Ardipithecus ramidus

Ardipithecus ramidus is a species of australopithecine from the Afar region of Early Pliocene Ethiopia 4.4 million
years ago (mya). A. ramidus, unlike modern hominids, has adaptations for both walking on two legs (bipedality) Ardipithecus ramidus
and life in the trees (arboreality). However, it would not have been as efficient at bipedality as humans, nor at Temporal range: Zanclean
arboreality as non-human great apes. Its discovery, along with Miocene apes, has reworked academic
understanding of the chimpanzee-human last common ancestor from appearing much like modern day
chimpanzees, orangutans and gorillas to being a creature without a modern anatomical cognate.

The facial anatomy suggests that A. ramidus males were less aggressive than those of modern chimps, which is
correlated to increased parental care and monogamy in primates. It has also been suggested that it was among the
earliest of human ancestors to use some proto-language, possibly capable of vocalizing at the same level as a
human infant. A. ramidus appears to have inhabited woodland and bushland corridors between savannas, and was
a generalized omnivore.

Contents
Taxonomy A. ramidus at the Museo Nacional de
Description Ciencias Naturales
Paleobiology
Scientific classification
Paleoecology
Kingdom: Animalia
See also
Phylum: Chordata
References
External links Class: Mammalia
Order: Primates
Suborder: Haplorhini
Taxonomy
Infraorder: Simiiformes
The first remains were described in 1994 by American anthropologist Tim Family: Hominidae
D. White, Japanese paleoanthropologist Gen Suwa, and Ethiopian
paleontologist Berhane Asfaw. The holotype specimen, ARA-VP-6/1, Subfamily: Homininae
comprised an associated set of 10 teeth; and there were 16 other paratypes Tribe: Hominini
identified, preserving also skull and arm fragments. These were unearthed
in the 4.4 million year (Ma) deposits of the Afar region in Aramis, Genus: †Ardipithecus
Ethiopia from 1992 to 1993, making them the oldest hominin remains at
Species: †A. ramidus
the time, surpassing Australopithecus afarensis. They initially classified it
as Australopithecus ramidus, the species name deriving from the Afar Binomial name
language ramid "root".[1] In 1995, they made a corrigendum
†Ardipithecus ramidus
recommending it be split off into a separate genus, Ardipithecus; the name
(White, Suwa & Asfaw, 1994)
Map showing discovery locations of stems from Afar ardi "ground" or "floor".[2] The 4.4 million year old
various australopithecines female ARA-VP 6/500 ("Ardi") is the most complete specimen.[3] Synonyms
Fossils from at least nine A. ramidus individuals at As Duma, Gona
Australopithecus ramidus
Western Margin, Afar, were unearthed from 1993 to 2003. The fossils were dated to between 4.32 and 4.51 million
years ago.[4]

In 2001, 6.5–5.5 million year old fossils from the Middle Awash were classified as a subspecies of A. ramidus by Ethiopian paleoanthropologist Yohannes
Haile-Selassie.[5] In 2004, Haile-Selassie, Suwa, and White split it off into its own species, A. kadabba.[6] A. kadabba is considered to have been the direct
ancestor of A. ramidus, making Ardipithecus a chronospecies.[7]

The exact affinities of Ardipithecus have been debated. White, in 1994, considered A. ramidus to have been more closely related to humans than
chimpanzees, though noting it to be the most ape-like fossil hominin to date.[1] In 2001, French paleontologist Brigitte Senut and colleagues aligned it more
closely to chimpanzees,[8] but this has been refuted.[5] In 2009, White and colleagues reaffirmed the position of Ardipithecus as more closely related to
modern humans based on dental similarity, a short base of the skull, and adaptations to bipedality.[9] In 2011, primatologist Esteban Sarmiento said that there
is not enough evidence to assign Ardipithecus to Hominini (comprising both humans and chimps),[10] but its closer affinities to humans have been reaffirmed
in following years.[11] White and colleagues consider it to have been closely related to or the ancestor of the temporally close Australopithecus anamensis,
which was the ancestor to Au. afarensis.[3]

