Tetrapod Zoology

In the previous post we looked at the small, island dwelling crocodilians of the south-west Pacific. I personally find it exciting that such animals were (in the case of at least some of the species) alive until just a few thousand years ago, that they were encountered by people, and that their remains have eluded detection until recent decades. The odds are high that further species await discovery. Here’s another article that originally appeared at Tet Zoo ver 1.

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Before we get to the new stuff, here’s something relevant to the mekosuchine article: a never-before-seen skull reconstruction of Mekosuchus whitehunterensis. This image was kindly provided by John Scanlon: John has a lot of new mekosuchine material and is due to publish on it at some stage. Anyway, moving on…

All of the island dwelling crocodilians I discussed in the previous post were members of the predominantly Australasian mekosuchine radiation. But there is one recently extinct crocodilian of the south-west Pacific that I didn’t mention, and which isn’t a mekosuchine. First reported by Charles De Vis in 1905, it’s a long-snouted form known from Pleistocene remains discovered at Busai on Murua, one of the Solomon Islands. Because of its long, slender jaws, De Vis regarded this animal as a gharial and named it Gavialis papuensis. It then languished in obscurity until 1982 when Ralph Molnar published a redescription.

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Molnar (1982) concluded that the Murua crocodilian almost certainly didn’t belong in the genus Gavialis, and that it was more likely closely related to Charactosuchus, Euthecodon or Ikanogavialis, with a relationship with the last named taxon being deemed most likely. That’s good news, because Ikanogavialis (best known for I. gameroi from Upper Miocene* Venezuela [a jaw segment from this taxon is figured at left]) is – while not the same thing as Gavialis – still undoubtedly a member of the gharial family, Gavialidae. South American Miocene Charactosuchus, while gharial-like, has been regarded as a highly unusual crocodylid of uncertain affinities (Langston 1965, Langston & Gasparini 1997), while Euthecodon – a uniquely African taxon, some species of which approached 10 m in length – is also a crocodylid, and perhaps a close relative of the living dwarf crocodiles (and we’ll discuss those more in a moment). Most recently, Rauhe et al. (1999) listed the Murua gharial as belonging to Ikanogavialis, and if this has been accepted then presumably we should refer to it as Ikanogavialis papuensis. The presence of this genus in both Venezuela and the south Pacific might seem odd given that living gharials are freshwater animals, but the fossil record shows that gharials formerly occurred widely in marine environments around the world.

* And, according to Langston & Gasparini (1997), not from the Pliocene as usually stated.

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Like the south Pacific mekosuchines, the Murua gharial was again fairly small, at 2-3 m long. Its fossils were associated with those of sea turtles and sirenians, so it was almost certainly marine. To the list of small crocodilians that inhabited the south-west Pacific, we can add gharials then. Whether the Murua gharial became extinct before humans colonised the region, or whether its extinction was caused by people, we again don’t know. Indeed the only known specimen’s exact geological age is unknown. Given this, and the many anthropogenic extinctions that occurred in the region, I can’t help but speculate that the Murua gharial survived into the Holocene, and that humans killed it off, but there’s no direct evidence for this. Regardless, it’s surprising that small, marine gharials survived so relatively late [the adjacent photo figures the living gharial species Gavialis gangeticus].

Aldabrachampsus dilophus

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Besides mekosuchines and gharials, we also know of a third crocodilian group that included island dwelling forms, and again the species concerned became extinct geologically recently. To look at the members of this group we need to move over to the Indian Ocean. And it’s here that we find the most recently named of all the crocodilians we’ve looked at: Aldabrachampsus dilophus, from Aldabra (Brochu 2006). Though recently named, Aldabrachampsus was actually first described in 1976, though at this time it was misidentified as representing a dwarf population of Crocodylus niloticus (Arnold 1976). In being of Pleistocene age, Aldabrachampsus is like both Volia from Fiji (see previous post) and the Murua gharial in being too old to have its extinction indisputably linked to the arrival of people. Arnold (1976) discussed environmental changes that occurred on Aldabra that may have caused the extinction of endemic reptiles, among them geckos, iguanas and skinks, with the most notable of them being the breaching of the atoll rim and subsequent habitat degradation that occurred about 4000 years ago [Osteolaemus tetraspis shown in adjacent image. Read on].

Various skull features make Aldabrachampsus unusual, including the shape of its premaxillae, and the orientation of its tooth row and external nostrils. However, its most obvious feature would almost certainly have been the convex crests that grew from the dorsolateral edges of the squamosal bones at the back of its skull. Some living crocodiles have crest-like projections in this region, but none have the prominent, elongate structures present in Aldabrachampsus. These crests explain the specific name, ‘dilophus’ meaning ‘with two crests’.

