john hawks weblog

paleoanthropology, genetics and evolution

Miocene ape evolution

  • The diet of Gigantopithecus

    Wed, 2011-10-26 13:07 -- John Hawks
    Synopsis: 
    Gigantopithecus was once imagined as an exclusive bamboo feeder, but evidence suggests a broader diet focused on fruits

    Gigantopithecus has often been described as a bamboo eater, based on analogy with another kind of large herbivore in China, the giant panda. Giant pandas have several specialized feeding adaptations to support their bamboo diet. The most famous of these is the expansion of what we would call a wrist bone in most mammals, a sesamoid bone associated with the distal radius. In giant pandas, this bone projects from the arm in a way that makes it function similar to an additional digit for the hand, a solution described as "the panda's thumb". This "thumb" is used to grip the bamboo stems so that the teeth can work through the indigestible fiber and woody portions of the bamboo stems into the softer shoots. As a famous example in the history of evolutionary biology, the panda's thumb was celebrated by the evolutionary biologist Stephen Jay Gould as a unique evolutionary solution.

    Some information on the dietary proportions of giant pandas is available from BBC News. The following is quoted from that site:

    Ninety nine per cent of a panda's diet is made up of 30 species of bamboo. The remaining one per cent is made up of other plants and meat. Their digestion of bamboo is very inefficient; pandas only digest about 20 per cent of the dry matter of bamboo, whereas most herbivores assimilate about 80 per cent. This means that they must eat large amounts to obtain their energy requirements. They can eat between 12 and 38kg of bamboo shoots, leaves and stems per 24 hour period.

    Giant pandas can maintain this dietary solution only by sustaining a high feeding rate. The digestibility of bamboo varies markedly across the year (Wei et al. 1999), and in the winter when new growth is rare or absent, there are very few nutrients available. Pandas do not have any significant digestion of the structural elements of cell walls or other fibers. They therefore must extract the proteins and simple carbohydrates from bamboo and pass the bulk as quickly as practicable. To this end, they have wide and flat molars and premolars compared to other bears. These are not teeth with high crowns and shearing surfaces. This makes them different from primates, like gorillas and colobus monkeys, that eat a high proportion of leaves and other vegetation. It seems that pandas are not really in the business of cutting fibrous bamboo into a pulp; but instead they crush the bamboo to extract as much of the cell contents as possible.

    Gigantopithecus also had broad, flat molars and premolars. These teeth had relatively thick enamel. Enamel thickness is a tricky indicator of diet, because there are actually advantages to having enamel that wears through completely during life. If the goal is to maintain an effective shearing surface on the tooth for cutting fibrous plant material, then thin enamel exposes the softer dentin, which wears faster. The wear gradient between the two maintains a topography to the tooth surface that is a better shearing implement than a flat, thick-enameled tooth. So the thick molar enamel in Gigantopithecus would not be very useful for shearing bamboo leaves into an undifferentiated mush. But those teeth might have been used to crush bamboo to extract the cell contents while leaving the mass mostly intact.

    The evidence suggests that Gigantopithecus differed from giant pandas in having a more varied diet. One of the world's experts on Gigantopithecus is the paleoanthropologist Russ Ciochon. He has
    a very nice article about the species which appeared in Natural History magazine in 1991. This nice review features the history of Gigantopithecus discoveries, our current understanding of their anatomy, diet, and history, and Ciochon's own attempts to find fossil Gigantopithecus in Vietnam.

    Ciochon describes looking for phytoliths on the teeth as evidence of diet. When the fossil teeth of Gigantopithecus were examined with scanning electron microscopy, dozens of phytoliths were found:

    More than half of the phytoliths we observed were long and needlelike and could be attributed to the vegetative part of grasses, possibly bamboo. The rest were conical or hat shaped, attributable to the fruits and seeds of dicotyledons. Piperno tentatively identified them as fruits from a tree of the family Moraceae, quite possibly durian or jackfruit, both of which are common throughout tropical Southeast Asia. This proved that Gigantopithecus had a varied diet, although we still suspect that bamboo was its staple food.

    This work is described in
    Ciochon et al. (1990) in PNAS, which includes scanning electron micrographs of the phytoliths.

    Of course the relative quantities of phytoliths do not directly address dietary composition, since different plants have different phytolith abundances. Likewise, one might speculate that the phytoliths on fossil teeth represent foods eaten near the time of death -- a "last meal" effect. This might explain the apparent evidence for one kind of fruit in the Gigantopithecus data: the individual died at the time that fruit was in season. In any event, Ciochon and colleagues (1990) conclude it likely that Gigantopithecus had a very broad diet, that nonetheless included bamboo as a staple. In support of this, they cite an examination of tooth wear by Daegling and Grine (1989 in abstract; later published in 1994 in SAJS) that found Gigantopithecus microwear to be similar to chimpanzees. Chimpanzees themselves eat a majority of fruit, with smaller proportions of leaves, insects, and meat.

    References:

    Wei F, Feng Z, Wang Z, Zhou A and Hu J. 1999. Use of the nutrients in bamboo by the red panda (Ailurus fulgens). J Zool Lond 248:535-541.

    Ciochon RL, Piperno DR and Thompson RG. 1990. Opal phytoliths found on the teeth of the extinct ape Gigantopithecus blacki: implications for paleodietary studies. Proc Natl Acad Sci U S A 87:8120-8124.
    JSTOR

    Dean MC and Schrenk F. 2003. Enamel thickness and development in a third permanent molar of Gigantopithecus blacki. J Hum Evol 45:381-387.

    Daegling DJ and Grine FE. 1994. Bamboo feeding, dental microwear, and diet of the Pleistocene ape Gigantopithecus blacki. S Afr J Sci 90:527-532.

