Morphometric affinities of Gigantopithecus.
AbstractMultivariate analyses, supplemented by univariate statistical methods, of measurements from mandibular tooth crown dimensions and the mandible of Gigantopithecus blacki, G. bilaspurensis, Plio-Plelstocene hominids, Homo erectus, and seven Neogene ape species from the genera Proconsul, Sivapithecus, Ouranopithecus, and Dryopithecus were used to assess the morphometric affinities of Gigantopithecus. The results show that Gigantopithecus displays affinities to Ouranopithecus and to the hominids, particularly the Plio-Plelstocene hominids, rather than to the apes. Ouranopithecus demonstrated dental resemblances to G. bilaspurensis and the Plio-Pleistocene hominids but mandibular similarities to the apes. Results of analyses of tooth and mandibular shape indices, combined with multivariate distance and temporal relationships, suggest that Ouranopithecus is a more likely candidate for Gigantopithecus ancestry than is Silvapithecus indicus. Shape and allometric differences between G. bilaspurensis and the robust australopithecines weaken the argument for an ancestral-descendant relationship between these groups. The results support the hypothesis that Gigantopithecus is an extinct side branch of the Hominidae.[End of Abstract]
Copy of an illustration of male and female Ouranopithecus jaws, both looking remarkably similar to the corresponding Gigantopithecus jaws. Unfortunately, this is the best copy of the illustration I can get at present.
Some of the features of Ouranopithecus are similar to hominids. Most notably, like some other Late Miocene remains, the canines are relatively smaller in size than in many apes, especially in females. With fairly thick molar enamel and low cusps, the molar teeth are not gorilla-like, but instead are more similar to later hominids. Sexual dimorphism is substantial between the teeth, however, and males have large canines and shearing lower premolars.
Miocene ape review, in brief
Fri, 2010-01-29 19:58 -- John Hawks
I've been trying to give a longer account of Miocene ape evolution here, by dribs and drabs. You can get most of the list under my "Miocene" category. Last fall's account of "Late Miocene apes from Africa" expands significantly on a topic that Harrison glosses -- what do we know about African apes during the time that chimpanzees, gorillas, and humans were diverging?
There are two phylogenetic issues related to Miocene apes that command a lot of attention. Where did hominins come from? -- that's one issue. The other is, how are Asian and African apes related?
This latter question is important because it helps to establish the timeline of orangutan divergence from us and subsequent evolution. It also determines the position of the European apes -- and thereby, whether their locomotor and dietary diversity is relevant to later ape evolution.
Harrison comes down in favor of the hypothesis that Dryopithecus and most other European apes were stem hominids -- that is, collaterals of all living great apes, not specially related to the African apes and humans. The main alternative to that view is that the European apes mostly represent the African side of an early Asia-Africa biogeographic split, such that chimpanzees, gorillas and humans (the hominines) descend from Dryopithecus or some similar lineage closely aligned with these European apes. Harrison is willing to admit the Late Miocene Greek ape Ouranopithecus to the African ape lineage, but in his view the earlier European apes belong to one or more side-branches of the great apes.
A question: If Harrison is correct here, does that affect the paleontological evidence for the human-orangutan divergence? That is, if Dryopithecus were a hominine, it is plausible that the Asian/African ape divergence actually happened in Eurasia not long before 12 million years ago. If Dryopithecus branched off before the African and Asian ape divergence, and Sivapithecus was derived from an African ape, does that make the divergence earlier or later?
I'll consider that over the weekend.