African hominin timeline (in mya)

View references
Before the discovery of Ardipithecus and other pre-Australopithecus hominins, it was assumed that the chimpanzee–human last common ancestor and
preceding apes appeared much like modern day chimpanzees, orangutans and gorillas, which would have meant these three changed very little over millions
of years. Their discovery led to the postulation that modern great apes, much like humans, evolved several specialized adaptations to their environment (have
highly derived morphologies), and their ancestors were comparatively poorly adapted to suspensory behavior or knuckle walking, and did not have such a
specialized diet. Also, the origins of bipedality were thought to have occurred due to a switch from a forest to a savanna environment, but the presence of
bipedal pre-Australopithecus hominins in woodlands has called this into question,[12] though they inhabited wooded corridors near or between savannas. It is
also possible that Ardipithecus and pre-Australopithecus were random offshoots of the hominin line.[13]

Description
Assuming subsistence was primarily sourced from climbing in trees, A. ramidus may not have exceeded 35–60 kg
(77–132 lb). "Ardi," a larger female specimen, was estimated to have stood 117–124 cm (3 ft 10 in–4 ft 1 in) and
weighed 51 kg (112 lb) based on comparisons with large-bodied female apes.[14] Unlike the later Australopithecus
but much like chimps and humans, males and females were about the same size.[3]

A. ramidus had a small brain, measuring 300–350 cc (18–21 cu in). This is slightly smaller than a modern bonobo or
chimp brain, but much smaller than the brain of Australopithecus–about 400–550 cc (24–34 cu in)–and roughly 20%
the size of the modern human brain. Like chimps, the A. ramidus face was much more pronounced (prognathic) than
modern humans.[15] The size of the upper canine tooth in A. ramidus males was not distinctly different from that of
females (only 12% larger), in contrast to the sexual dimorphism observed in chimps where males have significantly
larger and sharper upper canines than females.[3][16]

A. ramidus feet are better suited for walking than chimps. However, like non-human great apes, but unlike all
previously recognized human ancestors, it had a grasping big toe adapted for locomotion in the trees (an arboreal
lifestyle), though it was likely not as specialized for grasping as it is in modern great apes.[17][9] Its tibial and tarsal
lengths indicate a leaping ability similar to bonobos.[10] It lacks any characters suggestive of specialized suspension,
vertical climbing, or knuckle walking; and it seems to have used a method of locomotion unlike any modern great
ape, which combined arboreal palm walking clambering and a form of bipedality more primitive than
Reconstruction of Ardipithecus Australopithecus. The discovery of such unspecialized locomotion led American anthropologist Owen Lovejoy and
skeleton colleagues to postulate that the chimpanzee–human last common ancestor used a similar method of locomotion.[18][9]

The upper pelvis (distance from the sacrum to the hip joint) is shorter than in any known ape. It is inferred to have
had a long lumbar vertebral series, and lordosis (human curvature of the spine), which are adaptations for bipedality. However, the legs were not completely
aligned with the torso (were anterolaterally displaced), and Ardipithecus may have relied more on its quadriceps than hamstrings which is more effective for
climbing than walking.[19][7] However, it lacked foot arches and had to adopt a flat-footed stance. These would have made it less efficient at walking and
running than Australopithecus and Homo. It may not have employed a bipedal gait for very long time intervals.[3] It may have predominantly used palm
walking on the ground,[20] Nonetheless, A. ramidus still had specialized adaptations for bipedality, such as a robust fibularis longus muscle used in pushing
the foot off the ground while walking (plantarflexion),[17] the big toe (though still capable of grasping) was used for pushing off, and the legs were aligned
directly over the ankles instead of bowing out like in non-human great apes.[21]

