, but occur along the southwestern flank of Iki-Ilu Ridge, and are best represented by a section exposed at Sidiha Koma (section JGW15-1). The upper surface of the basalt flow at the base of this zone forms a broad low-lying surface, dissected by the Gafura River, extending into the base of the Daamé Valley. This sequence of basalts defines the Gafura Basalts-I and Gafura Basalts-II groups; it underlies the main sedimentary sequence and is thus stratigraphically distinct from the overlying flows represented as the Afar Stratoid Series. Within the lowermost exposures of the Gafura Basalts, a normal to reverse magnetostratigraphic reversal is recorded Given the age constraints of overlying strata, this reversal must be equal to or older than the age of the base of the Kaena Chron (3.127 million years).

The transition to overlying sediments of the Sidiha Koma area is marked by mudstones with ferruginised burrows interspersed with thin, poorly-sorted sands with a framework of basaltic lithic grains, occasionally containing abundant Gastropods, and some Bivalves. Near the top of Gafura sediments, additional basalt flows overlie the sediments locally, although these have not yet been attributed to one of the geochemically-defined groups. The fossiliferous sediments of Gafura underlie a widespread diatomaceous unit, the Iki-Ilu Diatomite, which can be mapped in regionally extensive exposures along the southwestern flank of Iki-Ilu Ridge, across its southern tip, and into the floor of the Seraitu Valley, making this a practical stratigraphic boundary. The Hinti Mageta Tuff (2.914 million years old), preserved within the Iki-Ilu Diatomite, provides an upper age limit to the Gafura sediments.

The middle stratigraphic unit is represented by the Seraitu lake beds, which often form bare, steep cliffs of largely mudstone outcrops representing lacustrine deposition. Alemseged et al. use the base of the Iki-Ilu Diatomite as the lower boundary of this zone, although most sections cannot be mapped in continuity within a measured stratigraphic distance to the diatomite. The upper boundary of this zone may not be defined by a single widespread marker but can be locally taken as the stratigraphically lowest basalt flow with chemical composition characteristic of the Uraitele-Garsele Dora Group, which may consist of several different flow units. The Uraitele-Garsele Dora Group is frequently accompanied by an overlying, distinctive and widespread air-fall tuff with well-preserved glass and small lapilli-sized pumices, the Goyana Tuff. Thus, the presence of one or both of these markers provides a working stratigraphic definition.

The sediments of the Seraitu lake beds are predominantly laminated clays which often contain abundant Ostracods, Gastropods and some Bivalves, as well as plant fragments and Fish remains. Diatoms in the Iki-Ilu Diatomite are somewhat recrystallised but identifiable to the genus Aulacoseira. Tephras are also numerous in the lake beds but characteristically thin (less than 30 cm), often air-fall occurrences, in which the primary glass is altered. Despite this, abundant feldspar crystal populations are preserved, providing two of the Argon⁴⁰/Argon³⁹ dates reported by Alemseged et al.. Besides the Hinti Mageta Tuff at the base of the Seraitu lake beds, two tuffs within the lake sediments provide precise ages: MLP14/SR-6 at 2.576 million years ago and MLP14/GOY-2 at 2.485 million years ago. In addition to the chronological information from these markers, two sections within the Seraitu lake beds zone record a magnetostratigraphic reversal which we interpret to be the Gauss/Matuyama, dated to 2.59 million years ago.

The third unit, Uraitele, includes limited sedimentary exposures in-between extensive and thick basalt flows of the Uraitele-Garsele Dora Group, Gafura Basalts-I, and Gafura Basalts-II groups, which outcrop in the Goyana and Uraitele areas. The sediments include some lenticular sandstones interpreted as fluvial channels, but are predominantly laminated mudstones with occasional Gastropod and Bivalve bearing sandstones formed on surfaces of the Uraitele-Garsele Dora Group basalt or within the mudstones. The upper boundary of the Uraitele zone is as yet undefined, as the section continues in a thick sequence of numerous basalt flows that extend into the ridges of the Magenta Mountains at the northern extent of the area (mapped as the Afar Stratoid Series). The uppermost flows form two chemically-distinct groups, termed the Gafura Basalts-I and Gafura Basalts-II groups.

As with the Gafura area, the sediments of the Uraitele area contain a number of reworked vitric tuffs generally lacking large feldspar populations, but having distinctive chemical composition with no known correlates from the Awash Valley. One of these vitric tephras, the Uraitele Tuff, has also produced populations of feldspars suitable for dating, and represents the most precise age of those presently analysed from the Mille-Logya area: 2.443 million years old.

