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Friday, 27 April 2012

Pongo and 'Fossil Pongo'

"The Mystery Ape" of Chinese Paleontology
Tyler Stone recently posted a blog entry on the identity of the Himalayan "Migu" or Wildman as being a survival of the creature often called "Fossil Pongo" in the literature. Tyler's article uses the recently suggested name "Pongo hooijeri" for what proponents also call "The Mystery Ape", and the Cryptid we are speaking of is what Bernard Heuvelmans used to call "Le Petit Yeti" (The smaller kind of 'Yeti') The publication of the description for "Pongo hooijeri" is quoted in the archived article reprinted below. The complete story is even more complicated than this will imply on first reading. Pongo is the scientific name for the orangutan.

Wednesday, August 18, 2010

Jianshi--Earliest Extra-African Hominin, Further Evidence of Late Occurring Lufengpithecus or Fossil Orangutan?

Liu et al. (2010) have recently published a study of three supposedly hominin teeth from the Longgudong cave site in Jianshi county, Hubei. The teeth augment the sample of 4 hominin-like teeth collected in and around Jianshi in the 1970s that were initially attributed to an australopithecine (Gao 1975). The assessment by Liu et al. (2010) supplements the report in the 2004 monograph on Jianshi (Zheng 2004) published as the first in a series commissioned as part of the State Key Project of the 9th Five Year Plan – Origin of Early Humans and Environmental Background. The 2004 monograph is a comprehensive study of the geology, stratigraphy and paleontology of the fossil bearing site. In it the three hominin-like teeth, collected in 2000, were assigned to the Javanese genus and species Meganthropus paleojavanicus, based primarily on metric criteria.

The Jianshi site is further distinguished by the presence of a relatively large number of Gigantopithecus blacki teeth (28 in toto) and an extensive and diagnostic mammalian fauna. Study of the micro- and macro-mammalian fossils indicates a biochronological age slightly younger than the Longgupo site at Wushan, Chongqing (former Sichuan) or the Gigantopithecus cave site at Liucheng, Guangxi. Paleomagnetic research, however, suggests that the hominin-like material falls below the Reunion Normal Polarity Event or in excess of 2.15 mya. As such the Jianshi specimens would represent the earliest record of an extra-African human presence in all of Eurasia.

I recently reposted an essay written over a decade ago, derived from my 1994 Dissertation, which suggested that the original Jianshi molars could be favorably compared to fossil orangutan teeth previously collected in South China and Java. I have also written extensively on the Longgupo mandibular fragment and shown how it is compatible with an attribution to Lufengpithecus. Russ Ciochon, who along with Huang Wanpo, et al. (1995), originally described the Longgupo specimen as an early hominin, has since backtracked and conceded its non-hominin nature (Ciochon 2009). Given the possibility that the new Jianshi specimens may be mimicking a hominin morphotype I think it is essential to compare the new specimens not only to other hominins known from Africa, Europe and Asia, as has been done by Liu et al. (2010), but with other hominoids, including Lufengpithecus, known from southern China and Southeast Asia. In this regard Schwartz et al. (1994, 1995) published a study of hominoid dental remains from a variety of Pleistocene age sites in Vietnam. These included teeth ascribed to Gigantopithecus, Homo and Pongo. Amongst these teeth are some attributed to a new species, Pongo hooijeri, described as,