Paleobiology
The reduced canine size and reduced skull robustness in A. ramidus males (about the same size in males and females) is typically correlated with reduced
male–male conflict, increased parental investment, and monogamy.[9][7] Because of this, it is assumed that A. ramidus lived in a society similar to bonobos
and ateline monkeys[16] due to a process of self domestication (becoming more and more docile which allows for a more gracile build). Because a similar
process is thought to have occurred with the comparatively docile bonobos from more aggressive chimps, A. ramidus society may have seen an increase in
maternal care and female mate selection compared to its ancestors.[22] Alternatively, it is possible that increased male size is a derived trait instead of basal (it
evolved later rather than earlier), and is a specialized adaptation in modern great apes as a response to a different and more physically exerting lifestyle in
males than females rather than being tied to interspecific conflict.[12]

Australian anthropologists Gary Clark and Maciej Henneberg argued that such shortening of the skull—which may have caused a descension of the larynx—
as well as lordosis—allowing better movement of the larynx—increased vocal ability, significantly pushing back the origin of language to well before the
evolution of Homo. They argued that self domestication was aided by the development of vocalization, living in a pro-social society, as a means of non-
violently dealing with conflict. They conceded that chimps and A. ramidus likely had the same vocal capabilities, but
said that A. ramidus made use of more complex vocalizations, and vocalized at the same level as a human infant due
to selective pressure to become more social. This would have allowed their society to become more complex. They
also noted that the base of the skull stopped growing with the brain by the end of juvenility, whereas in chimps it
continues growing with the rest of the body into adulthood; and considered this evidence of a switch from a gross
skeletal anatomy trajectory to a neurological development trajectory due to selective pressure for sociability.
Nonetheless, their conclusions are highly speculative.[23][22]

American primatologist Craig Stanford postulated that A. ramidus

behaved similarly to chimps, which frequent both the trees and the
ground, have a polygynous society, hunt cooperatively, and are the most
technologically advanced non-human.[24] However, Clark and
Chimp skull (note the large canines
Henneberg concluded that Ardipithecus cannot be compared to chimps,
and elongated face)
having been too similar to humans.[22] According to French
paleoprimatologist Jean-Renaud Boisserie, the hands of Ardipithecus
would have been dextrous enough to handle basic tools, though it has not been associated with any tools.[25]

Hypothetical restoration of a female
Ardipithecus using a hammer and anvil
to crack open a nut
The teeth of A. ramidus indicate that it was likely a generalized omnivore and fruit eater which predominantly
consumed C3 plants in woodlands or gallery forests. The teeth lacked adaptations for abrasive foods.[16][9][10]
Lacking the speed and agility of chimps and baboons, meat intake by Ardipithecus, if done, would have been
sourced from only what could have been captured by limited pursuit, or from scavenging carcasses.[26]

Paleoecology
Half of the large mammal species associated with A. ramidus at Aramis are spiral-horned antelope and colobine monkeys (namely Kuseracolobus and
Pliopapio). There are a few specimens of primitive white and black rhino species, and elephants, giraffes, and hippo specimens are less abundant. These
animals indicate that Aramis ranged from wooded grasslands to forests, but A. ramidus likely preferred the closed habitats,[27] specifically riverine areas as
such water sources may have supported more canopy coverage.[28] Aramis as a whole generally had less than 25% canopy cover.[13] There were exceedingly
high rates of scavenging, indicating a highly competitive environment somewhat like Ngorongoro Crater. Predators of the area were the hyenas Ikelohyaena
abronia and Crocuta dietrichi, the bear Agriotherium, the cat Dinofelis and Megantereon, the dog Eucyon, and crocodiles.[29] Bayberry, hackberry, and palm
trees appear to have been common at the time from Aramis to the Gulf of Aden; and botanical evidence suggests a cool, humid climate.[30] Conversely,
annual water deficit (the difference between water loss by evapotranspiration and water gain by precipitation) at Aramis was calculated to have been about
1,500 mm (59 in), which is seen in some of the hottest, driest parts of East Africa.[13]

Carbon isotope analyses of the herbivore teeth from the Gona Western Margin associated with A. ramidus indicate that these herbivores fed mainly on C4
plants and grasses rather than forest plants. The area seems to have featured bushland and grasslands.[31]

