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Homo rudolfensis
Temporal range:Pleistocene,2.5/1.9–1.85/1.55Ma
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Reconstruction of the KNM-ER 1470 skull
Reconstruction of the KNM-ER 1470 skull
Scientific classificationEdit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Primates
Suborder: Haplorhini
Infraorder: Simiiformes
Family: Hominidae
Subfamily: Homininae
Tribe: Hominini
Genus: Homo(?)
Species:
†H. rudolfensis
Binomial name
†Homo rudolfensis
(Alekseyev,1986)
Synonyms

Homo rudolfensisis an extinct species ofarchaic humanfrom theEarly Pleistoceneof East Africa about 2 million years ago (mya). BecauseH. rudolfensiscoexisted with several otherhominins,it is debated what specimens can be confidently assigned to this species beyond thelectotypeskull KNM-ER 1470 and other partial skull aspects. No bodily remains are definitively assigned toH. rudolfensis.Consequently, both itsgenericclassification and validity are debated without any wide consensus, with some recommending the species to actually belong to the genusAustralopithecusasA. rudolfensisorKenyanthropusasK. rudolfensis,or that it issynonymouswith the contemporaneous and anatomically similarH. habilis.

H. rudolfensisis distinguished fromH. habilisby larger size, but it is also argued that this species actually consists of maleH. habilisspecimens, assuming thatH. habiliswassexually dimorphicand males were much larger than females. Because no bodily remains are definitely identified, body size estimates are largely based on the stature ofH. habilis.Using this, maleH. rudolfensismay have averaged about 160 cm (5 ft 3 in) in height and 60 kg (130 lb) in weight, and females 150 cm (4 ft 11 in) and 51 kg (112 lb). KNM-ER 1470 had a brain volume of about 750 cc (46 cu in). Like other earlyHomo,H. rudolfensishad largecheek teethand thickenamel.

EarlyHomospecies exhibit marked brain growth compared toAustralopithecuspredecessors, which is typically explained as a change in diet with a calorie-rich food source, namely meat. Though not associated with tools, dental anatomy suggests some processing of plant or meat fiber before consumption, though the mouth could still effectively chew through mechanically challenging food, indicating tool use did not greatly affect diet.

Research history

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Homofamily tree showingH. rudolfensisandH. habilisat the base as offshoots of the human line[1]
Reconstruction ofH. rudolfensisbyMauricio Antón

The first fossils were discovered in 1972 alongLake Turkana(at the time called Lake Rudolf) in Kenya, and were detailed by Kenyan palaeoanthropologistRichard Leakeythe following year. The specimens were: a large and nearly complete skull (KNM-ER 1470, thelectotype) discovered by Bernard Ngeneo, a local; a rightfemur(KNM-ER 1472) discovered by J. Harris; an upper femur (proximal) fragment (KNM-ER 1475) discovered by fossil collectorKamoya Kimeu;and a complete left femur (KNM-ER 1481) discovered by Harris. However, it is unclear if the femora belong to the same species as the skull. Leakey classified them under the genusHomobecause he had reconstructed the skull fragments so that it had a large brain volume and a flat face, but did not assign them to a species. Because thehorizonthey were discovered in was, at the time, dated to 2.9–2.6 million years ago (mya), Leakey thought these specimens were a very early human ancestor.[2]This challenged the major model of human evolution at the time whereAustralopithecus africanusgave rise toHomoabout 2.5 mya, but ifHomohad already existed at this time, it would call for serious revisions.[3]However, the area was redated to about 2 mya in 1977 (the same time period asH. habilisandH. ergaster/H. erectus),[4]and more precisely to 2.1–1.95 mya in 2012.[5]They were first assigned to the specieshabilisin 1975 by anthropologistsColin GrovesandVratislav Mazák.In 1978, in a joint paper with Leakey and English anthropologistAlan Walker,Walker suggested the remains belong inAustralopithecus(and that the skull was incorrectly reconstructed), but Leakey still believed they belonged toHomo,though they both agreed that the remains could belong tohabilis.[6]