“Description and Diagnosis: Larger than most subspecies of fossil and living Pongo pygmaeus (see figs. 9-11), but differs from all known populations of P. pygmaeus in lacking significant crenulation on the occlusal surfaces of the molars and upper premolars, and on the basins of the lower premolars. Incisors are not known. Molar cusp disposition similar to that of P. pygmaeus, but the cusps themselves are puffier and more rounded occlusally as well as on their external slopes. The occlusal surfaces are thus more poorly defined, and the occlusal basins are more constricted. Discussion: The characters of the molars listed above distinguish these teeth very markedly from those of Pongo pygmaeus, and give them a very distinctive "gestalt." While it does appear that this new form belongs in the same general clade as the extant orangutan, it is by no means clear on the basis of current evidence that the two are very closely related. Pending better material, we feel a conservative interpretation is most appropriate, and have thus allocated this dentally distinctive primate to its own species of Pongo. It may well turn out, however, that the relationship between the two species is more distant than this implies, and it is possible that a separate generic designation for hooijeri will ultimately be warranted.”
Unfortunately only the holotype specimen, a lower M2, is figured. In its overall appearance it looks remarkably similar to M2’s from Lufeng and is analogous in features to the M1 from Longgupo. Its description and analysis is also consistent with previously known Lufengpithecus.

The two recently described molars from Jianshi, one upper and one lower, are extremely well worn and lack most diagnostic surface features. They, as well as the original set of four molars described from Jianshi, are relatively large but fall within the upper range of variation of the Lufeng sample of hominoid molars. In this regard the Longgupo specimen falls within the lower range of variation. Therefore in terms of dental metrics there is no reason to dismiss the Jianshi and Longgupo remains as potentially members of a single highly dimorphic species such as seen in Lufengpithecus.

Liu et al. (2010) use a bevy of geometric morphometric techniques to tease apart shape discriminations to sort the Jianshi teeth relative to a variety of hominin samples. In a cursory comparison of the new Jianshi molars with homologous teeth in Schwartz et als study it appears that the sort of comparisons made by Liu et al. (2010) should also be done with non-hominin taxa from contiguous regions of East and Southeast Asia.

Additionally, the Mohui and Chuifeng Cave sites in the Bubing Basin of southern Guangxi have produced an abundant early Pleistocene fauna with, respectively, 16 and 92 teeth assigned to Gigantopithecus blacki. Another set (n=17) of smaller teeth from Mohui have been tentatively assigned to Hominoidea (gen. et sp. indet.)(Wang et al. 2007). These are said to include molars similar to those from the Longgupo mandibular fragment (Huang et al., 1995). Wang et al. go on to say that this Mohui sample does not strongly resemble Pongo remains found in Wuyun Cave and other sites in China. In addition Chuifeng Cave has produced "24 relatively large teeth, with high crowned molars and a simple occlusal pattern, (that) differ from the low crowned and wrinkled molars of Pongo, as well as the very large teeth of G. blacki. These teeth are provisionally assigned to Hominoidea gen. et sp. indet." (Wang 2009) Unfortunately neither the Mohui nor Chuifeng specimens are figured in the pertinent articles. Another similar molar specimen has been reported from Sanhe Cave in Chongzuo, Guangxi (Jin et al. 2009). Given the fact that non-hominin, non-pongine teeth that have been mistaken for specimens of Homo are now found widespread throughout southern China and northern Vietnam, the possibility must be entertained that the Jianshi teeth also represent a non-hominin taxon.

Zhao et al. (2009) describe and discuss a set of orangutan-like teeth from the early Late Pleistocene (~110 kya) site of Mulanshan, Guangxi. They review similar specimens collected from throughout southern China and from Chinese drugstores since the 1930s. They reach similar conclusions as have others, that there are two distinct morphs within the sample. Specimens that are morphologically similar, but larger than, extant and subfossil orangutans from Indonesia (see also Wang et al. 2009) and those that are less crenulated with smoother, less complex occlusal surfaces. These latter specimens are said to be similar to Lufengpithecus but on average somewhat larger. Zhao et al. go on to say that there are also specimens within the Lufeng and Yuanmou hypodigms of Lufengpithecus that are similar to the more crenualted specimens collected from Pleistocene sites in southern China and northern Vietnam. They conclude that "In the absence of the skull and postcranial bones, it is difficult to establish new genus or species based on Pleistocene orangutan-like isolated teeth."