The mystery ape from Longgupo
For many years, I used Longgupo to promote this pre-erectus origin for H. erectus finds in Asia. But now, in light of new evidence from across southeast Asia and after a decade of my own field research in Java, I have changed my mind. Not everyone may agree; such classifications are always open to interpretation. But I am now convinced that the Longgupo fossil and others like it do not represent a pre-erectus human, but rather one or more mystery apes indigenous to southeast Asia's Pleistocene primal forest. In contrast, H. erectus arrived in Asia about 1.6 million years ago, but steered clear of the forest in pursuit of grassland game. There was no pre-erectus species in southeast Asia after all.I think it's interesting how much speculation Nature is willing to publish about hominid evolution in Asia. The 2005 review article by Robin Dennell and Wil Roebroeks, "An Asian perspective on early human dispersal from Africa," speculated that the origin and early evolution of Homo may have been in Asia, not Africa. And of course, several papers on the hobbits have included speculations about the pattern of early Homo in Asia, in pursuit of ways to derive Homo floresiensis from early hominids not yet found in Asia.
Ciochon's essay is part of this new tradition, but it bucks the trend. Instead of arguing that Asia was the home to an undiscovered diversity of hominids, he instead argues that the hominids have been overestimated (in part by himself) and that some fossils represent an undiscovered diversity of apes.
Ancient orangutans (Pongo) and Gigantopithecus are already known from China. Ciochon proposes a third lineage of great ape, one that would be similar to the earlier Lufengpithecus from China and Thailand:
Later, we had to field a serious proposal that Longgupo belonged to Lufengpithecus (4, 5). Although the age disparity remained troubling, the dental similarities could not be denied. I began to imagine a mystery ape as a possible solution to the problem.The "age disparity" Ciochon refers to is that Lufengpithecus is known from the Late Miocene and very earliest Pliocene, but not the Late Pliocene. Still, if the teeth look like Lufengpithecus, it seems probable that the "mystery ape" actually is a late-surviving Lufengpithecus, or at least a close relative. Reference 5 is a paper by Dennis Etler, Tracy Crummett and Milford Wolpoff, which is available (PDF) from Etler's excellent website. Wolpoff refers to this in his 1999 book, Paleoanthropology:
The Longgupo mandible is actually a fossil ape that is related to Lufengpithecus, the missing P3 was sectorial in shape. However the prestigious British Journal Nature hastily published it as a hominid, with a picture of the specimen on its cover, and subsequently refused to accept papers establishing its identity. The misidentifications actually started decades ago, when G.H.R. von Koenigswald identified an ancient australopithecine-like hominid from South China based on worn, isolated teeth, which he named Hemianthropus. These turned out to be worn postcanine teeth of a medium-sized Pongo species. The resemblances of the other materials to Australopithecus species were real enough, but they were not unique resemblances. A. Kramer and Zhang Yinyun have each shown there are no synapomorphies that support the hypothesis of Asian australopithecines.So it's not a new idea that Longgupo represents an ape, or that the ape was different in size and morphology from Pongo or Gigantopithecus. It is probably natural that early paleontologists might associate these ape teeth with the hominids -- until 40 years ago, most paleontologists thought that hominids went back far into the Miocene. They were wrong, but a mistake like "Hemianthropus" was a natural one. The opposite mistake -- "Meganthropus" as an australopithecine-like hominid -- was also a natural consequence of the assumption that an unrecognized hominid diversity existed in Asia. That assumption has outlived Meganthropus, as we've seen.
Ciochon adds the idea that the ape may also be represented at other contemporary or later sites, and is apparently unwilling to attribute them to Lufengpithecus, at least not yet. He does not mention the isolated upper incisor from Longgupo, but he does appear to accept the claim that the two stone artifacts from the site are intrusive elements that are not contemporary with the jaw. The same is probably true of the incisor, which Etler and colleagues found morphologically most like living East Asians.
