Hadar, Ethiopia
"Lucy", a 3.2-million-year-old Australopithecus afarensis fossil discovered in Hadar, Ethiopia | |
Location | Mille district, Afar Region, ![]() |
---|---|
Region | Eastern Africa |
Coordinates | 11°10′01″N 40°37′59″E / 11.167°N 40.633°E |
Type | Archaeological Paleontology |
History | |
Periods | Middle - Late Pliocene |
Site notes | |
Excavation dates | 1970, 1972, 1973, 1974, 2000 |
Archaeologists | Donald Johanson Maurice Taieb Denis Geraads Zeresenay Alemseged |
Condition | Excavated |
Ownership | Ethiopian Government |
Hadar or Hadar Formation (also spelled Qad daqar, Qadaqar; Afar "white [qidi] stream [daqar]")[1] is a paleontological fossil site located in Mille district, Administrative Zone 1 of the Afar Region, Ethiopia, 15 km upstream (west) of the A1 road's bridge across the Awash River (Adayitu kebele).[2]
It is situated on the southern edge of the Afar Triangle (part of East Africa's Great Rift Valley), along the left banks of the Awash River, between two minor tributaries, the eponymous Kada Hadar and the Kada Gona.[2] In 1972, Taieb organized a small exploratory reconnaissance of the Afar region to investigate more paleontological finds there. After six weeks of exploration, the party focused on the Hadar site.[3]
The site has yielded some of the most well-known hominin fossils, including "Lucy". These hominin fossils range in age from approximately 3.42 to 2.90 million years ago. These finds give us a greater understanding of hominin evolution during this period.
It is postulated that the specimens in the region were deposited by way of a large river system with associated crevasse channels/splays, deltas, and distributary channels, as well as periodic transgressions of paleolake Hadar located east of the research area (Aronson and Taieb, 1981, Tiercelin, 1986, Campisano and Feibel, in press) possibly related to geological activity or climatic cycles in at least the Kada Hadar Member (Yemane et al., 1996, Yemane, 1997, Campisano and Feibel, in press)."
According to Jon Kalb, early maps show caravan routes passing within 10 to 15 km of Hadar but not through it. The British explorer L.M. Nesbitt passed 15 km west of Hadar in 1928.[1]
Geology
The region's rocks consist mainly of mudstones, siltstones, fine-grained sandstones and volcanic tuffs. The region of Hadar has been divided into four geologic members — Basal (~3.8–3.42 Ma), Sidi Hakoma (~3.42–3.26 Ma), Denen Dora (~3.26–3.2 Ma), and Kada Hadar (<~3.2 Ma)—with three tuffs (Sidi Hakoma Tuff [SHT], Triple Tuff [TT] and Kada Hadar Tuff [KHT]) separating the four members.
The Sidi Hakoma member tends towards high rainfall and low seasonality. The overlying Denan Dora Member was a grassland habitat. Finally, the Kada Hadar Member was an even more open and arid habitat, as seen in the high abundance of antilopines, which frequent these types of terrains.[3]
Paleontology
The first paleo-geological explorations of the Hadar area were conducted by Maurice Taieb. He found Hadar in December 1970 by following the Ledi River, which originates in the highlands north of Bati to empty into the Awash River. Taieb recovered several fossils in the area and led a party back to Hadar in May 1972. In October 1973, 16 individuals with the International Afar Research Expedition (IARE) arrived at Hadar and camped there for two months during which the first hominin fossil was found. (Taieb claims in his 1985 book Sur la Terre des premiers Hommes to have discovered the Hadar fields in 1968, but Kalb argues that claim to be incorrect.)[4] The IARE party examined a series of sedimentary layers called the Hadar Formation, which was dated to the late Pliocene to early Pleistocene epochs (3.5 to 2.3 million years ago).[5]
The anthropologist Donald Johanson, a member of the 1973 expedition to Hadar, returned the next year and discovered the fossil hominin "Lucy" in the late fall of 1974.[6] He spotted a right proximal ulna in a gully, followed by an occipital bone, a femur, some ribs, a pelvis, and a lower jaw. Within two weeks, nearly 40% of the hominoid skeleton had been identified and cataloged.[7] Lucy is the most famous fossil to have been found at Hadar. Lucy is among the oldest hominin fossils ever discovered[6] and was later given the taxonomic classification Australopithecus afarensis. (The name 'Lucy' was inspired by the song "Lucy in the Sky with Diamonds" by The Beatles, which happened to be playing on the radio at base camp.)