See also
Australopithecus – Genus of hominin ancestral to modern Graecopithecus – Extinct hominin from Miocene Greece
humans Orrorin – Postulated early hominin discovered in Kenya
Ardi – Designation of the fossilized skeletal remains of an Paranthropus – Contested extinct genus of hominins
Ardipithecus ramidus
Sahelanthropus – Extinct hominid from Miocene Africa

References
1. White, T. D.; Suwa, G.; Asfaw, B. (1994). "Australopithecus
ramidus, a new species of early hominid from Aramis, Ethiopia"
(https://web.archive.org/web/20130413081635/http://www2.ku.ed
u/~lba/courses/articles/White%20ramadis94.pdf) (PDF). Nature.
371 (6495): 306–312. Bibcode:1994Natur.371..306W (https://ui.a
dsabs.harvard.edu/abs/1994Natur.371..306W).
doi:10.1038/371306a0 (https://doi.org/10.1038%2F371306a0).
PMID 8090200 (https://pubmed.ncbi.nlm.nih.gov/8090200).
S2CID 4347140 (https://api.semanticscholar.org/CorpusID:43471
40). Archived from the original (http://www2.ku.edu/~lba/courses/
articles/White%20ramadis94.pdf) (PDF) on 2013-04-13.
2. White, T. D.; Suwa, G.; Asfaw, B. (1995). "Corrigendum:
Australopithecus ramidus, a new species of early hominid from
Aramis, Ethiopia" (https://doi.org/10.1038%2F375088a0). Nature.
375 (6526): 88. doi:10.1038/375088a0 (https://doi.org/10.1038%2
F375088a0). PMID 7677838 (https://pubmed.ncbi.nlm.nih.gov/76
77838).
3. White, T. D.; Lovejoy, C. O.; Asfaw, B.; Carlson, J. P.; Suwa, G.
(2015). "Neither chimpanzee nor human, Ardipithecus reveals the
surprising ancestry of both" (https://www.ncbi.nlm.nih.gov/pmc/arti
cles/PMC4413341). Proceedings of the National Academy of
Sciences. 112 (16): 4877–4884.
Bibcode:2015PNAS..112.4877W (https://ui.adsabs.harvard.edu/a
bs/2015PNAS..112.4877W). doi:10.1073/pnas.1403659111 (http
s://doi.org/10.1073%2Fpnas.1403659111). PMC 4413341 (http
s://www.ncbi.nlm.nih.gov/pmc/articles/PMC4413341).
PMID 25901308 (https://pubmed.ncbi.nlm.nih.gov/25901308).
4. Semaw, S.; Simpson, S. W.; et al. (2005). "Early Pliocene
hominids from Gona, Ethiopia". Nature. 433 (7023): 301–305.
Bibcode:2005Natur.433..301S (https://ui.adsabs.harvard.edu/abs/
2005Natur.433..301S). doi:10.1038/nature03177 (https://doi.org/1
0.1038%2Fnature03177). PMID 15662421 (https://pubmed.ncbi.n
lm.nih.gov/15662421). S2CID 4431031 (https://api.semanticschol
ar.org/CorpusID:4431031).
5. Haile-Selassie, Y. (2001). "Late Miocene hominids from the
Middle Awash, Ethiopia". Nature. 42 (6843): 179–181.
Bibcode:2001Natur.412..178H (https://ui.adsabs.harvard.edu/abs/
2001Natur.412..178H). doi:10.1038/35084063 (https://doi.org/10.
1038%2F35084063). PMID 11449272 (https://pubmed.ncbi.nlm.n
ih.gov/11449272). S2CID 4432082 (https://api.semanticscholar.or
g/CorpusID:4432082).
6. Haile-Selassie, Y.; Suwa, G.; White, T. D. (2004). "Late Miocene
Teeth from Middle Awash, Ethiopia, and Early Hominid Dental
Evolution". Science. 303 (5663): 1503–1505.