Given that Aldabrachampsus was comparable in size to the smallest living crocodilians – that is, between 2 and 2.5 m long – it’s tempting to assume that, like so many island dwelling tetrapods, it was a dwarf. This would actually be odd for a crocodilian, given that other island dwelling forms are not dwarfed relative to their mainland relatives (as we saw in the previous post, island-dwelling mekosuchines were not dwarfs, as their mainland relatives were equally small). Indeed, stratigraphic evidence seems to contradict the possibility that Aldabrachampsus evolved its small size as a result of insular isolation: its remains come from sediments that were deposited shortly after a period of Aldabran submergence, so it’s unlikely to have evolved on the island. It presumably swam in from elsewhere.

The former Osteolaemine Empire

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What sort of crocodilian was Aldabrachampsus? It was a crocodylid, but there’s no indication that it was anything to do with the mekosuchines: instead, there are reasons for thinking that it was an osteolaemine. That is, a member of the same crocodylid clade as the west African dwarf crocodiles (Osteolaemus*) – see picture above – and the extinct Madagascan species Voay robustus (Brochu 1997, 2006, 2007). Some phylogenetic studies find that the osteolaemines also include Euthecodon, the bizarre gharial-like African taxon we met above, as well as Rimasuchus (Brochu 1997, 2007), a broad-snouted east African taxon that grew to 7 m or more in length [Rimasuchus skull shown here, © Koobi Fora Research Project and borrowed from here]. The African slender-snouted crocodile Crocodylus cataphractus might also be an osteolaemine, a view that would be in agreement with data suggesting that it needs removing from Crocodylus (the old generic name Mecistops Gray, 1844 is available: see McAliley et al. 2006), and additional fossil African crocodylids also seem to belong to this group. If this is all valid, then little Osteolaemus is a sorry remnant of a once diverse group that included several enormous species. Anyway, within this group, an affinity between Aldabrachampsus and Voay is particularly plausible given that both taxa share a vaulted palate and large squamosal crests.

* Though conventionally thought to include just a single living species (O. tetraspis), new data has caused some workers to regard a second species as valid (McAliley et al. 2006). This is O. osborni, a taxon from the Congo (first described in 1919 and given its own genus, Osteoblepharon) until recently regarded as a subspecies of O. tetraspis.

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Unlike the crests of Aldabrachampsus, those of Voay were large horn-like growths (see photo above, my hands for scale), and unlike both Aldabrachampsus and Osteolaemus, this species was large and comparable in size to a Nile crocodile Crocodylus niloticus. Originally described in 1872, Voay has been mostly considered synonymous with C. niloticus, but ‘this synonymy results from an inadequate initial description and from subsequent misidentifications of living C. niloticus from Madagascar as C. robustus‘ (Brochu & Storrs 1995). Brochu’s recent redescription and analysis confirms that the ‘true’ ‘Crocodylusrobustus – now Voay robustus (‘voay’ is the Malagasy word for crocodile) – is very different from the members the genus Crocodylus, and closest to Osteolaemus as discussed here (Brochu 2007).

Voay wasn’t as big as Euthecodon or Rimasuchus, reaching 4-5 m in length (Burness et al. (2001) estimated its weight as 170 kg). This size would have made it the largest predator on Madagascar, and given that prehistoric Madagascar was also home to giant eagles and fossas, the lemurs, elephant birds and other animals of the island would certainly have lived in fear of formidable predators.

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Again, what fascinates me most about Voay is how recently it was alive. So far as I can tell from the literature, an exact date for its extinction is unknown, and I’d be interested to know if it disappeared as part of the anthropogenic wave of extinctions that occurred on the island. Brochu (2007, p. 857) suggested this too, and also noted that C. niloticus (which does not have a confirmed prehistoric occurrence on Madagascar) might only have invaded the island after Voay became extinct. This is consistent with the fact that Crocodylus crocodiles seem to be highly opportunistic, and good at colonising new areas. When mekosuchines, osteolaemines and members of other groups go extinct, it seems that Crocodylus species quickly move in. But – do Crocodylus crocodiles really only move in afterwards, or are they linked in some way with the extinction of these ‘old endemics’? That’s a good question, and more research is needed [adjacent Nile crocs from wikipedia. Osteolaemus tetraspis below].

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And this is not the end of the story: what about the island-dwelling crocs of the Caribbean? Of the Mascarenes? Oh well, I can’t cover everything…

For a previous articles on extinct crocodilians see…

Refs – -

Arnold, E. N. 1976. Fossil reptiles from Aldabra Atoll, Indian Ocean. Bulletin of the British Museum (Natural History). Zoology 29, 85-116.

Brochu, C. A. 1997. Morphology, fossils, divergence timing, and the phylogenetic relationships of Gavialis. Systematic Biology 46, 479-522.