Thus, there are a number of possibilities, none of which are, as of yet, conclusive.

1. All the specimens discussed above including the Jianshi molars are variants of one or more species or subspecies of orangutan.

2. The specimens can be divided between a Pleistocene descendant of Lufengpithecus, fossil forms of Pongo or a previously unidentified "mystery ape".

3. Some of the specimens can be allocated to an early form of hominin, the rest to Pongo or some other combination of the above.

Below are some visual comparisons of relevant specimens:

Tham Hai Cave Upper RM1 or M2 (reversed)
"Immediately adjacent to Tham Khuyen, this cave was excavated toward the end of 1964 by a joint Vietnamese-German (DDR) team. Dating is uncertain, but the site is thought to be approximately contemporaneous with Tham Khuyen. A single tooth was attributed to Homo erectus (Cuong (sic) Nguyen, 1985). This is a relatively large upper right molar (BL width 15.41 mm; MDlength 13.47 mm), which is heavily worn. It almost certainly belongs to the "Morph 2" category of Schwartz et al. (1994), described below as a new species of Pongo." (Schwartz et al. 1995).

To the left is an upper right molar of Lufengpithecus hudienensis recovered from the Yuanmou Basin of Yunnan. The Yuanmou sites are dated between 7-8 mya. Note the similarity of the Yuanmou specimen's occlusal outline to the specimen of Homo erectus from Zhoukoudian (ZKD) shown below (specimens are not to scale).

Occlusal views of UM1s a) Jianshi b) Australopithecus c) ZKD H. erectus d) Sangiran H.erectus e) modern Chinese

Right Lower M1 or M2 (TK 65/123), Right Lower M1 or M2 (Jianshi PA 1277), Longgupo Right Lower M1 (not to scale).
Original Jianshi lower molars compared to Pongo lower molar from Tham Khuyen Cave Vietnam (center). Note median placement of longitudinal fissure and alignment of transverse fissures in all specimens (not to scale).
Original Jianshi lower molars compared to Lufengpithecus hudienensis from Yuanmou Basin, Yunnan and a specimen from Mohui originally described as "hominid" (not to scale).

Given the ambiguous nature of hominoid dentitions from the Mio-Plio-Pleistocene of East Asia the issues discussed above cannot be resolved until more diagnostic material is recovered.

Literature cited:

Ciochon, R. The mystery ape of Pleistocene Asia. 2009, Nature, 459:910-911.

Etler, DA. The Chinese Hominidae: New Finds, New Interpretations. 1994, PhD. Dissertation, University of California, Berkeley.

Etler, DA. Mystery ape: other fossils suggest that it's no mystery at all. 2009, Nature 460:684.

Etler, DA., Crummett, TL., Wolpoff, MH. Longgupo: Early Homo colonizer or Late Pliocene Lufengpithecus survivor in South China? 2001, Hum Evol. 16:1-12.

Gao, J. Australopithecine teeth associated with Gigantopithecus. Vertebrata PalAsiatica, 1975, 13(2):81-88.

Huang, W., Ciochon, R., Gu, Y., Larick, R., Fang, Q., Schwarcz, H., Yonge, C., de Vos, J. and Rink, W. Early Homo and associated artifacts from Asia. Nature, 1995, 378:275-78.

Jin, C., Qin D., Pan W., Tang Z., Liu, J., Wang, Y., Deng C., Zhang, Y., Dong, W., and Tong, H. A newly discovered Gigantopithecus fauna from Sanhe Cave, Chongzuo, Guangxi, South China, Chinese Science Bulletin, 2009, 54(5):788-797.

Liu W, Clarke R, Xing S. Geometric morphometric analysis of the early Pleistocene hominin teeth from Jianshi, Hubei Province, China. Sci China Earth Sci, 2010, 53: 1141–1152.