Bibcode:2004Sci...303.1503H (https://ui.adsabs.harvard.edu/abs/
2004Sci...303.1503H). doi:10.1126/science.1092978 (https://doi.o
rg/10.1126%2Fscience.1092978). PMID 15001775 (https://pubm
ed.ncbi.nlm.nih.gov/15001775). S2CID 30387762 (https://api.sem
anticscholar.org/CorpusID:30387762).
7. Lovejoy, C. O. (2014). "Ardipithecus and Early Human Evolution
in Light of Twenty-First-Century Developmental Biology". Journal
of Anthropological Research. 70 (3): 337–363.
doi:10.3998/jar.0521004.0070.301 (https://doi.org/10.3998%2Fjar.
0521004.0070.301). JSTOR 24394231 (https://www.jstor.org/stab
le/24394231). S2CID 84197134 (https://api.semanticscholar.org/
CorpusID:84197134).
 8. Senut, B.; Pickford, M.; Gommery, D.; et al. (2001). "Premier
hominidé du Miocène (formation de Lukeino, Kenya)" [First
hominid from the Miocene (Lukeino Formation, Kenya)]. Comptes
Rendus de l'Académie des Sciences (in French). 332 (2): 137–
144. Bibcode:2001CRASE.332..137S (https://ui.adsabs.harvard.e
du/abs/2001CRASE.332..137S). doi:10.1016/S1251-
8050(01)01529-4 (https://doi.org/10.1016%2FS1251-8050%280
1%2901529-4).
9. White, T. D.; Asfaw, B.; Beyene, Y.; Haile-Selassie, Y.; Lovejoy, C.
O.; Suwa, G.; WoldeGabriel, G. (2009). "Ardipithecus ramidus
and the Paleobiology of Early Hominids". Science. 326 (5949):
75–86. Bibcode:2009Sci...326...75W (https://ui.adsabs.harvard.ed
u/abs/2009Sci...326...75W). doi:10.1126/science.1175802 (http
s://doi.org/10.1126%2Fscience.1175802). PMID 19810190 (http
s://pubmed.ncbi.nlm.nih.gov/19810190). S2CID 20189444 (http
s://api.semanticscholar.org/CorpusID:20189444).
10. Sarmiento, E. E.; Meldrum, D. J. (2011). "Behavioral and
phylogenetic implications of a narrow allometric study of
Ardipithecus ramidus". HOMO. 62 (2): 75–108.
doi:10.1016/j.jchb.2011.01.003 (https://doi.org/10.1016%2Fj.jchb.
2011.01.003). PMID 21388620 (https://pubmed.ncbi.nlm.nih.gov/
21388620).
11. Kimbel, W. H.; Suwa, G.; Asfaw, B.; Rak, Y.; White, T. D. (2014).
"Ardipithecus ramidus and the evolution of the human cranial
base" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3903226).
Proceedings of the National Academy of Sciences. 111 (3): 948–
953. Bibcode:2014PNAS..111..948K (https://ui.adsabs.harvard.e
du/abs/2014PNAS..111..948K). doi:10.1073/pnas.1322639111 (h
ttps://doi.org/10.1073%2Fpnas.1322639111). PMC 3903226 (http
s://www.ncbi.nlm.nih.gov/pmc/articles/PMC3903226).
PMID 24395771 (https://pubmed.ncbi.nlm.nih.gov/24395771).
12. Lovejoy, C. O. (2009). "Reexamining Human Origins in Light of
Ardipithecus ramidus" (http://doc.rero.ch/record/211449/files/PAL
_E4439.pdf) (PDF). Science. 326 (5949): 74–74e8.
Bibcode:2009Sci...326...74L (https://ui.adsabs.harvard.edu/abs/2
009Sci...326...74L). doi:10.1126/science.1175834 (https://doi.org/
10.1126%2Fscience.1175834). PMID 19810200 (https://pubmed.
ncbi.nlm.nih.gov/19810200). S2CID 42790876 (https://api.semant
icscholar.org/CorpusID:42790876).