Given the above stratigraphic sequence, Alemseged et al. make some important interpretations of the basin’s history. Prior to about 3 million years ago, there is no evidence of an active depositional basin in the Mille-Logya region. Thus, during the period characteristic of most active lacustrine sedimentation at Hadar and Dikika (about 3.6–3 million years old), the sequence of Gafura Basalts and residual palaeosols at Mille-Logya suggests subaerial emplacement of basaltic lavas followed by periods of non-deposition and pedogenesis. The local onset of active subsidence and sedimentation is marked by early onset of shoreline and shallow lacustrine deposits overlying the uppermost Gafura Basalts-II Group. The subsequent lacustrine sequence culminates in a deep, well-mixed lake represented by an Aulacoseira-dominated Diatom facies within the Iki-Ilu Diatomite and invertebrate fossil rich mudstones preserved throughout the Seraitu lake beds zone. Few terrestrial indicators are present except occasional coarser-grained facies suggestive of shorelines or brief subaerial exposure, where occasional rhizoliths are preserved and Vertebrate fossils are slightly more common. This lacustrine setting is persistent throughout the exposures and continues into the overlying Uraitele sediments, the lowermost of which are characterised by  gastropod-bearing shoreline facies developed on the surface of basalt flows. Ultimately, the lacustrine phase ends with thin intervening sediments between basalt flows of the Gafura Basalts Group, which have been associated with the fissural system of the axial rift.

An archaeological survey was conducted in conjunction with palaeontological reconnaissance. Archaeological visibility is extremely low due to the combination of a thick colluvial cover, relatively few exposed sections and the fact that most of the sedimentary deposits are lacustrine in origin. Nevertheless, all of the fossiliferous localities and their surroundings were examined on multiple occasions. In general, lithic artifacts are infrequent and scattered at very low density. The only exception comes from Seraitu Dida where slopes on two adjacent ridges with sediments above the Uraitele Tuff have numerous handaxes and Levallois cores and flakes made on a fairly consistent coarse-grained but good quality volcanic material. While artifact densities were relatively high in this area, no concentrations suggested a source for this material that has temporal constraint. Nevertheless, Alemseged et al. excavated three trenches at the crest of one of the ridges above the Uraitele Tuff. In one of these trenches, about one meter into a layer of gravel, they found a single Levallois flake. A maximum age is provided by the Uraitele Tuff (2.443 million years old) but the minimum age remains unconstrained. Archaeological exploration will continue.

The stratigraphic setting provides a framework to interpret fossil data recovered from the three units. Fossil concentrations at Mille-Logya are sparse and relatively difficult to locate. Yet, after four field seasons, the fossil collection currently includes 2287 specimens, of which 1835 were collected while the rest were observed and documented on site Fossil collections at the Mille-Logya Project followed a standardised protocol in order to minimize collection bias. The identifiable specimens comprise 62 Cercopithecidae, 4 Hominidae, 33 Proboscidea, 10 Camelidae, 165 Suidae, 135 Hippopotamidae, 36 Giraffidae, 944 Bovidae, 218 Equidae, 21 Rhinocerotidae, 20 Carnivora, 17 Birds, and some Rodents, Fish, Turtles, and Crocodiles.

Hominins were recovered from four different localities and are represented by a left and right proximal ulnae (MLP-786 & MLP-1617 respectively: 2.6–2.8 million years old; from two different localities, thus not from the same individual), a calvarium fragment (MLP-1469: 2.6–2.8 million years old) and a diagnostic and complete upper second molar crown (MLP-1549: 2.4–2.5 million years old). The molar, found in two pieces that refit cleanly, measures 14 mm and 12.5 mm buccolingually and mesiodistally, respectively, falling within the known range of Australopithecus afarensis as well as early Homo as represented by A.L. 666-1 from the younger horizons at Hadar dated at 2.33 million years old. The buccolingual and mesiodistal dimensions overlap with those of early Homo and are closest to the mean values. The occlusal outline, which is dominated by the two mesial cusps, is rhomboidal with the longest axis running from the distolingual to mesiobuccal corners. The distobuccal corner is truncated. The tooth is moderately worn with no cuspal dentine exposure. The lingual wear flattens the protocone and hypocone and polishes the lingual margin leading to a rather homogenised region of the lingual half of the tooth. In contrast, the paracone and metacone are not as worn and the buccal margin is still sharp. The distal marginal ridge and distal fovea are quite perceptible, but the mesial marginal ridge is largely worn down leaving only a hint of the mesial fovea. In occlusal view, running buccal to the protocone and distal to the paracone is a large buccal groove that dominates other grooves and is positioned mesiobuccal to a lingual groove that is much smaller.