KNM-ER 1470 was much larger than the Olduvai remains, so the termsH. habilissensu lato( "in the broad sense" ) andH. habilissensu stricto( "in the strict sense" ) were used to include or exclude the larger morph, respectively.[7][8]In 1986, English palaeoanthropologist Bernard Wood first suggested these remains represent a differentHomospecies, which coexisted withH. habilisandH. ergaster/H. erectus.CoexistingHomospecies conflicted with the predominant model of human evolution at the time which was that modern humans evolved in a straight line directly fromH. ergaster/H. erectuswhich evolved directly fromH. habilis.[9]In 1986, the remains were placed into a new species,rudolfensis,by Russian anthropologistValery Alekseyev[10](but he used the genusPithecanthropus,which was changed toHomothree years later by Groves).[11]In 1999, Kennedy argued that the name was invalid because Alekseyev had not assigned aholotype.[12]Pointing out that this is in fact not mandatory, Wood the same year nevertheless designated KNM-ER 1470 as the lectotype.[13]However, the validity of this species has also been debated on material grounds, with some arguing thatH. habiliswas highlysexually dimorphiclike modern non-human apes, with the larger skulls classified as "H. rudolfensis"actually representing maleH. habilis.[8][13]In 1999, Wood and biological anthropologist Mark Collard recommended movingrudolfensisandhabilistoAustralopithecusbased on the similarity of dental adaptations. However, they conceded that dental anatomy is highly variable amonghomininsand not always reliable when formulating family trees.[14]

KNM-ER 1802

In 2003, Australian anthropologist David Cameron concluded that the earlieraustralopithecineKenyanthropus platyopswas the ancestor ofrudolfensis,and reclassified it asK. rudolfensis.He also believed thatKenyanthropuswas more closely related toParanthropusthanHomo.[15]In 2008, a re-reconstruction of the skull concluded it was incorrectly restored originally, though agreed with the classification asH. rudolfensis.[16]In 2012, British palaeoanthropologistMeave Leakeydescribed the juvenile partial face KNM-ER 62000 discovered inKoobi Fora,Kenya; noting it shares several similarities to KNM-ER 1470 and is smaller, she assigned it toH. rudolfensis,and, because prepubescent male and female bones should be indistinguishable, differences between juvenileH. rudolfensisand adultH. habilisspecimens support species distinction. She also concluded that the jawbone KNM-ER 1802, an important specimen often used in classifying other specimens asH. rudolfensis,actually belongs to a different (possibly undescribed) species,[17]but American palaeoanthropologistTim D. Whitebelieves this to be premature because it is unclear how wide the range of variation is in early hominins.[7]The 2013 discovery of the 1.8 Ma GeorgianDmanisi skullswhich exhibit several similarities with earlyHomohave led to suggestions that all contemporary groups of earlyHomoin Africa, includingH. habilisandH. rudolfensis,are the same species and should be assigned toH. erectus.[18][19]There is still no wide consensus on howrudolfensisandhabilisrelate toH. ergasterand descendent species.[20]

Beyond KNM-ER 1470, there is disagreement on which specimens actually belong inH. rudolfensisas it is difficult to assign with accuracy remains that do not preserve the face and jaw.[7][8]NoH. rudolfensisbodily elements have been definitively associated with a skull and thus to the species.[21]Most proposedH. rudolfensisfossils come from Koobi Fora and date to 1.9–1.85 mya. Remains from theShungura Formation,Ethiopia, and Uraha, Malawi, are dated as far back as 2.5–2.4 mya, which would make it the earliest identified species ofHomo.The latest potential specimen is KNM-ER 819 dating to 1.65–1.55 mya.[21]: 210 