13. Cerling, T. E.; Levin, N. E.; Quade, J. (2010). "Comment on the
Paleoenvironment of Ardipithecus ramidus" (https://doi.org/10.11
26%2Fscience.1185274). Science. 328 (5982): 1105.
Bibcode:2010Sci...328.1105C (https://ui.adsabs.harvard.edu/abs/
2010Sci...328.1105C). doi:10.1126/science.1185274 (https://doi.o
rg/10.1126%2Fscience.1185274). PMID 20508112 (https://pubm
ed.ncbi.nlm.nih.gov/20508112).
14. Lovejoy, C. O.; Suwa, G.; Simpson, S. W.; Matternes, J. H.; White,
T. D. (2009). "The Great Divides: Ardipithecus ramidus Reveals
the Postcrania of Our Last Common Ancestors with African
Apes". Science. 326 (5949): 73–106.
Bibcode:2009Sci...326..100L (https://ui.adsabs.harvard.edu/abs/2
009Sci...326..100L). doi:10.1126/science.1175833 (https://doi.or
g/10.1126%2Fscience.1175833). PMID 19810199 (https://pubme
d.ncbi.nlm.nih.gov/19810199). S2CID 19629241 (https://api.sema
nticscholar.org/CorpusID:19629241).
15. Suwa, G.; Asfaw, B.; Kono, R. T.; Kubo, D.; Lovejoy, C. O.; White,
T. D.; et al. (2009). "The Ardipithecus ramidus skull and its
implications for hominid origins" (http://doc.rero.ch/record/211453/
files/PAL_E4442.pdf) (PDF). Science. 326 (5949): 68, 68e1–
68e7. Bibcode:2009Sci...326...68S (https://ui.adsabs.harvard.edu/
abs/2009Sci...326...68S). doi:10.1126/science.1175825 (https://d
oi.org/10.1126%2Fscience.1175825). PMID 19810194 (https://pu
bmed.ncbi.nlm.nih.gov/19810194). S2CID 19725410 (https://api.s
emanticscholar.org/CorpusID:19725410).
16. Suwa, G.; Kono, R. T.; Simpson, S. W.; Asfaw, B.; Lovejoy, C. O.;
White, T. D.; et al. (2009). "Paleobiological implications of the
Ardipithecus ramidus dentition" (http://doc.rero.ch/record/211460/f
iles/PAL_E4443.pdf) (PDF). Science. 326 (5949): 69, 94–99.
Bibcode:2009Sci...326...94S (https://ui.adsabs.harvard.edu/abs/2
009Sci...326...94S). doi:10.1126/science.1175824 (https://doi.org/
10.1126%2Fscience.1175824). PMID 19810195 (https://pubmed.
ncbi.nlm.nih.gov/19810195). S2CID 3744438 (https://api.semanti
cscholar.org/CorpusID:3744438).
17. Lovejoy, C. O.; Latimar, B.; Suwa, G.; Asfaw, B.; White, T. D.
(2011). "Combining Prehension and Propulsion: The Foot of
Ardipithecus ramidus" (http://doc.rero.ch/record/211468/files/PAL
_E4445.pdf) (PDF). Science. 326 (5949): 72–72e8.
doi:10.1126/science.1175832 (https://doi.org/10.1126%2Fscienc
e.1175832). PMID 19810198 (https://pubmed.ncbi.nlm.nih.gov/19
810198). S2CID 26778544 (https://api.semanticscholar.org/Corpu
sID:26778544).
18. Lovejoy, C. O.; Simpson, S. W.; White, T. D.; Asfaw, B.; Suwa, G.
(2009). "Careful Climbing in the Miocene: The Forelimbs of
Ardipithecus ramidus and Humans Are Primitive" (http://doc.rero.c
h/record/211452/files/PAL_E4441.pdf) (PDF). Science. 326
(5949): 70–70e8. Bibcode:2009Sci...326...70L (https://ui.adsabs.h
arvard.edu/abs/2009Sci...326...70L).
doi:10.1126/science.1175827 (https://doi.org/10.1126%2Fscienc