Ciochon suggests that some of the Hemianthropus collection may be his mystery ape:
Von Koenigswald viewed Hemanthropus as a distant relative of African Australopithecus. Later research revealed that these were worn or atypical orangutan teeth and Hemanthropus was quickly abandoned. But, had von Koenigswald actually discovered evidence of the mystery ape? In October 2005, I examined the original Hemanthropus collection. Among the many worn orangutan teeth I found several small ape teeth that very closely resembled the mystery ape teeth from Mohui. Perhaps von Koenigswald was the first to lay hands on the mystery ape.It's not an easy task to sort through large samples of teeth trying to sort them into sets. Particularly not with these teeth -- sure, Gigantopithecus falls right out, but worn orangutan teeth aren't very easy to tell from hominids, much less "mystery apes." Ciochon ends his essay with a plan to revisit the existing samples of teeth, trying to document the variation in the mystery lineage. Sounds like a good topic for a TV show. There's a historical angle, lots of museums, a personal hook, reversal of fortune, the whole "mystery ape" thing....
Meanwhile, the introductory paragraph at the top of the post raised two issues, not one. The first is the focus of the rest of the essay: Longgupo represents a third ape in Pleistocene China; smaller than both Gigantopithecus and Pongo. The second idea is covered briefly near the end of the essay, but I think it deserves more consideration. Is it true that humans reached China 1.6 million years ago, and then "steered clear of the forest"?
Here's what Ciochon writes:
Homo erectus, it seems from this perspective, hunted grazing mammals on open grasslands, and did not or could not penetrate the dense subtropical forest. In fact, there is no record of early hominins living in tropical or subtropical forested environments in Africa or Asia.
In resolving the mystery, two other Asian sites come to mind: Jianshi (Hubei province, China) and Tham Khuyen (Lang Son province, Vietnam). At both sites, teeth labelled variously as Australopithecus, H. erectus and Meganthropus are most likely to be the mystery ape instead. Others have come to similar conclusions; a 2009 paper identifies a tooth from Sanhe Cave (Chongzuo, Guangxi province, China) as belonging to an unidentified ape.The map accompanying the article is mysteriously depauperate of actual early hominid sites in China. Considering their locations relative to the proposed distribution of subtropical forest in Pleistocene China, I don't see an immediate objection to the hypothesis. The earliest Chinese archaeological sites, from the Nihewan basin near Beijing (Majuangou and Xiaochangliang) and also from around the Yellow River (Gongwangling and Xihoudu), are north of the Stegodon--Ailuropoda fauna. Yuanmou may have been forested at this time, but the hominid teeth there appear to be later (Hyodo et al. 2002), when the Ciochon's forest-plains biogeographic proposal may no longer hold. Josette Sarel and colleagues (2009) report on stone tools from Baerya Cave, which does preserve the Stegodon--Ailuropoda fauna, but these are so far undated and the stratigraphy has not been worked out. For all we know, the association is no clearer than at Longgupo, but that may change.
The other early Chinese sites with hominid teeth, Ciochon suggests are not hominids -- Mohui and Sanhe. Since he has examined the Mohui teeth (Wang et al. 2007), this isn't an idle speculation, and it would be odd for humans to drop their teeth around these sites without dropping a single stone tool. If he's right, that would make the earliest clear evidence of human occupation of South China into the Middle Pleistocene in age.
So, it's an interesting generalization. It remains to be seen how true it may be -- was early Homo really limited to a biogeographic strip of plains and savanna as it left Africa, or were the early humans more broadly adapted -- or adaptable?
ca 15-14.5 Million years ago, dispersal of the Proconsuls and early apes Out Of Africa. early apes are most similar to macaque monkeys in size and appearance but they already have the standard types of molar teeth that apes and humans still have.
13 to 14 Million years ago approximately. Dryopithecine radiation.Sivapithecine apes make a first appearance in the Siliwak deposits of India
11 to 12.5 million years ago according to the stage indicated. Apes with more recognisably ape-shaped bodies and adaptations begin to spread, including Sivapithecus and its allies.
9 to 10.5 million years ago
This would represent Ouranopithecus' age except that it persisted longer than the indicated date. The majority of apelike creatures are dying out in Europe and the Near East, becoming established in tropical Asia and Africa instead.
Middle Miocene dispersals of apes.