- . 2006. A new miniature horned crocodile from the Quaternay of Aldabra Atoll, western Indian Ocean. Copeia 2006, 149-158.

- . 2007. Morphology, relationships, and biogeographical significance of an extinct horned crocodile (Crocodylia, Crocodylidae) from the Quaternary of Madagascar. Zoological Journal of the Linnean Society 150, 835-863.

- . & Storrs, G. W. 1995. The giant dwarf crocodile: a reappraisal of ‘Crocodylus’ robustus from the Quaternary of Madagascar. In Patterson, Goodman & Sedlock (eds) Environmental Change in Madagascar, p. 70.

Burness, G. P., Diamond, J. & Flannery, T. 2001. Dinosaurs, dragons, and dwarfs: the evolution of maximal body size. Proceedings of the National Academy of Sciences 98, 14518-14523.

Langston, W. 1965. Fossil crocodilians from Colombia and the Cenozoic history of the Crocodilia in South America. University of California Publications in Geological Sciences 52, 1-169.

- . & Gasparini, Z. 1997. Crocodilians, Gryposuchus, and the South American gavials. In Kay, R. F., Madden, R. H., Cifelli, R. L. & Flynn, J. J. (eds) Vertebrate Paleontology in the Neotropics: The Miocene fauna of La Venta, Colombia. Smithsonian Institution Press (Washington, D.C.), pp. 113-154.

McAliley, L. R., Willis, R. E., Ray, D. A., White, P. S., Brochu, C. A. & Densmore, L. D. 2006. Are crocodiles really monophyletic? – Evidence for subdivisions from sequence and morphological data. Molecular Phylogenetics and Evolution 39, 16-32.

Molnar, R. E. 1982. A longirostrine crocodilian from Murua (Woodlark), Solomon Sea. Memoirs of the Queensland Museum 20, 675-685.

Rauhe, M., Frey, E., Pemberton, D. S. & Rossman, T. 1999. Fossil crocodilians from the Late Miocene Baynunah Formation of the Emirate of Abu Dhabi, United Arab Emirates: osteology and palaeoecology. In Whybrow, P. J. & Hill, A. (eds) Fossil Vertebrates of Arabia. Yale University Press (New Haven), pp. 163-185.

Comments

  1. #1 Jerzy
    May 14, 2009

    Thanks for the interesting post!

    It is a bit of miracle, that tiny atoll could support crocodiles for long enough to evolve into a separate species.

    I guess that giant tortoises and flightless rails mean that Aldabra was never permanently occupied by man before European discovery.

  2. #2 Dartian
    May 14, 2009

    what fascinates me most about Voay is how recently it was alive. So far as I can tell from the literature, an exact date for its extinction is unknown, and I’d be interested to know if it disappeared as part of the anthropogenic wave of extinctions that occurred on the island. Brochu (2007, p. 857) suggested this too

    Bickelmann & Klein (2009) disagree with Brochu and are of the opinion that climate change, rather than human persecution, wiped out V. robustus. The case they make isn’t particularly strong, though.

    Incidentally, regarding the length of Voay robustus, both Brochu (2007:855) and Bickelmann & Klein (2009:13) suggest that this species was rather less than 4 m in length.

    Oh well, I can’t cover everything…

    In response to this I can only paraphrase a certain current world leader: Yes you can!

    Reference:

    Bickelmann, C. & Klein, N. 2009. The late Pleistocene horned crocodile Voay robustus (Grandidier & Vaillant, 1872) from Madagascar in the Museum für Naturkunde Berlin. Fossil Record 12, 13-21.

  3. #3 Darren Naish
    May 14, 2009

    Thanks for that: I wasn’t aware of the Bickelmann & Klein paper (if anyone has a pdf, please pass it on).

  4. #4 Metalraptor
    May 14, 2009

    Is it just me, or does that skull of Mekosuchus at the top look suspiciously like a mammal?

  5. #5 Jorge Velez-Juarbe
    May 14, 2009

    what about the island-dwelling crocs of the Caribbean?

    Maybe I should sit down and do something about it…

    How much of the Murua crocodylian was found?

    There’s also something referred tentatively to Ikanogavialis from the Late Miocene Baynunah Formation, identified from part of a dentary (Rauhe et al., 1999).

    Rauhe, M., E. Frey, D. S. Pemberton & T. Rossmann. 1999. Fossil Crocodilians from the Late Miocene Baynunah Formation of the Emirate of Abu Dhabi, United Arab Emirates: Osteology and Paleoecology; pp.163-185 in P. J. Whybrow & A. Hill (eds.), Fossil Vertebrates of Arabia. Yale University Press.

  6. #6 Darren Naish
    May 14, 2009

    Yeah, Rauhe et al. (1999) is cited in the article above (indeed, these are the authors that identified the Murua gharial as Ikanogavialis), but I forgot to add it to the references. Will correct this now, thanks.