e.1175827). PMID 19810196 (https://pubmed.ncbi.nlm.nih.gov/19
810196). S2CID 37251630 (https://api.semanticscholar.org/Corpu
sID:37251630).
19. Lovejoy, C. O.; Suwa, G.; Spurlock, L.; Asfaw, B.; White, T. D.
(2009). "The Pelvis and Femur of Ardipithecus ramidus: The
Emergence of Upright Walking" (http://doc.rero.ch/record/211462/f
iles/PAL_E4444.pdf) (PDF). Science. 326 (5949): 71–71e6.
Bibcode:2009Sci...326...71L (https://ui.adsabs.harvard.edu/abs/2
009Sci...326...71L). doi:10.1126/science.1175831 (https://doi.org/
10.1126%2Fscience.1175831). PMID 19810197 (https://pubmed.
ncbi.nlm.nih.gov/19810197). S2CID 19505251 (https://api.semant
icscholar.org/CorpusID:19505251).
20. Prang, T. C. (2019). "The African ape-like foot of Ardipithecus
ramidus and its implications for the origin of bipedalism" (https://w
ww.ncbi.nlm.nih.gov/pmc/articles/PMC6491036). eLife. 8:
e44433. doi:10.7554/eLife.44433 (https://doi.org/10.7554%2FeLif
e.44433). PMC 6491036 (https://www.ncbi.nlm.nih.gov/pmc/articl
es/PMC6491036). PMID 31038121 (https://pubmed.ncbi.nlm.nih.
gov/31038121).
21. Simpson, S. W.; Levin, N. E.; Quade, J.; Rogers, M. J.; Semaw, S.
(2019). "Ardipithecus ramidus postcrania from the Gona Project
area, Afar Regional State, Ethiopia". Journal of Human Evolution.
129: 1–45. doi:10.1016/j.jhevol.2018.12.005 (https://doi.org/10.10
16%2Fj.jhevol.2018.12.005). PMID 30904038 (https://pubmed.nc
bi.nlm.nih.gov/30904038).
22. Clark, Gary; Henneberg, Maciej (2015). "The life history of
Ardipithecus ramidus: A heterochronic model of sexual and social
maturation" (https://doi.org/10.1515%2Fanre-2015-0009).
Anthropological Review. 78 (2): 109–132. doi:10.1515/anre-
2015-0009 (https://doi.org/10.1515%2Fanre-2015-0009).
23. Clark, G.; Henneberg, M. (2017). "Ardipithecus ramidus and the
evolution of language and singing: An early origin for hominin
vocal capability". Homo. 68 (2): 101–121.
doi:10.1016/j.jchb.2017.03.001 (https://doi.org/10.1016%2Fj.jchb.
2017.03.001). PMID 28363458 (https://pubmed.ncbi.nlm.nih.gov/
28363458).
24. Stanford, C. B. (2012). "Chimpanzees and the Behavior of
Ardipithecus ramidus". Annual Review of Anthropology. 41: 139–
149. doi:10.1146/annurev-anthro-092611-145724 (https://doi.org/
10.1146%2Fannurev-anthro-092611-145724).
25. Boisserie, J.-R. (2010). "Ardipithecus ramidus and the birth of
humanity" (https://www.persee.fr/doc/ethio_0066-2127_2010_nu
m_25_1_1420). Annales d'Éthiopie. 25: 271–281.
doi:10.3406/ethio.2010.1420 (https://doi.org/10.3406%2Fethio.20
10.1420).
26. Sayers, K.; Lovejoy, C. O. (2014). "Blood, Bulbs, and Bunodonts:
On Evolutionary Ecology and the Diets of Ardipithecus,
Australopithecus, and Early Homo" (https://www.ncbi.nlm.nih.gov/
pmc/articles/PMC4350785). Quarterly Review of Biology. 89 (4):
319–357. doi:10.1086/678568 (https://doi.org/10.1086%2F67856
8). PMC 4350785 (https://www.ncbi.nlm.nih.gov/pmc/articles/PM