Schwartz, J. H., Vu, T. L., Nguyen, L. C., Le, T. K., and Tattersall, I. A diverse hominoid fauna from the late middle Pleistocene breccia cave of Tham Khuyen, Socialist Republic of Vietnam, 1994, Anthropologica Papers of the American Museum of Natural History, 73:1-11.

Schwartz, J. H., Vu, T. L., Nguyen, L. C., Le, T. K. A review of the Pleistocene hominoid fauna of the Socialist Republic of Vietnam (excluding Hylobatidae). Anthropological Papers of the American Museum of Natural History, 1995, 76:1-23.

Wang, W. New discoveries of Gigantopithecus blacki teeth from Chuifeng Cave in the Bubing Basin, Guangxi, south China. Journal of Human Evolution, 2009, 57:229–240.

Wang, W., Potts, R., Hou, Y., Chen, Y., Wu, H., Yuan B., and Huang, W. Early Pleistocene hominid teeth recovered in Mohui cave in Bubing Basin, Guangxi, South China, 2005, Chinese Science Bulletin, 50(23):2777—2782.

Wang, CB., Zhao, L., Jin, C., Hu, Y. and Wang, CS. Quantitative study of Pleistocene fossil orangutan teeth from China and their taxonomic significance. 2009, Acta Anthropologica Sinica 28(2):192-200.

Wang, W., Potts, R., Yuan, B., Huang, W., Cheng H., Edwards, R. L., Ditchfield, P. Sequence of mammalian fossils, including hominoid teeth, from the Bubing Basin caves, South China, 2007, Journal of Human Evolution 52:370-379.

Zhao, L., Wang, C., Jin, C., Qin, D. and Pan, W. Fossil Orangutan-like hominoid teeth from late Pleistocene human site of Mulanshan cave in Chongzuo of Guangxi and implications on taxonomy and evolution of orangutan, 2009, Chinese Sci Bull, 54: 3924―3930.

Zheng S H. Jianshi Hominid Site. Beijing: Science Press, 2004, 1–412.

Zhou, G. Supplementary notes on the baby skull of Sivapithecus yunnanensis -- with discussions on "Wushan Man." Memoirs of Beijing Natural History Museum, 1999, 58:111-123.

We have touched on these sample teeth before in an earlier posting discussing possible Australopithecines from the Orient and at that time I was perfectly happy to allow that the teeth were "Meganthropus" and that "Meganthropus" was most likely a form of Australopithecus/Paranthropus robustus. However the article does touch on the much larger issue and the one that impacts all current reports of Mainland-Asian apes as well, the general classification of the category "Fossil Pongo". Because in reality the "Fossil Pongo" is at the base of the whole "Mystery Ape" problem. Simply because, whatever it might be, it cannot be a standard orangutan (Pongo) in any way whatsoever. And, whatever else might be true, it also cannot be a late-surviving species of the general Sivapithecus or Lufengpithecus because those are Miocene genera and inappropriate for a Pleistocene species. Ordinarily even if we are dealing with a lazarus taxon, The same genus name is not used for two distinct species which lived ten million years apart (or more) and had no fossil material representative of the intervening time.

There is a consistent problem in that researchers will speak of the fossil mainland Asian apes as being "Pongo" and even as being an unending parade of new subspecies for Pongo pygmaeus, the living Sumatran orangutan, solely on the basis of dental similarity to the living orangutans. As mentioned in the article above, the dental anatomy of orangutans and possibly quite unrelated fossil apes, and even early humans is similar enough that distinguishing the species is rather hazardous. There is an exellent review of the fossil material in the document which originally named "Pongo hooijeri" but it also uses the default of calling all the fossils subspecies of "Pongo pygmaeus", and then goes on to name four new subspecies on the basis of variations in the teeth. The largest of these teeth are easily twice the size as the average teeth of the living orangutans. Clearly, something is very wrong with the classification. See the pdf:

I have consistently been one in the minority who have pointed out that if these "Fossil Pongo" creatures were as large as gorillas and lived entirely upon the ground (since they were much too large to live in the trees as do regular orangutans), then they simply are not orangutans at all, no matter what their teeth look like. From the size of the teeth, we are talking about creatures that could commonly weigh 200 to 400 pounds in life: the females are at least as large as a modern male orangutan, and the males could probably regularly stand over six feet tall when upright. Their joints, musculature, tendons and the shape of their hands and feet must all have differed from modern orangutans and must have been used consistently in different ways owing to the different lifestyle. And even if you do make a special allowance for another species of Pongo under the name hooijeri, there is no way that you can consider the other fossil types as belonging to the same species as modern orangutans (counting both Bornean and Sumatran orangutans as the same species) Moreover, there is also no way that they can even be included in the same GENUS. The modern orangutan is an arboreal specialist with a number of peculiar specializations adapting it for life in the trees. Take away all of those specialized adaptations abnd you simply are not talking orangutans any more.

There shall be another part following this one in order to deal with the relationships of "Fossil Pongo" to the related (but much earlier) Lufengpithecus and Sivapithecus. But for right now there is also another article which touches on the idea of a "Mystery ape" assumed to be the same as this "Pongo hooijeri" and discusses th idea that the fossils had Australopithecine affinities:

Nature 459, 910-911 (18 June 2009) | doi:10.1038/459910a; Published online 17 June 2009

The mystery ape of Pleistocene Asia

Russell L. Ciochon1
  1. Russell L. Ciochon is chair of anthropology at the University of Iowa, Iowa City, Iowa 52242, USA. This Essay is based on a contribution to the book Out of Africa I: Who, When and Where? (eds, Fleagle, J. G. et al. Springer, 2009).
Fossil finds of early humans in southeast Asia may actually be the remains of an unknown ape. Russell Ciochon says that many palaeoanthropologists — including himself — have been mistaken.
Fourteen years ago, a Nature paper by my colleagues and I described a 1.9-million-year-old human jaw fragment from Longgupo in Sichuan province, China1. The ancient date in itself was spectacular. Previous evidence had suggested that human ancestors arrived in east Asia from Africa about 1 million years ago, in the form of Homo erectus. Longgupo nearly doubled that estimate. But even more exciting — and contentious — was our claim that the jaw was related to H. habilis, a species of distinctly African origin. If this descendant of H. habilis had arrived so early into southeast Asia, then it probably gave rise to H. erectus in the Far East, rather than H. erectus itself sweeping west to east.
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.
The Longgupo site, discovered in 1984, lies 20 kilometres south of the Yangtze River in eastern Sichuan. At the beginning of the Pleistocene (1.8 million to 10,000 years ago) this cave sat near the northern range of a subtropical forest as rich with life as any in contemporary Africa. Unsurprisingly, the mammalian fossils dug up from Longgupo belonged to the subtropical StegodonAiluropoda fauna found throughout the subtropical forested region south of China's Qinling Mountains (see map). The name comes from two common members — the extinct elephant-like Stegodon and the bear-like giant panda, Ailuropoda. It includes primates such as the extinct giant ape Gigantopithecus, as well as the ancestors of the living orangutan (Pongo) and gibbon (Hylobates).
The mystery ape of Pleistocene Asia
But Longgupo also yielded a mystery jaw fragment, including the fourth premolar and the first molar. Although obviously primate, the worn enamel surfaces made precise classification difficult. Some had called it an ape whereas others saw an early human. In 1992, colleagues and I were invited to Longgupo to provide a reliable age determination and to help understand the palaeoanthropology.
The 1.8-million–2-million-year-old jaw was smaller than that of any known orangutan, living or extinct. We also compared it with primate dental fossils from the site of Lufeng, in neighbouring Yunnan province. Lufengpithecus was of the right size and general morphology, but the age was wrong: Lufeng and similar sites belonged to the late Miocene period, about 7 million–9 million years ago. Some possible stone tools found at the site seemed to support a human classification. Asian H. erectus was the obvious possibility, but the size, tooth proportions and root structure were not quite right. Dissatisfied with the usual regional comparisons, we looked to Longgupo's possible links with early African humans such as H. habilis, whose Great Rift Valley fossils are as old as 2.3 million years. Our Nature announcement1 thus presented the Longgupo jaw as a newcomer to the StegodonAiluropoda fauna: an African hominin more primitive than H. erectus.

