The Ostrich: From Whence this Derpy Horror?

Male and two female ostriches Struthio camelus, copyright Yathin S. Krishnappa.

Ostriches are widely known for two things: firstly, that they are the largest living bird by a quite respectable margin, and secondly, that they look ridiculous. Seriously, is there anyone out there who can look at the animals in the picture above and not think them ludicrous. Though I am, admittedly, invoking the luxury of distance: my uncle spent a year or two raising ostriches back during the brief boom of ostrich farming in New Zealand in the early 2000s, and I can say from experience that what looks humorous from afar is, close up, intimidating in a way no other animal is. They're just so tall*.

*Not to mention their well-earned reputation for gobbling down any item that attracts their attention. There are numerous stories out there demonstrating that wearing jewellery in an ostrich enclosure is a bad idea.

The modern ostrich is commonly regarded as a single species, Struthio camelus, found in savannah and semi-desert habits around Africa. There are some grounds for recognising the Somali ostrich S. molybdophanes of the Horn of Africa as a separate species—it is both genetically and morphologically divergent from other ostrich populations (for instance, its skin is blue rather than pink or red), and there is a small amount of overlap in range between Somali and typical ostriches—but this question remains open. Other subspecies are the North African ostrich S. c. camelus, the southern ostrich S. c. australis, and the Masai ostrich S. c. massaicus of Tanzania and Kenya. A fifth subspecies, the Arabian ostrich S. c. syriacus, became extinct around 1940, though it is worth noting that mitochondrial DNA extracted from specimens of Arabian ostriches in the British Museum did not separate them from the North African ostrich (Robinson & Matthee 1999). Ostriches can not really be confused with any other modern bird: not only is their remarkable size (matched by the remarkable size of their eggs), but they are the only birds to have reduced the number of toes to just two, with only the third and fourth toes of the standard bird foot remaining. This feature is generally presumed to be related to their cursorial lifestyle.

More evidence that ostriches are just daft. Copyright Georges Olioso.

Ostriches are members of the group of birds known as palaeognaths, that also includes such birds as the emu, kiwis, cassowaries, rheas and tinamous (the flightless members of the palaeognaths are commonly referred to as the ratites, but recent studies have cast doubt on whether flightlessness in the palaeognaths has a single origin). Phylogenetic relationships within the palaeognaths have been shuffled about considerably over the years (and even now are probably not really settled), but it is generally agreed that ostriches probably diverged from their nearest living relatives a long time ago (Burleigh et al. [2015], for instance, place them as the sister taxon to all other palaeognaths). Just how long ago we can't really say, the early fossil record of ostriches (and palaeognaths in general) being pretty dire. The heron-sized middle Eocene Palaeotis weigelti from Germany has been suggested to be a direct relative of ostriches but the evidence for this is equivocable (Mayr 2009). The earliest undoubted ostrich is the early Miocene Struthio coppensi from Namibia, and this is already similar enough to modern ostriches to be placed in the same genus.

Fossil ostriches are known from southeastern Europe to China, and survived across much of Asia until the Pleistocene. Several species have been named, but the usual vagaries of preservation make it debatable how many are distinct. Matters are complicated by several 'species', such as the Ukrainian Struthio chersonensis, that have been named based on fossil eggshells, raising questions as to whether such names can or should be applied to associated body fossils. Also unknown are the phylogenetic relationships between modern and fossil ostriches: whether the Eurasian ostriches represented a single or multiple dispersals out of Africa, or even whether ostriches may have originated in Eurasia*.

*It was suggested at one point that ostriches may have originally come from India, only dispersing to Africa after the subcontinent latched onto the rest of Eurasia. Support for this was based on the phylogenetic hypotheses that ostriches and the South American rheas formed an exclusive clade within the palaeognaths, and that the divergence of the flightless ratites was directly influenced by the division of the Gondwanan supercontinent (a South American-African connection being inconsistent with Africa being the first part of Gondwana to be separated). As support for both these arguments has declined, the need to somehow get ostriches out of Africa has evaporated.

The earliest known ostrich, the aforementioned Struthio coppensi, was a smaller and more slender bird than the modern ostrich, but some fossil species were larger. Perhaps the tallest ostrich species was S. oldowayi of the Tanzanian Pleistocene, which had a femur about a third as long again as the modern species. The femur of the Georgian Plio-Pleistocene S. dmanisiensis was not quite as long as that of S. oldowayi but it was considerably more robust, suggesting a proportionally solidly-built bird (Vekua 2013). Struthio brachydactylus (which sometimes moonlights as S. chersonensis) from the Miocene of Ukraine was also robustly built, albeit probably no taller (if not shorter) than a modern ostrich, but its main distinction lies in it taking the toe reduction seen in other ostriches even further. The fourth toe was reduced, with more weight placed on the third toe, making this species functionally almost single-toed (Boev & Spassov 2009).

REFERENCES

Boev, Z., & N. Spassov. 2009. First record of ostriches (Aves, Struthioniformes, Struthionidae) from the late Miocene of Bulgaria with taxonomic and zoogeographic discussion. Geodiversitas 31 (3): 493–507.

Burleigh, J. G., R. T. Kimball & E. L. Braun. 2015. Building the avian tree of life using a large-scale, sparse supermatrix. Molecular Phylogenetics and Evolution 84: 53–63.

Mayr, G. 2009. Paleogene Fossil Birds. Springer.

Robinson, T. J., & C. A. Matthee. 1999. Molecular genetic relationships of the extinct ostrich, Struthio camelus syriacus: consequences for ostrich introductions into Saudi Arabia. Animal Conservation 2: 165–171.