Hominin remains from the MLP area. (a), (b) are left and right proximal ulnae (MLP-786 & MLP-1617, respectively from two different localities, thus not from the same individual); (c) calvarium fragment (MLP-1469); (d) a diagnostic and complete upper second molar crown (MLP-1549). Alemseged et al. (2020).

An asymmetric and rhomboidal occlusal outline of the upper second molar has been reported to characterise Homo erectus and Homo habilis but is rare in Australopithecus afarensis. The Mille-Logya Project M2 possesses these features but is buccolingually broad unlike Homo erectus. Based on size and average enamel thickness in addition to diagnostic occlusal features, Alemseged et al. attribute it to Homo sp. With an age of 2.4–2.5 million years, this molar represents one of the oldest specimens of this genus and expands the earliest Homo sample from the Afar, which currently includes only LD 350-1 from Ledi Geraru at 2.8 million years and A.L. 666-1 from Hadar at 2.33 million years.

MLP-1549 bucco-lingual breadth and mesio-distal length in mm compared to mean values for many Hominin species. Alemseged et al. (2020).

The calvarium fragment is probably from the parietal; the bone is relatively thin and different from what is seen in middle Pleistocene specimens such as Bodo. MLP-1617 and MLP-786 are fragmentary proximal ulnae mainly preserving the olecranon and the trochlear notch. The two specimens differ in size and degree of preservation. While MLP-786 is larger, MLP-1617 is better preserved, especially its trochlear notch where the maximum mediolateral breadth, including the radial notch, is about 28 mm. In MLP-786, most of the distal aspect of the trochlear notch including the radial notch is broken away. In both, various quadrants of the trochlear notch are mildly concave. Maximum radial notch dimensions are 13.4mm anteroposteriorly and 9.4 mm superoinferiorly in MLP-1617. In both specimens, the olecranon process is moderately projected proximally and slightly more pronounced in MLP-786, as is seen in other Hominins. The olecranon proximodistal height is 7.3mm and 8.6mm respectively and similar to that of A.L. 438-1 (Australopithecus afarensis). The trochlear keel is mild in MLP-1617 and not perceptible in MLP-786. Posteriorly, MLP-1617 and MLP-786 measure 15.3mm and 18.5 mm, respectively at the middle of the trochlear notch and proximally they measure 20.8mm and 23.6 mm. The two ulnae can readily be assigned to Hominini based on the extent of the olecranon process and the orientation and extent of the trochlear and radial notches. Many features and dimensions discussed by Michelle Drapeau to characterise most fossil Hominins are preserved (mainly in MLP-1617) but further discrimination is not possible and these bones are therefore attributed to Hominini indet.

Molar crown dimensions. Buccolingual and mesiodistal dimensions of MLP-1549 compared to values in Australopithecus anamensis, Australopithecus afarensis, Australopithecus africanus, Paranthropus boisei, Paranthropus robustus, Homo habilis, Homo erectus & A.L. 666-1(Homo sp. from Hadar dated to 2.33 million years). (a) Upper second molar bucco-lingual dimensions in mm; (b) upper second molar mesio-distal dimensions in mm. Alemseged et al. (2020).

Cercopithecines, especially Theropithecus, are abundant though mostly represented by fragmentary teeth and extremities of postcranials. The Mille-Logya Project Theropithecus is characterised by diagnostic high crowned, bilophodont molars with deeply incised notches and clefts. Based on limited existing material, molar size appears larger than in representatives of Theropithecus oswaldi from the older members of the Hadar Formation. Some teeth displaying similar features are smaller; we interpret them as females of the same species. There are also several teeth that do not display these diagnostic Theropithecus features, which Alemseged et al. assign to Cercopithecinae indet. pending recovery of more complete specimens.

Enamel fragments belonging to Deinotherium are frequently encountered, and a nearly complete skull was excavated at Uraitele. Dental elements and postcranials of Elephantidae are common as well, and some were collected; a complete M3 from Uraitele best matches Elephas recki shungurensis known from Omo Shungura Members C to F, but there are no clear boundaries between successive subspecies to allow more precise biochronological attribution.