Pre-erectus claims

We weren't the first or last to suggest that a pre-erectus African hominin migrated to east Asia. In the 1940s and 1950s pre-erectus African claims were made for fossils from Sangiran, on Java, Indonesia. Early in Sangiran's long history of H. erectus discoveries, a couple of massive jaws seemed similar to those of South African australopiths — they were coined 'Meganthropus'. But as more fossils were discovered at Sangiran, it became clear that the Meganthropus jaws were merely a local variant of H. erectus.
More than a decade later, with some distance from the subject, the teeth looked distinctly more ape-like.
Just this year, claims for a pre-erectus African in Asia have also surfaced to explain the evolution of Indonesia's Homo floresiensis, popularly known as the Liang Bua 'hobbit'. Discovered in 2003, and dated to just 95,000 to 17,000 years ago, the Liang Bua skeleton is a diminutive species significantly different from all other known humans. The discoverers proposed that the diminutive H. floresiensis evolved from a southeast Asian H. erectus group that became isolated on Flores: faced with limited resources, the erectus group dwarfed to match the small-island conditions. However, recent studies of Liang Bua wrist and foot bones reveal primitive anatomies reminiscent of H. habilis or Australopithecus, again leading some to propose a pre-erectus African origin for the species. The problem is that no comparable wrist or foot bones are known for H. erectus, making it impossible at this time to exclude a local variant of H. erectus as the ancestor of the Liang Bua 'hobbit'.
So our claim of a pre-erectus African hominin living in east Asia fell into a long line of such arguments. It was met with healthy scepticism. We were first faced with the response that Longgupo was an orangutan, but we were able to show that the two teeth lay significantly outside the orangutan range of variation2, 3. Later, we had to field a serious proposal that Longgupo belonged to Lufengpithecus4, 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.

Then in spring 2005, I met with Wang Wei, director of the Guangxi Natural History Museum, to examine his collection of 33 primate teeth from Mohui cave in Bubing Basin, south China. Wang's excavations produced an excellent sample of the StegodonAiluropoda fauna. Quickly I could see that 15 teeth were those of Gigantopithecus, and 10 were probably Pongo. The remaining eight specimens did not fit with any known east Asian Pleistocene primate.
Some 15 years earlier, I had worked hard to show that Gigantopithecus had crossed paths with H. erectus; I wrote a book in 1990 proposing this relationship (Other Origins: The Search for the Giant Ape in Human Prehistory) and a few years later had documented evidence of the species' co-existence. In my mind the two were firmly linked. But more than a decade after the discovery, with some distance from the subject, the teeth in Wang's lab looked distinctly more ape-like than hominin.