C4350785). PMID 25510078 (https://pubmed.ncbi.nlm.nih.gov/25
510078).
27. White, T. D.; et al. (2009). "Macrovertebrate Paleontology and the
Pliocene Habitat of Ardipithecus ramidus" (http://doc.rero.ch/recor
d/211327/files/PAL_E4424.pdf) (PDF). Science. 326 (5949): 67–
93. doi:10.1126/science.1175822 (https://doi.org/10.1126%2Fsci
ence.1175822). PMID 19810193 (https://pubmed.ncbi.nlm.nih.go
v/19810193). S2CID 14837552 (https://api.semanticscholar.org/C
orpusID:14837552).
28. Gani, M. R.; Gani, N. D. (2011). "River-margin habitat of
Ardipithecus ramidus at Aramis, Ethiopia 4.4 million years ago" (h
ttps://doi.org/10.1038%2Fncomms1610). Nature
Communications. 2: 602. Bibcode:2011NatCo...2..602G (https://u
i.adsabs.harvard.edu/abs/2011NatCo...2..602G).
doi:10.1038/ncomms1610 (https://doi.org/10.1038%2Fncomms16
10). PMID 22186898
(https://pubmed.ncbi.nlm.nih.gov/22186898).
29. Louchart, A.; Wesselman, H.; Blumenschine, R. J.; et al. (2009).
"Taphonomic, Avian, and Small-Vertebrate Indicators of
Ardipithecus ramidus Habitat". Science. 326 (5949): 66–66e4.
Bibcode:2009Sci...326...66L (https://ui.adsabs.harvard.edu/abs/2
009Sci...326...66L). doi:10.1126/science.1175823 (https://doi.org/
10.1126%2Fscience.1175823). PMID 19810192 (https://pubmed.
ncbi.nlm.nih.gov/19810192). S2CID 13825029 (https://api.semant
icscholar.org/CorpusID:13825029).
30. WoldeGabriel, G.; Ambrose, S. H.; Barboni, D.; et al. (2009). "The
Geological, Isotopic, Botanical, Invertebrate, and Lower
Vertebrate Surroundings of Ardipithecus ramidus". Science. 326
(5949): 65–65e5. Bibcode:2009Sci...326...65W (https://ui.adsabs.
harvard.edu/abs/2009Sci...326...65W).
doi:10.1126/science.1175817 (https://doi.org/10.1126%2Fscienc
e.1175817). OSTI 971297 (https://www.osti.gov/biblio/971297).
PMID 19810191 (https://pubmed.ncbi.nlm.nih.gov/19810191).
S2CID 11646395 (https://api.semanticscholar.org/CorpusID:1164
6395).
31. Levin, N. E.; Simpson, S. W.; Quade, J.; Cerling, T. E.; Frost, S. R.
(2008). "Herbivore enamel carbon isotopic composition and the
environmental context of Ardipithecus at Gona, Ethiopia". The
Geology of Early Humans in the Horn of Africa. 446. Geological
Society of America Special Paper. pp. 215–234.
doi:10.1130/2008.2446(10) (https://doi.org/10.1130%2F2008.244
6%2810%29). ISBN 9780813724461.

External links
Reconstruction (http://gurche.com/ardipithecus-ramidus) by John Gurche
The Smithsonian Institution's Human Origins Program (http://humanorigins.si.edu/evidence/human-fossils/species/ardipithecus-ramidus)
Discovering Ardi - Discovery Channel (https://web.archive.org/web/20130318211338/http://dsc.discovery.com/tv-shows/other-shows/vide
os/other-shows-discovering-ardi-videos.htm)
Human Timeline (Interactive) (http://humanorigins.si.edu/evidence/human-evolution-timeline-interactive) – Smithsonian, National
Museum of Natural History (August 2016).

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