SourceDepartment of Palaeontology, Natural History Museum, London, UK. firstname.lastname@example.org <email@example.com>
AbstractThe earliest record of fossil apes outside Africa is in the latest early Miocene of Turkey and eastern Europe. There were at least 2, and perhaps 4, species of ape, which were found associated with subtropical mixed environments of forest and more open woodland. Postcranial morphology is similar to that of early Miocene primates and indicates mainly generalized arboreal quadrupedal behaviours similar to those of less specialized New World monkeys such as Cebus. Robust jaws and thick enamelled teeth indicate a hard fruit diet. The 2 best known species of fossil ape are known from the site of Paşalar in Turkey. They have almost identical molar and jaw morphology. Molar morphology is also similar to that of specimens from Germany and Slovakia, but there are significant differences in the anterior teeth of the 2 Paşalar species. The more common species, Griphopithecus alpani, shares mainly primitive characters with early and middle Miocene apes in Africa, and it is most similar phenetically to Equatorius africanus from Maboko Island and Kipsaramon. The second species is assigned to a new species of Kenyapithecus, an African genus from Fort Ternan in Kenya, on the basis of a number of shared derived characters of the anterior dentition, and it is considered likely that there is a phylogenetic link between them. The African sites all date from the middle Miocene, similar in age to the Turkish and European ones, and the earliest emigration of apes from Africa coincides with the closure of the Tethys Sea preceding the Langhian transgression. Environments indicated for the African sites are mixtures of seasonal woodlands with some forest vegetation. The postcrania of both African taxa again indicate generalized arboreal adaptation but lacking specialized arboreal function. This middle Miocene radiation of both African and non-African apes was preceded by a radiation of arboreal catarrhine primates in the early Miocene, among which were the earliest apes. The earliest Miocene apes in the genus Proconsul and Rangwapithecus were arboreal, and because of their association with the fruits of evergreen rain forest plants at Mfwangano Island, it would appear that they were forest adapted, i.e. were living in multi-storied evergreen forest. The same or similar species of the same genera from Rusinga Island, together with other genera such as Nyanzapithecus and the small ape Limnopithecus, were associated with plants and animals indicating seasonal woodland environments, probably with gallery forest forming corridors alongside rivers. While the stem ancestors of the Hominoidea were almost certainly forest adapted, the evidence of environments associated with apes in the later part of the early Miocene and the middle Miocene of East Africa indicates more seasonal woodlands, similar to those reconstructed for the middle Miocene of Paşalar in Turkey. This environmental shift was probably a requisite for the successful emigration of apes out of Africa and made possible later movement between the continents for much of the middle Miocene, including possible re-entry of at least one ape lineage back into Africa.
Copyright 2007 S. Karger AG, Basel.
A new Late Miocene great ape from Kenya and its implications for the origins of African great apes and humans
- Yutaka Kunimatsua,b,
- Masato Nakatsukasac,
- Yoshihiro Sawadad,
- Tetsuya Sakaid,
- Masayuki Hyodoe,
- Hironobu Hyodof,
- Tetsumaru Itayaf,
- Hideo Nakayag,
- Haruo Saegusah,
- Arnaud Mazurieri,
- Mototaka Saneyoshij,
- Hiroshi Tsujikawak,
- Ayumi Yamamotoa, and
- Emma Mbual
- bTo whom correspondence should be addressed. E-mail: firstname.lastname@example.org
- Author contributions: Y.K., M.N., Y.S., and E.M. designed research; Y.K., M.N., Y.S., T.S., M.H., H.H., T.I., H.N., A.M., M.S., H.T., and A.Y. performed research; Y.K., Y.S., T.S., M.H., H.H., T.I., H.N., H.S., A.M., M.S., H.T., and A.Y. analyzed data; and Y.K., M.N., Y.S., and H.N. wrote the paper.
- The authors declare no conflict of interest.
- This article is a PNAS Direct Submission.
- This article contains supporting information online at www.pnas.org/cgi/content/full/0706190104/DC1.
- © 2007 by The National Academy of Sciences of the USA