    Ungar P. 1998. Dental allometry, morphology and wear as evidence for diet in fossil primates. Evol Anthropol 6:205-217.

  • Meet Gigantopithecus

    Tue, 2011-10-25 00:09 -- John Hawks
    Synopsis: 
    Laboratory introduction to the species Gigantopithecus blacki, with discussion of its body size relative to gorillas and robust australopithecines.

    Gigantopithecus blacki was, as its name implies, a gigantic ape from the Pleistocene of China. Its remains consist only of teeth and jaws, but these are of a tremendous size, with the largest specimens nearly twice the dimensions of male gorilla teeth and jaws. A similar, slightly smaller jaw is known from the Miocene of northern India, and has been called Gigantopithecus bilaspurensis [1].

    Here you see casts of some of the teeth of Gigantopithecus blacki. Assuming that Gigantopithecus had the same proportion of tooth size and body mass as living apes, these Chinese remains would suggest a body mass of over 400 kg for the largest individuals. But should we assume a model of body size like that of today's large great apes, such as the orangutan and gorilla? Or should we assume a model in which Gigantopithecus had enlarged jaws and teeth relative to its mass, as is the case in the extinct robust australopithecines?

    Examine the Gigantopithecus teeth in comparison to modern gorilla teeth and jaws, and the teeth and jaws of Australopithecus boisei and Australopithecus robustus. How do the femora of A. robustus compare to the gorilla femur? How do the molars of these species compare? Which do you think is the better model for Gigantopithecus, and what would you predict as the body mass of this extinct species?


    References

    1. Simons EL, Ettel PC. Gigantopithecus. Scientific American. 1970;222:77–85.
  • The homoplastic apes

    Sat, 2011-02-19 16:08 -- John Hawks

    Bernard Wood and Terry Harrison have published a review paper in Nature[1], arguing that the extent of anatomical convergence among Miocene apes makes it difficult to reconstruct their relationships. The keyword of their essay is "homoplasy", a word for the situation when characters evolve convergently, in parallel, or in reverse. When parallelism or convergence have been common enough, we will find it difficult to use morphological characters to test hypotheses about the phylogeny of species. The butt of their essay is Ardipithecus, with extension to Sahelanthropus and Orrorin:

    We emphasize that we are not claiming that the presence of homoplasy in and around the hominin clade, and the other methodological and analytical limitations of phylogenetic analyses noted above, doom all efforts to recover evolutionary relationships to failure. Nor are we claim- ing that Ar. ramidus, S. tchadensis and O. tugenensis are definitely not hominins. We do, however, advocate that those palaeoanthropologists whose considerable and much valued efforts in the field are rewarded with fossils as significant as those from Aramis, Toros Menalla, Lukeino and Malapa acknowledge the potential shortcomings of their data when it comes to generating hypotheses about relationships.

    The main points won't be news to many readers. One long-time correspondent called the essay "idea homoplasy", focusing as it does on the same issues that I covered here in 2009, and the evidence for craniodental parallelism among Miocene apes that we reviewed in our 2006 paper on Sahelanthropus [2]. That's probably too kind respecting my role in this matter, but I suppose it is one of the chief drawbacks of blogging that I pass by many of these opportunities to do perspective and review articles for top-tier journals. But still, why wait two years to make a point that can really be made in an afternoon, and reach many, many more readers? I mean, Nature wants $32 for this review. Seriously.

    I find it nonetheless interesting to see Nature take up the subject, however belatedly, and Wood and Harrison ably cover some of the problems of convergence in Miocene apes. Harrison is an expert on Oreopithecus, and the paper includes four paragraphs describing its relevance to the topic of homoplasy in fossil apes. To me, this is a key comparison that deserves a longer treatment. You can find a bit more information in my 2009 Ardipithecus coverage ("The Ardipithecus pelvis"), but I'm not really the person to do a thorough job of it. I would hope that someone will return to the issue at greater length, but I think that would require access to the Ardipithecus pelvis reconstruction, which is not available for independent inspection.

    In light of the Pliocene goat-man lunacy the other day, Daniel Lende ties the satire directly to this little Ardipithecus dustup. He points to the dueling quotes in this Science News article by Bruce Bower:

    “Researchers have to stop publishing papers that say, essentially, ‘This fossil is an early hominid, so suck it up and accept it,’” [Bernard] Wood says. “Nature and Science could change this practice overnight if they wanted to.”

    ...

    “With no new data, no new ideas, no new methods, no new hypothesis, no new experiments, no new fossils, not even a new classification, this paper will leave everybody wondering what’s happened to the peer review process at Nature,” [Tim] White says.

    And then there's the write-up by Katherine Harmon, who pulls quotes from Nature's podcast on the paper:

    Tim White, of the University of California, Berkeley, and one of the lead authors on the 2009 Ardi papers, called the new article a "six page illustrated op-ed piece" in the Nature podcast. He maintains that "whole functional complexes"—not just individual characteristics—that were described in his team's papers link Ardi to humans "to the exclusion of the great apes."

    Oh, goodness. I'm not entirely sure what is to be done about these folks. The thing about the 17-year inquiry into the "one large goat" theory, is that I bet they made the CT reconstructions and dental measurements of the goat-men available for inspection.


    References

  • Miocene ape review, in brief

    Fri, 2010-01-29 19:58 -- John Hawks

    Science has a short essay by Terry Harrison this week about Miocene ape evolution: "Apes among the tangled branches of human origins."

    This is the sort of article that shows just how frustrating the Miocene apes can be. It's short, probably not even 1500 words, and it has more than 20 genus names stacked into it -- and he doesn't even list Proconsul and its Early Miocene ilk.

    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.

    References:

    Harrison T. 2010. Apes among the tangled branches of human origins. Science 327:532-534. doi:10.1126/science.1184703

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