Teething problems

Without the assumption that Gigantopithecus and H. erectus lived together, everything changed: if early humans were not part of the StegodonAiluropoda fauna, I had to envision a chimpanzee-sized ape in its place — either a descendant of Lufengpithecus, or a previously unknown ape genus. The Mohui mystery teeth surely belonged to an unknown ape, as did Longgupo, and other human-like teeth often identified from similar cave fossils. Although I no longer consider the Longgupo jaw to be human, the two stone tools still stand as described. They must have been more recent additions to the site.
The mystery ape concept is bolstered by looking at definitive H. erectus finds in east Asia. Our knowledge comes mainly from two sites: Zhoukoudian near Beijing, which lies well north of the primal forest, and Sangiran in Java, which lies well south of it. Each site represents hundreds of thousands of years of H. erectus occupation: Sangiran beginning as early as 1.6 million years ago, Zhoukoudian beginning about 780,000 years ago6. Neither site preserves StegodonAiluropoda fauna or mystery ape teeth. 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 conclusions7; a 2009 paper identifies a tooth from Sanhe Cave (Chongzuo, Guangxi province, China) as belonging to an unidentified ape8.
In this call to reassess historical assemblages, it is worth remembering the story of 'Hemanthropus'. Legendary fossil collector Ralph von Koenigswald created this hominin taxon in 1957, based on isolated fossil teeth found in apothecary shops across southeast Asia. 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.
The question remains: is there only one mystery ape or possibly more? It seems that there was as much diversity of apes in the southeast Asian Pleistocene as in Africa today. In modern Africa there is one large ape (the gorilla) and two smaller apes (the chimpanzee and bonobo); in Asia during the Pleistocene and recent times, we have one very large ape (Gigantopithecus), one large ape (the orangutan), at least one smaller ape (mystery ape) and finally a tiny ape (the gibbon).
The next step is to consider the mystery ape fossils as a group and see how they fit into the evolutionary history of the range of southeast Asian apes. Wang will head up this team effort, along with Chinese and international colleagues, including myself. Museum collections holding potential mystery-ape evidence will be examined, including those in Hanoi, Jianshi, Beijing and Frankfurt. Wang's ongoing excavations at cave sites in Guangxi's Bubing Basin are yielding new evidence with every passing day. Possibly, there will be a chance to announce a new southeast Asian fossil ape in some future issue of this journal.
Further reading
Bettis, E. A. III et al. J. Hum. Evol. 56, 11-24 (2009).
Ciochon, R. L. et al. Proc. Natl Acad. Sci. USA 93, 3016-3020 (1996).
Colbert, E. H. & Hooijer, D. R. Bull. Amer. Mus. Nat. Hist. 102, 1-134 (1953).
Gao, J. Vertebrata PalAsiatica 13, 81-88 (1975).
Jungers, W. L. et al. Nature 459, 81-84 (2009).
Li, H., Yang, X., Heller, F. & Li, H. Quat. Res. 69, 250-262 (2008).
Lieberman, D. E. Nature 459, 41-42 (2009).
Pei, W.-C. Vertebrata Palasiatica 1, 9-24 (1957).
Rink, W. J., Wei, W., Bekken, D. & Jones, H. L. Quat. Res. 69, 377-387 (2008).
Shen, G., Gao, X., Gao, B. & Granger, D. E. Nature 458, 198-200 (2009).
Tocheri, M. W. et al. Science 317, 1743-1745 (2007).
Wang, W. et al. J. Hum. Evol. 52, 370-379 (2007).
Zhang, Y. Acta Anthropol. Sin. 3, 85-92 (1984).
Zheng, S. H. Jianshi Hominid Site (Beijing Science Press, 2004).
See also News, page 899.


  1. Huang, W. et al. Nature 378, 275–278 (1995). | Article | PubMed | ISI | ChemPort |
  2. Schwartz, J. H. & Tattersall, I. Nature 381, 201–202 (1996). | Article | PubMed | ISI | ChemPort |
  3. Huang, W. et al. Nature 381, 202 (1996). | Article |
  4. Wu, X. Acta Anthropol. Sin. 19, 1–10 (2000).
  5. Etler, D. A. et al. Hum. Evol. 16, 1–12 (2001). | Article |
  6. Ciochon, R. L. & Bettis, E. A. III Nature 458, 153–154 (2009). | Article | PubMed | ChemPort |
  7. Schwartz, J. H. et al. Anthropol. Pap. Amer. Mus. Nat. Hist. 76, 1–24 (1995).
  8. Jin, C. et al. Chinese Sci. Bull. 54, 788–797 (2009). | Article |

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