  7. #7 David Marjanović
    May 14, 2009

    I guess that giant tortoises and flightless rails mean that Aldabra was never permanently occupied by man before European discovery.

    Of course. Has never been doubted.

    Is it just me, or does that skull of Mekosuchus at the top look suspiciously like a mammal?

    Proportion-wise it’s interesting in that respect, yes. Reminds me strongly of the notosuchians.

  8. #8 Zach Miller
    May 14, 2009

    Agreed. When I first saw the picture, I thought Darren was going to be talking about notosuchians!

    Potentially stupid question: What are those prong-like bones sticking down off the bottom of the skull? I know lots of crocs (if not all) had them, but I’ve never known what they’re called, or their function.

  9. #9 Jorge Velez-Juarbe
    May 14, 2009

    What are those prong-like bones sticking down off the bottom of the skull?

    Zach: Those are the Pterygoid + Ectopterygoid: I’m not knowledgeable about crocodylian musculature, but these might provide attachment area for some of the muscles of mastication.

    BTW: Darren is that skull of Voay robustus you’re holding AMNH 3101?

  10. #10 Jura
    May 15, 2009

    The pterygoids provide attachment for the muscles of the same name. They don’t play a role in mastication (as no crocodile masticates), but instead provide power for the incredibly fast “snapping” ability of crocodylian jaws.

  11. #11 John Scanlon FCD
    May 15, 2009

    Suspiciously like a mammal, or notosuchian, or, for example, something like Cynognathus (recently shown here by Laelaps). The steep taper of the mandible to a very shallow chin allows the tip of the jaws to reach ground level, and seems one of the strongest bits of evidence for terrestriality (suggests feeding on small items on a surface). It’s a specialization of Mekosuchus, not present in things that appear to be closely related like Quinkana, Trilophosuchus and Baru.

  12. #12 Jerzy
    May 15, 2009

    I wonder if V. robustus could be extant and overlooked among C. niloticus. How many people looked closely at Madagascar crocodiles?

  13. #13 johannes
    May 15, 2009

    > The African slender-snouted crocodile Crocodylus
    > cataphractus

    Called Panzerkrokodil in German, which is rather confusing, because Bakker uses the pseudo-germanicism Panzercroc for the paleogene ziphodont croc *Pristichampsus*.

  14. #14 David Marjanović
    May 15, 2009

    How many people looked closely at Madagascar crocodiles?

    Those that have are probably all dead ;-)

  15. #15 Sordes
    May 15, 2009

    I once asked Chris Brochu about the size of Voay robustus, and he wrote me, that there were no specimens bigger than 3m. This is especially interesting, because there is no indication that there was ever a giant form of crocodile at Madagascar, because sometimes you find the notion that remains of an extinct form of the nile croc up to 10 m were found there. It is most probably nothing than an error which found its way into literature. Modern nile crocs don´t reach 10m, and those from Madagascar, which are genetically identical to those from mainland Africa, don´t grow that big too. There were no extinct forms of nile crocodiles at Madagascar, and the only native and now extinct crocodile was very small.

  16. #16 LeeB
    May 15, 2009

    Jerzy, you wonder if V. robustus could be extant and just overlooked.
    Well at least some crocodiles in Madagascar live underground in rivers that flow through caves.
    So who knows.

    And also of interest, the first person to write about Socotra 2000 years ago mentioned crocodiles, giant lizards and various large tortoises.

    Perhaps when palaeontologists get around to looking there they might make some interesting discoveries.
    Especially as there are apparently caves that animals could have fallen into and got preserved in.

    LeeB.

  17. #17 Alan Kellogg
    May 16, 2009

    Speaking of looking for extinct or exant crocodilians in places they just might be, but hadn’t been looked for before; has anybody taken a gander at the Mesopotamian Valley for croc sign?

  18. #18 dinosauricon
    May 17, 2009

    Hey guys can I just check something what is the size of Quinkana because I have seen sources which claim that specimens grew up to 5m but other sources say that it only grew up to 3m. Thanks in advance.

  19. #19 David Marjanović
    May 17, 2009

    sometimes you find the notion that remains of an extinct form of the nile croc up to 10 m were found there.

    Perhaps someone misread “10 feet” (3.05 m), and then one copied from another…

  20. #20 John Scanlon FCD
    May 17, 2009

    Quinkana babarra > Q. fortirostrum > others, size seems to have peaked in the Pliocene. I haven’t got all the papers on the genus, but Steve Salisbury was talking about a 7 m Quinkana in the mid-90′s which I assume was an estimate for babarra.

  21. #21 Jorge Velez-Juarbe
    May 19, 2009
    what about the island-dwelling crocs of the Caribbean?

    Maybe I should sit down and do something about it…


    Here it is.

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