In 1975, Donald Johanson made another discovery at a nearby site in Hadar: 216 specimens from approximately 17 individuals, most likely related and varying in age, called AL 333 (colloquially referred to as the "First Family").
About thirty years later in nearby Dikika, another Australopithecus afarensis fossil skeleton was found in a separate outcrop of the Hadar Formation across the Awash River from Hadar. The skeleton is of a three-year-old girl later named "Selam," which means peace in Amharic Ethiopian languages.
Specimens and inferences
![](http://upload.wikimedia.org/wikipedia/commons/thumb/3/3e/Hadar%2C_Ethiopia_%3B_Australopithecus_afarensis_1974_discovery_map.png/250px-Hadar%2C_Ethiopia_%3B_Australopithecus_afarensis_1974_discovery_map.png)
In 1973 and 1974 when the first anatomical discoveries were made, their size and shape pointed towards a variety of taxa, but further research has confirmed that only one hominin taxon is present here. The first find there was a fossil knee joint estimated to date from 3.4 million years ago. Since then, the Hadar research area has yielded 370 specimens of A. afarensis, one specimen of Homo, and 7571 additional vertebrate specimens.
The specimens recovered display a variety of different primitive cranial post features, which indicate A. afarensis is distinct from other species of Australopithecus: small cranial capacity, palate similar to African apes (parallel tooth rows, shallow, long from front to back, narrow from side to side), primitive occipital, basal cranium anatomy, high frequency of unicuspid third premolars, prognathic face, and primitive mandibular anatomy. Postcranially, the pelvis, knee, ankle, and foot indicate habitual, terrestrial bipedalism, but ape-like curved finger and foot bones are retained ancestral ape-like features.[3]
Fossil content
Color key
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Notes Uncertain or tentative taxa are in small text; |
Other specimens discovered from the Hadar Formation contain a vast diversity of bovid species found in Africa. The bovids found in the formation included the Aepycerotini (Aepyceros), Alcelaphini (Damalborea and Parmularius), Antilopini (Gazella), Bovini (Ugandax and Pelorovis[?]), Caprini (Budorcas), Cephalophini, Hippotragini (Oryx), Neotragini (Raphicerus[?] and Madoqua), Reduncini (Kobus), and Tragelaphini (Tragelaphus)[8].[9][10] Artiodactyls outside the bovid family were present within the formation as well, namely the giraffids (Giraffa and Sivatherium), Hippopotamidae (Hippopotamus), and suids (Kolpochoerus, Notochoerus, and Nyanzachoerus).[11] While a definitive list of carnivorans found within the Hadar Formation has yet to be compiled, confirmed genera that were found within the Hadar Formation include canids (Canis and Nyctereutes), felids (Dinofelis,[12] Leptailurus, Felis, Homotherium, and Panthera), hyaenids (Chasmaporthetes, Ikelohyaena, Crocuta, Hyaena, and cf. Pliocrocuta),[13] herpestids (Herpestes and cf. Helogale), mustelids (Mellivora, Enhydriodon,[14] and cf. Poecilogale), and the viverrid (cf. Civettictis). Mammals within the formation outside the artiodactyl and carnivoran families include a bat (indeterminate), the leporid (Lepus), the equid (Eurygnathohippus),[15] rhinoceroses (Ceratotherium and Diceros),[16] old world primates (Parapapio, Theropithecus, and Cercopithecoides), proboscideans (the deinothere Deinotherium and elephants Elephas, Loxodonta, and Mammuthus) [17][18][19] old world porcupines (Hystrix and Xenohystrix),[20] murid rodents (Gerbilliscus, Acomys, Golunda, Oenomys, Praomys, Saidomys, Millardia, and Mus),[21] the spalacid Tachyoryctes, a squirrel indet., and an aardvark species. Taxons within other classes are present within the Hadar Formation as well, such as birds (Plectropterus, Balearica, Anhinga, and Struthio) and reptiles (Crocodylus, Python, Varanus, and Bitis).