The mystery ape from Longgupo

In last week's Nature, Russell Ciochon has a remarkable essay:

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?

References:

Ciochon RL. 2009. The mystery ape of Pleistocene Asia. Nature 459:910:911.
doi:10.1038/459910a

Ciochon R, Long VT, Larick R, González L, Grün R, de Vos J, Yonge C, Taylor L,

Yoshida H, Reagan M. 1996. Dated co-occurrence of Homo erectus and
Gigantopithecus from Tham Khuyen Cave, Vietnam. Proc Nat Acad Sci 93:3016-3020.

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

Hyodo M, Nakaya H, Urabe A, Sagua H, Xue S, Yin J, Ji X. 2002. Paleomagnetic dates

of hominid remains from Yuanmou, China, and other Asian sites. J Hum Evol 43:27-41.
doi:10.1006/jhev.2002.0555
Sarel J, Zhang P, Weng Z. 2009. Recent discoveries in Baerya Cave (Bijie District,
Northern Province of Guizhou, China). Antiquity 83 (online).

Wang W, Potts R, Yuan B, Huang W, Cheng H, Edwards RL, Ditchfield P. 2007.

Sequence of mammalian fossils, including hominoid teeth, from the Bubing Basin caves,
South China. J Hum Evol 52: 370-379. doi:10.1016/j.jhevol.2006.10.003

Zhu RX, Potts R, Xie F, Hoffman KA, Deng CL, Shi CD, Pan YX, Wang HQ, Shi RP,
Wang YC, Shi GH, Wu NQ. (2004). New evidence of the earliest human presence at
high northern latitudes in Northeast Asia. Nature 431:559-562.