Vekua, A. 2013. Giant ostrich in Dmanisi fauna. Bulletin of the Georgian National Academy of Sciences 7 (2): 143–148.

Taxon of the Week: Leg or Breast?

This week's highlight taxon is one that is very familiar to me as a New Zealander, except it's not really. I've heard of these creatures since I was a little lad, and representations of them have been almost everywhere I've gone. I've never actually seen one. I don't know anyone who's every seen one. Probably no-one has seen one for hundreds of years, in fact. The Dinornithiformes are but a memory these days, long since converted into quarter-pack meals for Polynesian settlers. Ka ngaro i te ngaro a te moa - lost as the moa is lost.

The taxonomy of moa is complicated, but at present there are eleven species recognised as valid*. The order was unique to New Zealand - the "Australian" Dinornis queenslandiae De Vis, 1884, was based on a partial femur in the Queensland Museum, but this bone is now believed to have come from New Zealand and is assigned to Pachyornis elephantopus. In the past, Dinornithiformes has been divided into two families - the lightly built, more cursorial genus Dinornis in its own family and the other smaller and/or more heavily built genera in the Emeidae, but phylogenetic analysis has shown that Dinornis is nested within the Emeidae (Worthy & Holdaway, 2002).

*Eleven species were recognised in Worthy & Holdaway (2002), the most recent major review of Dinornithiformes. Bunce et al. (2003) reduced the number of species of Dinornis from three to two (see below), but Baker et al. (2005) increased the number of species of Megalapteryx from one to two.

The photo at the top of the page (from Wikipedia) shows the reconstructed Dinornis in the Auckland Museum. This specimen has been around for some time - it was built in 1913, though when the Natural History section of the museum was rebuilt it lost the tussock-land diorama it had previously inhabited (if I recall correctly) and moved into a glass case. The Auckland Museum moa stands about three metres tall, and modern interpretations would, unfortunately, label this a severely inaccurate reconstruction. It has been mounted in an unnaturally elevated stance, and should have been much more low-slung. A more realistic reconstruction was shown on a recent stamp issue, shown below (from New Zealand Birds):



Such a lowered reconstruction significantly lowers the height, but we're still looking at about two metres for the tallest Dinornis specimens. A moa of this size may have wighed over 150 kg (Worthy & Holdaway, 2002). Other species were smaller - the smallest was Megalapteryx didinus at probably about 40 kg. The super-heavy Pachyornis elephantopus was considerably shorter than large Dinornis, but may have weighed about the same amount. The image below comes from Nature, and shows three moa species against a 1.8 metre tall human. From left to right, the moas are a female Dinornis novaezealandiae, Megalapteryx didinus and Pachyornis elephantopus.



Despite their extinction prior to European settlement in New Zealand, a surprising amount of molecular data has been gleaned from ancient DNA studies of moa. Among other things, said molecular analyses have demonstrated that Dinornis displayed the highest degree of size dimorphism known from any bird (Bunce et al., 2003). Previously Dinornis had been divided into three species on the basis of size. Bunce et al. tested DNA from Dinornis remains for female-specific markers (birds differ from mammals in that it is the female that possesses different sex chromosomes [ZW], while the male has identical sex chromosomes [ZZ]). They found that all specimens that had been assigned to the smaller 'species' Dinornis struthoides were male, while all specimens of the larger 'species' D. novaezealandiae and D. giganteus were female. Molecular phylogenetic analysis also showed that these specimens were intermingled, with the major divide in the genus being not by size but by geography - the North Island and South Island populations (both containing representatives from all three 'species') were distinct, and were recognised as the separate (but morphologically indistinguishable) species Dinornis novaezealandiae (North Island) and D. robustus (South Island). While females showed a great deal of variation in size, the largest females would have been about 280% of the weight and 150% of the height of the largest males.

There is very little reliable information on the life habits of moa. By the time of European settlement, it had been long enough since the extinction of the moa that records of it in Maori oral tradition were seemingly few and far between, and those that were present had become significantly mythologised. Early researchers such as Owen and Haast interpreted moa as birds of open country, comparing them to modern ostriches and emus. However, the extensive New Zealand grasslands and fernlands these authors pointed to hadn't existed prior to human settlement, so moa were undoubtedly forest birds, foragers rather than grazers as also shown by preserved gizzard contents. Dinornis and Pachyornis seem to have had more fibrous diets with gizzards containing twigs and fibrous plants such as Phormium (the New Zealand flax) while Emeus and Euryapteryx with less robust bills had more selective diets of fruit and leaves. Interestingly, a well-preserved specimen of Euryapteryx geranoides showed a massive intrathoracic loop in the trachea, 1.2 metres long. In other birds such loops are associated with the ability to make loud, far-carrying calls, so moa (or at least Euryapteryx) would have been quite vocal birds in life.

REFERENCES

Baker, A. J., L. J. Huynen, O. Haddrath, C. D. Millar & D. M. Lambert. 2005. Reconstructing the tempo and mode of evolution in an extinct clade of birds with ancient DNA: The giant moas of New Zealand. Proceedings of the National Academy of Sciences of the USA 102(23): 8257-8262.

Bunce, M., T. H. Worthy, T. Ford, W. Hoppitt, E. Willerslev, A. Drummond & A. Cooper. 2003. Extreme reversed sexual size dimorphism in the extinct New Zealand moa Dinornis. Nature 425: 172-175.

Worthy, T. H., & R. N. Holdaway. 2001. The Lost World of the Moa: Prehistoric life of New Zealand. Indiana University Press: Bloomington (Indiana).