[22][23][24][25][26][27] [28][29]
Mammals
Artiodactyla
Bovidae
Bovids of the Hadar Formation | |||||
---|---|---|---|---|---|
Taxa | Species | Presence | Material | Notes | Images |
Aepyceros | A. datoadeni | Kada Hadar Member at Hadar. | Near-complete skull lacking parts of horn cores(AL 787-1 KH1).[9] | Extinct relative of modern impala. | |
Beatragus | B. sp. | Detached braincase with two horn cores.[9] | Extinct relative of modern hirola. | ||
Budorcas | B. churcheri[30] | Sida Hakoma member.[9] | Complete skull with horns (AL 136-5 DD2)[9] | Extinct relative of takin. | |
Damalops | D. sidihakomai.[31] | Sidi Hakoma member. | AL 208–7, a skull with horn cores, right P4–M3, and left M2–3. | Extinct genus of Alcelaphinae. | |
Damalborea | D. elisabethae | Basal Member and Sidi Hakoma member.[9] | A skull with horn cores (AL 208-07 SH3).[9] | Extinct genus of Alcelaphine bovids. | |
D. grayi | Denen Dora and Kada Hadar Member. [9] | Multiple postcranial fragments such as horn cores, maxilla, and mandibles.[9] | |||
Gazella | G. harmonae | Kada Hadar Member at Hadar.[9] | Skull remains such as isolated teeth, maxilla, frontlet, and occipital (AL 444-16). [9] | Extinct relatives of modern gazelles | |
G. cf. janenschi | Denen Dora Member. | Horn cores (AL 302-6 DD).[9] | |||
G. cf. praethomsoni | Denen Dora and Kada Hadar Member | Few Horn core specimens.[9] | |||
Hippotragini | Praedamalis deturi | Kada Hadar, Sida Hakoma, and Denen Dora member at Hadar. | Multiple specimens based on horn cores and lower teeth.[9] | Ancestor of modern day oryx. | |
Kobus | K. hadarensis | Sidi Hakoma, Kada Hadar and Denen Dora Member. | Extinct relatives of antelopes. | ||
K. oricornus | Denen Dora and Kada Hadar Member. | Several specimens of braincase and horn core. [9] | |||
Madoqua | M. sp | ||||
Raphicerus | R. sp | Kada Hadar and Sidi Hakoma. | Mandibles and horn cores.[9] | ||
Parmularius | P. pachyceras | Sidi Hakoma and Kada Hadar Member.[9] | Braincases and horn cores.[9] | Large extinct alcelphines related to topis and hartebeests | |
Pelorovis | P. sp. | ||||
Tragelaphus | T. lockwoodi | Basal Member and Sidi Hakoma Member. | Braincase with complete horn cores (AL 142-3 SH2).[9] | Extinct relatives of spiral-horned antelopes. | |
T. rastafari | Basal, Sidi Hakoma, Denen Dora, and Kada Hadar Member. | Several horn core specimens. | |||
Ugandax | U. coryndonae | Denen Dora and Sidi Hakoma member at Hadar. | Complete partial skull and horns (AL 194-1 DD1)[9] | Ancestor of Cape buffalo | ![]() |
Giraffidae
Giraffids of the Hadar Formation | |||||
---|---|---|---|---|---|
Taxa | Species | Presence | Material | Notes | Images |
Giraffa | G. jumae | Sidi Hakoma member and Denen Dora Member. |
Partial teeth, vertebrae, ossicones, limb bones, and mandibles. [24] | Extinct relatives of modern giraffe. | |
G. stillei | Sidi Hakoma member and Denen Dora Member. |
