31 August 2011

Characters that Support the Great Ape Clades

Anyone familiar with the current state of great ape phylogeny knows that the following structure is well-supported:
  • Great apes are a clade.
    • Orangutans are a subclade of great apes.
    • African great apes are a subclade of great apes.
      • Gorillas are a subclade of African great apes.
      • Mangani are a subclade of African great apes.
        • Humans are a subclade of mangani.
        • Chimpanzees are a subclade of mangani.
And most such people probably know that the primary evidence for this structure is molecular. But there has to be morphological data to back this up, right?

Great Apes
I've been trying to hunt down such morphological data, but it's been a bit hard. There really aren't that many morphology-based cladistic studies of primates, and the few that exist either exclude humans or focus on stem-humans more than living great apes.

An example of a study that looks at a wide array of fossil and living primates, but fails to include humans:
  • Rossie & Seiffert (2006). Continental paleobiogeography as phylogenetic evidence. Pages 469–522 in Lehman & Fleagle (eds.) Primate Biogeography: Progress and Prospects. Springer, New York. 546pp. isbn:0387298711
An example of a study that includes some living great apes, but focuses on stem-humans:
  • Strait & al. (1997). A reappraisal of early hominid phylogeny. Journal of Human Evolution 32(1):17–82. pmid:9034954
I've compiled some shared character lists from these:



CHARACTERS SHARED BY CHIMPANZEES AND HUMANS, BUT NOT GORILLAS (Strait & al. 1997):
  • Inferior orbital margin never rounded laterally
  • Sagittal crest not present in males*
  • Postorbital constriction moderate or slight
  • Narrow supraglenoid gutter width
  • External auditory meatus positioned medially (not laterally)
  • Consistently small mandibular cross-section area at M1
  • Narrow mandibular extramolar sulcus
  • Very small premolar crown area
  • Very small molar crown area
* Fossils show this to be a convergence.

CHARACTERS SHARED BY CHIMPANZEES AND GORILLAS, BUT NOT ORANGUTANS OR GIBBONS (Rossie & Seiffert 2006):
  • Upper premolar cusp heteromorphy reduced
  • P3 MD length narrow
  • M1-M2 waisting present
  • M1-M2 crown tapering distal
  • Talon size large relative to trigon
  • Radial head outline smaller flat area
  • Dorsal ridge on radius present
  • Os centrale fused*
  • Metacarpal dorsal ridges present
  • Hamate hamulus distally projecting
  • Posterior calcaneoastragular joint faces dorsally
  • Supraorbital region strong inflated tori
  • Frontal sinus present*
* I've verified that these are present in humans, too. I'm sure most or all of the others are as well, but I haven't had time to check.

CHARACTERS SHARED BY CHIMPANZEES, GORILLAS, AND ORANGUTANS, BUT NOT GIBBONS (Rossie & Seiffert 2006):
  • I2 cingulum present
  • P4 crown shape sub-ovoid
  • P4 MD length narrow
  • p3 mesiolingual beak present
  • p3 sectoriality poor
  • p4 length very broad
  • m1-m3 distal fovea wide
  • Transverse process inclination dorsally and caudally inclined
  • Transverse processes on pedicle
  • Four lumbar vertebrae
  • Deltoid insertion on distal half
  • Radial styloid process reduced
  • Trapezium-MC1 joint sellar
  • MC1 prox articulation lateral extension
  • Hamate distolateral edge interrupted by ligament pit
  • Tibial condylar facets medial larger
  • Femoral condyles medial larger
  • Talar head width greater than height
  • Anterior calcaneal length less than one third
  • Heel tubercle present
  • Infraorbital surface of maxilla slopes anteroinferiorly
  • Snout length protruding
  • Palate shape parallel to tooth rows
  • Orbit inferior margin flush with cheek
  • Orbit position high
  • Nasal shape narrow superiorly
  • Horizontal palatine process narrow

What I'd like to do eventually is synthesize these into easily-readable lists of characters supporting the major great ape clades (great apes, African great apes, mangani). The lists are in pretty sloppy form right now, though. Things left to do:
  1. Double-check against the character matrices.
  2. Find the human state for the relevant characters in Rossie & Seiffert 2006.
  3. Weed out the symplesiomorphies and homoplasies.
  4. Translate these into less technical language, insofar as possible.

5 comments:

  1. Did you take a look at the Paper from Strait & Grine which they published in 2004 in the Journal of human evolution ("Inferring hominoid and early hominid phylogeny: the role of fossil species")?
    The charcter matrix they used is essentially the same as in the study from 1997, but in this study they also included Gibbon and Orang-Utan specimens. I haven't really read this study though so I can't tell you, if it includes any valuable Information for your work.

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  2. I have that one, but I forgot to take a look. Thanks for reminding me about it.

    Of course, that one only looks at craniodental characters, and Rossie 7 Seiffert only look at skeletal characters. It'd be nice to find something that looks at soft tissue as well.

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  3. Very interesting post. It's great to see support for these groups "we all know" exist.

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  4. Mike: "It'd be nice to find something that looks at soft tissue as well."

    You want soft tissue studies, eh? Then you might want to check out:

    Gibbs, S., Collard, M. & Wood, B. 2002. Soft-tissue anatomy of the extant hominoids: a review and phylogenetic analysis. Journal of Anatomy 200, 3-49.

    and

    Diogo, R. & Wood, B. 2011. Soft-tissue anatomy of the primates: phylogenetic analyses based on the muscles of the head, neck, pectoral region and upper limb, with notes on the evolution of these muscles. Journal of Anatomy 219, 273–359.

    (Yes; that last paper - which also has data for non-hominoid primates - really is 87 pages long.)

    These (and their reference lists) ought to get you started at least...

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  5. Those refs look very promising. Thanks, Dartian!

    ReplyDelete