Nonetheless,H. rudolfensisandH. habilisgenerally are recognised members of the genus at the base of the family tree, with arguments for synonymisation or removal from the genus not widely adopted.[22]Though it is now largely agreed upon thatHomoevolved fromAustralopithecus,the timing and placement of this split has been much debated, with manyAustralopithecusspecies having been proposed as the ancestor. The discovery ofLD 350-1,the oldestHomospecimen, dating to 2.8 mya, in theAfar Regionof Ethiopia may indicate that the genus evolved fromA. afarensisaround this time. The species LD 350-1 belongs to could be the ancestor ofH. rudolfensisandH. habilis,but this is unclear.[23]Based on 2.1 million year old stone tools fromShangchen,China, possibly an ancestral species toH. rudolfensisandH. habilisdispersed across Asia.[24]

Anatomy

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Skull

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KNM-ER 1470H. rudolfensis(left) vs KNM-ER 1813H. habilis(right)

In 1973, Mr. Leakey had reconstructed the skull KNM-ER 1470 with a flat face and a brain volume of 800 cc (49 cu in).[2]In 1983, American physical anthropologistRalph Hollowayrevised the base of the skull and calculated a brain volume of 752–753 cc (45.9–46.0 cu in).[25]For comparison,H. habilisspecimens average about 600 cc (37 cu in), andH. ergaster850 cc (52 cu in).[26]Anthropologist Timothy Bromage and colleagues revised the face again at a 5° incline (slightlyprognathic) instead of completely flat, but pushed thenasal boneback directly beneath thefrontal bones.He then said it was possible to predict brain size based on just the face and (disregarding the braincase) calculated 526 cc (32.1 cu in), and chalked up the errors of Leakey's reconstruction to a lack of research of the biological principles of facial anatomy at the time as well asconfirmation bias,as a flat-faced reconstruction of the skull aligned with the predominant model of human evolution at the time. This was refuted by American palaeoanthropologistJohn D. Hawksbecause the skull remained more or less unchanged except for the 5° rotation outwards.[27]Bromage and colleagues returned in 2008 with a revised skull reconstruction and brain volume estimate of 700 cc (43 cu in).[16]

UR 501, the oldestH. rudolfensisspecimen[28][29]

Fossils have generally been classified intoH. rudolfensisdue to large skull size, flatter and broader face, broadercheek teeth,more complex toothcrownsandroots,and thickerenamelcompared toH. habilis.[28]EarlyHomoare characterised by larger teeth compared to laterHomo.The cheek teeth of KNM-ER 60000, a jawbone, in terms of size are on the lower end for earlyHomo,except for the thirdmolarwhich is within range. The molars increase in size towards the back of the mouth. The tooth rows of KNM-ER 1470, KNM-ER 60000, and KNM-ER 62000 are rectangular, whereas the tooth row of KNM-ER 1802 is U-shaped, which may indicate that these two morphs represent different species,[17]or demonstrate the normal range of variation forH. rudolfensisjaws.[7]In UR 501 from Uraha, Malawi—the oldestH. rudolfensisspecimen dating to 2.5–2.3 mya—thetooth enamelthickness is the same as in other earlyHomo,but the enamel on the molars is almost as thick asParanthropusmolars (which have some of the thickest enamel of any hominin). Such a wide variation in enamel thickness across the cheek teeth is not exhibited in KNM-ER 1802, which may indicate regional differences amongH. rudolfensispopulations.[28][29]

Build

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Body size estimates ofH. rudolfensisandH. habilistypically conclude a small size comparable to australopithecines. These largely depend on theH. habilispartial skeleton OH 62 estimated at 100–120 cm (3 ft 3 in – 3 ft 11 in) in height and 20–37 kg (44–82 lb) in weight.H. rudolfensisis thought to be bigger thanH. habilis,but it is unclear how big this species was as no bodily elements have been definitively associated with a skull.[30]Based on just the KNM-ER 1470 skull, maleH. rudolfensiswere estimated to have been 160 cm (5 ft 3 in) in height and 60 kg (130 lb) in weight, and females 150 cm (4 ft 11 in) and 51 kg (112 lb).[26]