Partial teeth, ossicones, limb bones, and mandibles. [24] | |||
Sivatherium | S. maurusium | Denen Dora Member, Kada Hadar Member, and Sidi Hakoma Member. | Partial mandibles and horns[24] | An okapi-like giraffid. |
Hippopotamidae
Hippopotamids of the Hadar Formation | |||||
---|---|---|---|---|---|
Taxa | Species | Presence | Material | Notes | Images |
Hexaprotodon | H. protoamphibius | Denen Dora Member. | Partial skull and teeth. | Extinct relatives of modern pygmy hippo. | |
H. coryndoni | Sidi Hakoma Member.[28] | Partial skull and teeth. | |||
Hippopotamus | H. afarensis | Basal, Sidi Hakoma, Kada Hadar, and Denen Dora Member.[28] | Complete postcranial skull and mandibles. | Extinct subspecies related to Common Hippopotamus, formally named Trilobophorus afarensis. |
Suidae
Suidaes of the Hadar Formation | |||||
---|---|---|---|---|---|
Taxa | Species | Presence | Material | Notes | Images |
Kolpochoerus | K. afarensis | Kada Hadar Member, Sidi Hakoma Member, and Denen Dora Member.[11] | Postcranial skulls, teeth fragments and mandibles. | ||
Notochoerus | N. eulius | Kada Hadar, Lower and Upper Sidi Hakoma Member, and Denen Dora Member.[11] | Postcranial skull, teeth, and mandibles. | ||
Nyanzachoerus | N. kanamensis | Basal member and Sidi Hakoma Member.[28] | Complete cranium, mandibles, and dentition. | Extinct genus of pig family. | |
N. pattersoni | Basal Member, Lower and Upper Sidi Hakoma Member, and Denen Dora Member.[11] | Complete skulls, mandibles, and dentition.[11] |
Carnivora
Canidae
Canids of the Hadar Formation | |||||
---|---|---|---|---|---|
Taxa | Species | Presence | Material | Notes | Images |
Nyctereutes | N. lockwoodi | Sidi Hakoma and Denen Dora Member at Dikika.[23] | Nearly complete skull (DIK-31-1).[32] | Extinct relative of modern Racoon dog. |
Felidae
Felids of the Hadar Formation | |||||
---|---|---|---|---|---|
Taxa | Species | Presence | Material | Notes | Images |
Dinofelis | D. aronoki | Kada Hadar Member[12] | Postcranial member: distal radius (AL-363-20)[12] | A sabertooth cat. | ![]() |
D. petteri | Sidi Hakoma Member and Denen Dora Member. [12] | Postcranial member: partial humerus (AL 168-30), proximal ulna (AL 262-9), and right astragalus (Al 173-11)[12] | A sabertooth cat. | ||
Felis | F. sp. | ||||
Homotherium | H. hadarensis | Denen Dora Member[23] | Complete cranium skull, mandible fragments (DIK-96-1), teeth, and forelimb.[23] | A sabertooth cat. | |
Megantereon | M. sp. | ||||
Leptailurus | L. sp. | ||||
Panthera | P. sp. |
Hyaenidae
Hyaenidaes of the Hadar Formation | |||||
---|---|---|---|---|---|
Taxa | Species | Presence | Material | Notes | Images |
Chasmaporthetes | C. sp. | Sidi Hakoma member and Denen Dora member at Hadar. [13] | Hunting Hyena. | ||
Crocuta | C. dietrichi | Sidi Dakoma member, Denen Dora member, and Basal member at Hadar.[13] | Fragmented mandibles (DIK-32-1).[23][13] | Extinct relatives of modern Hyena. | |
C. eturono | Kada Hadar member. | partial maxilla (DIK-73-1).[23] | |||
Hyaena | H. sp. | ||||
Ikelohyaena | I. abronia | Sidi Hakoma member at Hadar.[13] | Early member of the Parahyena lineage and the earliest close relative of the modern striped hyena. | ||
Pliocrocuta | P. cf. perierri | Sidi Hakoma and Denen Dora Member. |
Mustelidae
Mustelids of the Hadar Formation | |||||
---|---|---|---|---|---|
Taxa | Species | Presence | Material | Notes | Images |