A single mandibular fragment belongs to an Aardvark (Orycteropus).

The family Camelidae is extremely rare in the East African Plio-Pleistocene, and the discovery of ten specimens at Mille-Logya is remarkable. One of them is the only partial skull known from this part of Africa to date and detailed analysis of this skull has been published elsewhere.

Though remains of Hippopotamidae are common throughout the sequence, there is only one species; it has a hexaprotodont dentition, with a slightly smaller i2 than i1 and i3, thus resembling early forms of the ‘aff. Hippopotamus protamphibius'. It is clearly different from Hexaprotodon bruneti, from the Hata Member of the Bouri Formation at about 2.5 million years old, which has a very large i3. Yet, formal identification must await the revision of the Turkana and Hadar material. No Tetraprotodont dentition has been recovered from the Mille-Logya Project area.

Notochoerus is the most common genus of Suid. Several complete third molars, mostly from the Gafura unit, are smaller than those of Notochoerus scotti from Omo C and later members, and match those of Notochoerus euilus from Omo B or the Hadar Formation. The morphology and mesiodistal length of their molars are similar to those in Notochoerus clarki, which coexists with Notochoerus scotti at Omo, but are broader and Alemseged et al. attribute them to Notochoerus euilus. Kolpochoerus is less common and all specimens are of comparable size. Dimensions of the third molars, mostly from the Seraitu lake beds, are close to the upper end of the range for Kolpochoerus afarensis from the Hadar Formation, or to the lower end of the range for Kolpochoerus limnetes from Shungura D-E, and are closest to specimens from Shungura B and C. They are also somewhat larger than those of Kolpochoerus philippi, from Matabaietu at about 2.5 million years old. There is no definite evidence of Nyanzachoerus, nor of Metridiochoerus.

Both Giraffa and Sivatherium are represented, but are rare. The relatively more common Giraffa is not particularly large, and a second, smaller species (Giraffa cf. gracilis) is also present.

Bovids are by far the most common Mammals, and almost half of the identifiable specimens, mostly represented by teeth, belong to Alcelaphini. They are followed in decreasing order of abundance by the Reduncini, Bovini, Aepycerotini, Antilopini, and Tragelaphini. Horn-cores are encountered relatively frequently, but are seldom associated with other cranial parts. An Alcelaphin that resembles Damaliscus ademassui from Gamedah dated to about 2.5 million years and perhaps a primitive Wildebeest (Connochaetes sp.) are present at Uraitele. Alemseged et al. assign the most common Alcelaphin to Damalborea, a genus that is present throughout the Hadar Formation. Although variation at Hadar is great, the Mille-Logya Project form is distinctive in its short, twisted horn-cores with homonymous torsion. It could be an evolved form of Damalborea grayi from the Denen Dora Member at Hadar. An unidentified, very small Alcelaphin is reminiscent of the one that first appears in the Kada Hadar 2 submember. There are at least two species of Reduncins. The less common one is probably Kobus sigmoidalis, best known from the Turkana basin and recently reported from Ledi-Geraru. he specific identity of the more common reduncin is not clear; it resembles Kobus oricornus from Omo Shungura, West Turkana, Koobi Fora, and Hadar. Surprisingly, this taxon is absent in the nearby Ledi-Geraru. A Bovini horn-core from the middle part of the section is comparable to the type of Pelorovis kaisensis, from Kaiso village in Uganda, about 2.5 million years old. Aepyceros is common but species identification is difficult due to incompleteness. One of the horncores is larger than those from the Kada Hadar Member. Gazella, the only Antilopin so far recovered, is represented by several long and slender horn-cores resembling Gazella harmonae from the Kada Hadar Member at Hadar, and probably Omo Shungura Member F and Olduvai Bed I. Most Tragelaphin horn-cores are from the younger part of the exposures and resemble Tragelaphus nakuae in their moderate torsion and in the presence of a low supraoccipital ridge of braincase. A very long horn-core is reminiscent of a specimen from Omo-160 in Shungura Member C. Overall, the Tragelaphin material suggests an age of 2.6–2.3 mullion years. A second and rare species from the middle part of the sequence is similar to Tragelaphus gaudryi of which an ancestral form appears in Omo Shungura Member C.

Though Rhinos are rare at the Mille-Logya Project, both the grazer Ceratotherium and the browser (or mixed-feeder) Diceros are encountered.