For specimens that might beH. rudolfensis:thefemurKNM-ER 1472 which may also beH. habilisorH. ergasterwas estimated at 155.9 cm (5 ft 1 in) and 41.8 kg (92 lb), thehumerusKNM-ER 1473 162.9 cm (5 ft 4 in) and 47.1 kg (104 lb), the partial leg KNM-ER 1481 which may also beH. ergaster156.7 cm (5 ft 2 in) and 41.8 kg (92 lb), thepelvisKNM-ER 3228 which may also beH. ergaster165.8 cm (5 ft 5 in) and 47.2 kg (104 lb), and the femur KNM-ER 3728 which may beH. habilisorP. boisei153.3 cm (5 ft) and 40.3 kg (89 lb).[30]It is generally assumed that pre-H. ergasterhominins, includingH. rudolfensisandH. habilis,exhibitedsexual dimorphismwith males markedly bigger than females. However, relative female body mass is unknown in either species.[21]

Early hominins, includingH. rudolfensis,are thought to have had thick body hair coverage like modern non-human apes because they appear to have inhabited cooler regions and are thought to have had a less active lifestyle than (presumed hairless) post-ergasterspecies, and so probably required thick body hair to stay warm.[31]The juvenile specimen KNM-ER 62000, a partial face, has the same age landmarks as a 13 to 14 year old modern human, but more likely died at around 8 years of age due to the presumed faster growth rate among early hominins based on dental development rate.[17]

Culture

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Reconstruction of KNM-ER 1470 skull and jaw

It is typically thought that the diets of earlyHomohad a greater proportion of meat thanAustralopithecus,and that this led to brain growth. The main hypotheses regarding this are: meat is energy- and nutrient-rich and put evolutionary pressure on developing enhanced cognitive skills to facilitate strategic scavenging and monopolise fresh carcasses, or meat allowed the large and calorie-expensive ape gut to decrease in size allowing this energy to be diverted to brain growth. Alternatively, it is also suggested that earlyHomo,in a drying climate with scarcer food options, relied primarily on undergroundstorage organs(such astubers) and food sharing, which facilitated social bonding among both male and female group members. However, unlike what is presumed forH. ergasterand laterHomo,short-statured earlyHomowere likely incapable ofendurance runningand hunting, and the long andAustralopithecus-like forearm ofH. habiliscould indicate earlyHomowere stillarborealto a degree. Also, organisedhunting and gatheringis thought to have emerged inH. ergaster.Nonetheless, the proposed food-gathering models to explain large brain growth necessitate increased daily travel distance.[32]Largeincisorsize inH. rudolfensisandH. habiliscompared toAustralopithecuspredecessors implies these two species relied on incisors more. The large,Australopithecus-like molars could indicate more mechanically challenging food compared to laterHomo.Thebodies of the mandiblesofH. rudolfensisand other earlyHomoare thicker than those of modern humans and all living apes, more comparable toAustralopithecus.The mandibular body resists torsion from thebite forceor chewing, meaning their jaws could produce unusually powerful stresses while eating.[21]

H. rudolfensisis not associated with any tools. However, the greater molarcuspreliefinH. rudolfensisandH. habiliscompared toAustralopithecussuggests the former two used tools to fracture tough foods (such as pliable plant parts or meat), otherwise the cusps would have been more worn down. Nonetheless, the jaw adaptations for processing mechanically challenging food indicates technological advancement did not greatly affect their diet. Large concentrations of stone tools are known from Koobi Fora. Because these aggregations are coincident with the emergence ofH. ergaster,it is probableH. ergastermanufactured them, though it is not possible to definitively attribute the tools to a species becauseH. rudolfensis,H. habilis,andP. boiseiare also well known from the area.[21]The oldest specimen ofHomo,LD 350-1,is associated with theOldowanstone toolindustry,meaning this tradition had been in use by the genus since near its emergence.[33]

EarlyH. rudolfensisandParanthropushave exceptionally thick molars for hominins, and the emergence of these two coincides with a cooling and aridity trend in Africa about 2.5 mya. This could mean they evolved due to climate change. Nonetheless, in East Africa, tropical forests and woodlands still persisted through periods of drought.[28]H. rudolfensiscoexisted withH. habilis,H. ergaster,andP. boisei.[21]

See also

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References

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