Enhydriodon | E. dikikae | The Basal and Sidi Hakoma Members at Dikika site.[14][23] | Partial skull (DIK-56-9) and humerus (DIK-78-1).[14][23] | The extinct giant river otter. | |
Lutra | L. hearsti | Lower part of Sidi Hakoma Member at Dikika site. | Upper teeth fragments (DIK-50–35).[23] | Extinct relative of otters | |
Mellivora | M. sp. | ||||
Poecilogale | P. sp |
Herpestidae
Herpestids of the Hadar Formation | |||||
---|---|---|---|---|---|
Taxa | Species | Presence | Material | Notes | Images |
Herpestes | H. sp. | Dikika site. | Tooth fragments. | A mongoose. |
Viverridae
Viverrids of the Hadar Formation | |||||
---|---|---|---|---|---|
Taxa | Species | Presence | Material | Notes | Images |
Civettictis | C. sp. | A civet. |
Proboscidea
Deinotheriidae
Deinotheriidae of the Hadar Formation | |||||
---|---|---|---|---|---|
Taxa | Species | Presence | Material | Notes | Images |
Deinotherium | D. bozasi | Denen Dora Member, Kada Hadar Member and Sidi Hakoma Member. | Complete cranium skull and teeth fragments. | A large extinct elephant-like proboscidean. | ![]() |
Elephantidae
Elephantids of the Hadar Formation | |||||
---|---|---|---|---|---|
Taxa | Species | Presence | Material | Notes | Images |
Elephas | E. ekorensis | Sidi Hakoma and Denen Dora members of the Hadar Formation at Hadar and Dikika. [18] | Mandibles and molar fragments. | Extinct lineage related with Asian elephant. | |
Loxodonta | L. adaurora | Denen Dora Member and Sidi Hakoma Member at Hadar.[17] | Mandibles and molar fragments. | An extinct species of a modern-day African Elephant | |
L. exoptata | Denen Dora Member. | Undescribed Molars.[17] | |||
Mammuthus | M. sp. | Sidi Hakoma and Denen Dora Member. | Undescribed Molars. [17] | a mammoth relative. | |
Palaeoloxodon recki | P. r. brumpti | Sidi Hakoma Member.[18] | Mandibles and molar fragments. | An extinct relative of Elephants that are native to Africa. | ![]() |
Perissodactyla
Equidae
Equids of the Hadar Formation | |||||
---|---|---|---|---|---|
Taxa | Species | Presence | Material | Notes | Images |
Eurygnathohippus | E. afarense | Kada Hadar Member. | Partial skull (AL 363-18) and mandibles.[15] | Extinct relatives of Hipparionine horse. | |
E. hasumense | Denen Dora Member. | Partial skull (AL 340-8) and mandibles.[15] |
Rhinocerotidae
Rhinocerotidaes of the Hadar Formation | |||||
---|---|---|---|---|---|
Taxa | Species | Presence | Material | Notes | Images |
Ceratotherium | C. mauritanicum | Sidi Hakoma member at Lower Awash of Dikika. | Complete cranium skull (AL-129-25).[16] | Ancestor of modern White Rhinoceros. | |
Diceros | D. praecox | Sidi Hakoma member at Lower Awash of Dikika. | Incomplete cranium skull fragments and jawbones.[16] | A direct ancestor of modern Black Rhinoceros, formally named Ceratotherium praecox. |
Primates
Cercopithecidae
Cercopithecids of the Hadar Formation | |||||
---|---|---|---|---|---|
Taxa | Species | Presence | Material | Notes | Images |
Cercopithecoides | C. meaveae | Sidi Hakoma member. | Partial skeleton (A.L. 2-64 and A.L. 222-14 ).[25] | A colobine monkey. | |
Rhinocolobus | R. turkanaensis | Sidi Hakoma and Denen Dora Members. | Preserved mandibles and teeth fragments.[25] | Extinct relatives of colobus monkey. | |
Parapapio | P. cf. jonesi | Sidi Hakoma and Kada Hadar Members. | Complete postcranial skull, partial limbs, teeth fragments, and mandibles.[25] | A prehistoric baboon. | |