Equids are fragmentary but quite common. Alemseged et al. tentatively attribute all remains to a single species of Hipparion. A complete set of upper incisors shows that the I3 is not reduced, and the lingual grooves are shallow, in contrast to what is seen in derived hipparions of the cornelianus group. A remarkable feature is the absence or poor development of the ectostylid on many lower teeth. A moderately worn and well-preserved set of molars shows no ectostylids at all. Postcranial dimensions are close to the lower end of the Hadar range where skulls show that at least two species are represented.

Some postcranials potentially representing multiple taxa belong to Hyaenidae. Two metapodials and a tooth belong to a Felid, cf. Dinofelis. In addition, a distal radius belongs to a Servalsised Felid.

Antoine Louchart identified a large Ostrich and a member of the Anatidae, perhaps Sarkidiornis melanotos or Plectropterus gambensis. Large Ostriches have been mentioned from a number of Pleistocene Old World sites; they are likely attributable to Struthio asiaticus.

Crocodile teeth are widespread, and a few specimens represent Euthecodon.

Kathlyn Stewart identified Bagrid and Clariid Fish.

The Hadar Formation fauna, documented in the nearby sites of Hadar, Dikika, and Ledi-Geraru, has been widely studied and offers a very good reference for the new material from Mille-Logya. The Mille-Logya fauna points to a generally younger, late Pliocene age but shares a number of taxa with those from the Hadar Formation, where many have a wide chronological range. Of biochronological significance are the Antelopes, Damalborea and Kobus cf. oricornus. Gazella harmonae is also shared with Hadar, although this species has wide chronological and geographic ranges. Another indicator of a similar age is the Hexaprotodont Hippopotamid, present in the middle part of the sequence at Mille-Logya. The Suids also fall largely within the size range seen in the Hadar Formation, but most diagnostic specimens are encountered in the lower and middle parts of the sequence. It should be noted, however, that Kolpochoerus from the Seraitu lake beds is more consistent with the older absolute ages of this unit than with the younger ones. The absence of Nyanzachoerus suggests that the Mille-Logya Project assemblage postdates most of the fauna from Hadar. Although the above is generally true, there are differences within the Mille-Logya fauna indicating that sites in the southern portion of the research area are older than those in the north.

As in most African Pliocene sites, Bovids are the most common group followed by Equids and Suids. Primates are fairly common but rare compared to those at Hadar and Dikika. One striking feature of the Mille-Logya fauna is the high prevalence of Equids, particularly relative to Suids. At Hadar and Dikika the reverse is consistently the case. While this is true when the whole Mille-Logya assemblage is considered as a unit, looking at the different horizons reveals a different pattern. Gafura (roughly 2.9 to 2.8 million years old) contains fauna that is similar to that of Hadar where the proportion of Bovids compared to Suids and Equids is not very high and also where Suids are more common than Equids. In contrast, the Seraitu lake beds (roughly 2.8 to 2.6 million years old) and Uraitele zone (roughly 2.5 to 2.4 million years old) contain more Bovids while Equids overtake Suids. This suggest that the older fauna in Gafura might have followed the migration of the Hadar Lake Basin northeast around 3 million years ago, resulting in faunal similarities with Hadar. Relative faunal abundance however seems to have been altered in the younger horizons of the Seraitu lake beds and Uraitele (after abou 2.9 million years ago), leading to a faunal assemblage indicative of more open conditions. This is supported by the overall abundance of Bovids, particularly Alcelaphins, and Equids probably indicating an in situ faunal turnover.

Alemseged et al. used all Mille-Logya Project specimens identifiable to genus to compute the Sørenson (also known as Dice) faunal dissimilarity index for each pairwise comparison among the faunal zones. The results indicate that Seraitu and Uraitele are more compositionally similar to one another at the genus level than either of these zones is to Gafura. Alemseged et al. further conducted a correspondence analysis on taxon abundances in order to compare the Mille-Logya Project faunal zones with assemblages from the Hadar Formation at Hadar and Dikika. They restricted their analysis to seven bovid tribes (Aepycerotini, Alcelaphini, Antilopini, Bovini, Hippotragini, Reduncini, Tragelaphini), the Suid genera Notochoerus and Kolpochoerus, and all Equidae identified only to family. These relatively broad taxonomic categories were chosen to reduce the influence of inter-observer variation in taxonomic identifications. The correspondence analysis demonstrates that the Gafura assemblage is distinct from the Seraitu and Uraitele assemblages, with Gafura showing a high abundance of Notochoerus, and the Seraitu and Uraitele assemblages showing a high abundance of Alcelaphini and Antilopini, which are open-habitat indicator taxa.