Theropithecus oswaldi[33] | T. o. darti | Basal, Denen Dora, Kada Hadar, and Sidi Hakoma Member. | Postcranial skulls differentiate between male and female.[25] | Extinct relatives of Gelada baboons | |
T. o. ecki | Denen Dora Member. | A female cranium skull and mandible. | |||
T. o. oswaldi | Sidi Hakoma and Kada Hadar Member. | Postcranial skulls differentiate between male and female.[25] |
Hominidae
Hominids of the Hadar Formation | |||||
---|---|---|---|---|---|
Taxa | Species | Presence | Material | Notes | Images |
Australopithecus | A. afarensis | Basal Member, Denen Dora Member, Kada Hadar Member, and Sidi Hakoma Member. | Partial skeletons such as small cranial capacity, primitive occipital, basal cranium anatomy, high frequency of unicuspid third premolars, prognathic face, and primitive mandibular anatomy. | An australopithecine hominid. | ![]() ![]() |
Rodentia
Rodents of the Hadar Formation | |||||
---|---|---|---|---|---|
Taxa | Species | Presence | Material | Notes | Images |
Acomys | A. cf. lavocati | Mandible specimens.[21] | A spiny mouse. | ||
A. cf. lemniscomys | right mandible and a broken tooth root.[21] | ||||
Gerbilliscus | G. sp. | A right maxillary fragment with M2-3 and a right mandible fragment with m1-3.[21] | A relative of gerbil species. | ||
Golunda | G. gurai. | An Murine rodent that are closely related to Indian bush rat. | |||
Hystrix | H. cf. makapanensis. | Sidi Hakoma Member.[20] | An Old World porcupine. | ||
Mastomys | M. cf. minor | Partial skull and mandibles.[21] | Extinct relative of the multimammate mouse. | ||
Millardia | M. coppensi | Extinct relatives of soft-furred rat. | |||
M. taiebi | |||||
Mus | M. indet | Unknown relative of Murini. | |||
Oenomys | O. tiercelini. | An extinct relative of modern Rusty-nosed rat. | |||
Pelomys | P. cf dietrichi | mandibles. | a groove-toothed swamp rats. | ||
Praomys | P. sp. | Sidi Hakoma Member. | A.L. 894-35993, a right mandible with m1-3.[21] | A Muridae species. | |
Saidomys | S. afarensis | Sidi Hakoma Member. | An extinct relative of a muroid mouse. | ||
Tatera | T. sp. | Sidi Hakoma and Denen Dora Member. | Extinct relative of Indian gerbil | ||
Thallomys | T. cf. quadrilobatus | A complete skull with both mandibles.[21] | Extinct relatives of the rat family that are endemic to Africa. | ||
Thryonomys | T. swinderianus | A cane rat. | |||
Xenohystrix | X. cf. crassidens | Sidi Hakoma and Denen Dora Member. | An old-world porcupine. | ||
Xerus | X. sp. | Kada Hadar Member. | An unstriped ground squirrel. |
Birds
Birds of the Hadar Formation | |||||
---|---|---|---|---|---|
Taxa | Species | Presence | Material | Notes | Images |
Anhinga | A. sp. | ||||
Balearica | B. sp. | ||||
Plectropterus | P. sp. | ||||
Struthio | S. sp. |
Reptiles
Reptiles of the Hadar Formation | |||||
---|---|---|---|---|---|
Taxa | Species | Presence | Material | Notes | Images |
Bitis | B. sp. | ||||
Crocodylus | C. sp. | ||||
Python | P. sp. | ||||
Varanus | V. sp. |
See also
References
- ^ a b Jon Kalb Adventures in the Bone Trade (New York: Copernicus Books, 2001), p. 83
- ^ a b E. N. Dimaggio et al., "Tephrostratigraphy and depositional environment of young (<2.94 Ma) Hadar Formation deposits at Ledi-Geraru, Afar, Ethiopia", Journal of African Earth Sciences 112A (December 2015), pp. 234–250 (Figure 2), doi:10.1016/j.jafrearsci.2015.09.018.