Correspondence analysis on taxon abundances comparing the Mille-Logya Project faunal zones with assemblages from the Hadar Formation at Hadar and Dikika. Results demonstrate that the Gafura assemblage is distinct from the Seraitu and Uraitele assemblages, with Gafura showing a high abundance of Notochoerus, and the Seraitu and Uraitele assemblages showing a high abundance of Alcelaphini and Antilopini, which are open-habitat indicator taxa. Alemseged et al. (2020).

It is possible that some of the observed taxonomic differences between the collecting areas of successive ages are due to small sample size, but some have clear biochronological significance. Alemseged et al. interpret the presence of Damaliscus cf. ademassui, Connochaetes sp., and perhaps Kobus sigmoidalis, at Uraitele to suggest that the latter is younger than Gafura and Seraitu. In addition, it is not yet certain if the Tragelaphus from Gafura is a 'typical' Tragelaphus nakuae, and whether Damalborea survived into the Uraitele horizon.

In sum, the overall similarity of the Mille-Logya fauna, especially from the older Gafura sites, with that from the Hadar Formation is remarkable. Also, in spite of the apparent younger age, it contrasts with younger sites in the Middle and Lower Awash. For instance, the Bovid assemblage of Bouri Hata at about 2.5 million years before present includes a long list of Bovid taxa for which there is no evidence at Mille-Logya: Beatragus, cf. Numidocapra, cf. Rabaticeras, Megalotragus, Hippotragus, Oryx, and Tragelaphus strepsiceros. The nearby Ledi-Geraru area contains sediments whose ages are very similar to those of Mille-Logya, but their faunal assemblage also looks different, containing Beatragus, Syncerus and Ugandax, but lacking Kobus oricornus, the most common Reduncini at Mille-Logya.

In regards to palaeoenvironments, Tragelaphins are rare as are Giraffes, while hipparions and reduncins are common. Alcelaphins are by far the most abundant Bovids. The relative abundance of the otherwise rare Camelus is also noteworthy. On the whole, this assemblage points to an open savanna or grassland with little woody cover. This in conjunction with the presence of Homo at Mille-Logya may suggest that the earliest members of Homo were associated with more open environments than Australopithecus was. The in situ faunal change at Mille-Logya may be linked to environmental and climatic factors that may have also caused Homo to emerge in or disperse to the region. Further field work and faunal analysis with better taxonomic resolution and use of additional proxies will help to better elucidate the palaeoenvironmental and palaeoecological conditions of this new site and its relevance to the understanding our origins.

Although the Afar Depression has contributed uniquely to our understanding of the biological and cultural evolution of Hominins and faunal evolution more broadly over the past 6 million years, palaeontological data have been sparse in the region from the stratigraphic time interval (2.9–2.4 million years ago) represented by the Mille-Logya sediments. Alemseged et al.'s work shows the unique nature of the faunal assemblage and composition at the new site. The results suggest a northeast migration of the Hadar Basin and the creation of a new depocenter at Mille-Logya with continuity of the lake deposits and shoreline sediments that preserve a fauna similar to that from Hadar. Furthermore, Alemseged et al. have identified three different fossiliferous units in this project area suggesting an in situ faunal change. Yet, relative to the Hadar Formation fauna, which is older than 3 million years, Mille-Logya has a large proportion of Alcelaphin Bovids and Equids, indicating that the area likely included more open habitats after 3 million years ago. The absence of early Metridiochoerus, Menelikia and Australopithecus from Mille-Logya suggests that these habitats may not have been suitable for these taxa. The presence of Homo is instead suggestive of an adaptive shift in the transition between Australopithecus and Homo to settings with overall drier and more open conditions.

Looking forward, more research at Mille-Logya Projest will allow better documentation of the geological setting and palaeontological content of the poorly known post-Hadar-Dikika periods to better articulate the environmental setting of Human evolution. For the first time, Alemseged et al. now have a better understanding of the cause for local cessation of sedimentation in the Hadar-Dikika areas after approximately 2.9 million years ago. The new data set will serve as the basis for within-site faunal comparison and for testing competing spatial and temporal faunal and environmental change hypotheses. The presence of Hominin remains, including Homo associated with a diverse fauna presents a unique opportunity to address key questions that pertain to our genus and factors that led to its emergence and subsequent biological and cultural evolution.

See also...

Online courses in Palaeontology. 

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