- ^ a b c Johanson, Donald (1 March 2017). "The paleoanthropology of Hadar, Ethiopia". Comptes Rendus Palevol. Des gènes à la culture / From genes to culture. 16 (2): 140–154. doi:10.1016/j.crpv.2016.10.005. ISSN 1631-0683.
- ^ Halstead, L. B. (1984). A la recherche du passé: la vie sur Terre, des origines aux premiers hommes. Hachette. ISBN 2010096029. OCLC 25125386.
- ^ Feibel, Craig S.; Christopher J. Campisano (2004). "Sedimentary Patterns in the Pliocene Hadar Formation, Afar Rift, Ethiopia".
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(help) - ^ a b Hogenboom, Melissa. "The 'Lucy' fossil rewrote the story of humanity". Retrieved 25 October 2018.
- ^ "Lucy's Story | Institute of Human Origins". iho.asu.edu. Retrieved 25 October 2018.
- ^ Reed, Kaye & Bibi, Faysal. (2011). Fossil Tragelaphini (Artiodactyla: Bovidae) from the Late Pliocene Hadar Formation, Afar Regional State, Ethiopia. Journal of Mammalian Evolution. 18. 57-69. 10.1007/s10914-010-9146-6.
- ^ a b c d e f g h i j k l m n o p q r s t Geraads, D.; Bobe, R.; Reed, K. (2012). "Pliocene Bovidae (Mammalia) from the Hadar Formation of Hadar and Ledi-Geraru, Lower Awash, Ethiopia". Journal of Vertebrate Paleontology, 32(1): 180–197. http://www.jstor.org/stable/41407716
- ^ Reed, K. E. (2008). "Paleoecological Patterns at the Hadar Hominin Site, Afar Regional State, Ethiopia". Journal of Human Evolution, 54(6): 743–768. https://doi.org/10.1016/J.JHEVOL.2007.08.013
- ^ a b c d e Cooke, H.B. (1978). Pliocene-Pleistocene Suidae from Hadar, Ethiopia.
- ^ a b c d e Werdelin, Lars & Lewis, Margaret. (2001). A revision of the genus Dinofelis (Mammalia, Felidae). Zoological Journal of the Linnean Society. 132. 147 - 258. 10.1111/j.1096-3642.2001.tb02465.x.
- ^ a b c d e Werdelin, L., & Lewis, M. E. (2008). New Species of Crocuta from the Early Pliocene of Kenya, with an Overview of Early Pliocene Hyenas of Eastern Africa. Journal of Vertebrate Paleontology, 28(4), 1162–1170. http://www.jstor.org/stable/20491047
- ^ a b c Geraads, Denis; Alemseged, Zeresenay; Bobe, René; Reed, Denné (2011). "Enhydriodon dikikae, sp. nov. (Carnivora: Mammalia), a gigantic otter from the Pliocene of Dikika, Lower Awash, Ethiopia". Journal of Vertebrate Paleontology. 31 (2): 447–453. doi:10.1080/02724634.2011.550356. S2CID 84797296.
- ^ a b c Gilbert, William & Bernor, Raymond. (2009). Equidae. 10.1525/california/9780520251205.003.0006.
- ^ a b c Geraads, D. (2005). "Pliocene Rhinocerotidae (Mammalia) from Hadar and Dikika (Lower Awash, Ethiopia), and a Revision of the Origin of Modern African Rhinos". Journal of Vertebrate Paleontology, 25(2), 451–461. http://www.jstor.org/stable/4524458
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