Enantiornithine nesting colony, reconstruction by Julio Lacerda.

Like modern birds, and like their close relatives among the theropod dinosaurs, the birds of the Mesozoic Era laid eggs and, we reasonably infer, made nests. But what else do we know about reproductive behaviour in Mesozoic birds? Essentially, we know very little, and by “very little” I actually mean “just about nothing”. A new paper just published in Naturwissenschaften by Gareth Dyke, Mátyás Vremir, Gary Kaiser and myself describes an assemblage of bird eggshell and bones that is amazing in its scale, and fascinating in terms of its behavioural implications (Dyke et al. 2012). It shows, for the first time, that some Mesozoic bird species – only distantly related to modern birds – formed enormous breeding colonies.

Enantiornithine embryo, still in its egg, described by Zhou & Zhang (2004). The large wing feathers on this late-stage embryo indicate precocial (or superprecocial) habits for this species.

A few embryos and eggs belonging to the Mesozoic bird group known as Enantiornithines are already known (Elzanowski 1981, Mikhailov 1996, Grellet-Tinner & Norell 2002, Schweitzer et al. 2002, Zhou & Zhang 2004).

The most impressive of these fossils has to be the embryonic, partially feathered Lower Cretaceous Chinese enantiornithine [shown here], discovered within a near-complete egg, and described by Zhou & Zhang (2004). We can infer from the proportions and bone anatomy of this embryo and others that these bird species were precocial or even superprecocial: that is, the hatchlings were able to walk, fly and generally look after themselves within a day, or even within a few hours, of hatching.

But this doesn’t tell us anything about parental behaviour, since parents can still interact substantially with chicks in species where the chicks are precocial (though they don’t in all cases). And most of the other things you might ask about reproductive behaviour in Mesozoic birds remain unknown and often based on inference. Were egg clutches large or small? Where were nests located? What were the nests like? Did adults brood their eggs, or bury them? Did Mesozoic birds nest as lone individuals, in groups, or in colonies?

Given that Mesozoic birds were diverse in terms of body size and shape, and in ecology, they would presumably have been diverse in reproductive biology, just as modern birds are. Indeed, given that the Mesozoic members of Avialae (the bird lineage of Maniraptora) include early forms extremely similar to the little deinonychosaurs that were their close relatives, flightless walkers and runners, flightless, toothed seabirds, finch-, thrush-, crow- and vulture-sized terrestrial species, and early members of the ratite, duck, gamebird and neoavian lineages, it’s plausible that Mesozoic birds possessed a spectrum of reproductive behaviour, extending from ‘dinosaur-like’ at one end to ‘modern bird-like’ at the other.

Romania: fossil treasure trove

Our new research concerns a discovery made in the latest Cretaceous (Maastrichtian) Sebeş* Formation of Transylvania in western Romania. Already the Sebeş Formation (long erroneously considered Oligocene or Miocene in age!) is well known as a source of diverse Late Cretaceous fossils, including pleurodiran turtles, azhdarchid pterosaurs, ornithopods and the amazing dromaeosaurid theropod Balaur bondoc (Csiki et al. 2010, Vremir 2010).

* Pronounced something like “seb-esh”.

The Oarda de Jos sites (Sebeş River, Transylvania) as it looked when I visited it in 2011.

The sediments here were deposited by a large, meandering river system, surrounded by floodplains and forests with large trees. This was a continental environment*, with sedimentological evidence showing that long dry seasons were punctuated by short wet ones when extreme flooding sometimes occurred.

* This doesn’t mean “on a continent’, since the region was actually a large island at the time. Rather, it means that the environment was well inland, and without any estuarine or marine influence.

The entire Od accumulation as collected, prior to preparation. Scale bar = 50 mm. From Dyke et al. (2012).

It’s the detailed mapping and exploration of the Sebeş Formation by Mátyás Vremir of the Transylvanian Museum Society that has resulted in so many amazing Sebeş Formation discoveries. And it’s one of these that forms the focus of our new paper. In sediments exposed on the banks of the Sebeş River, Mátyás discovered a large, lens-shaped chunk of calcareous mudstone packed full of thousands of broken eggshell fragments. Unfortunately (but – I hope – understandably), the concretion couldn’t be extracted as a single mass; rather, it broke into lots of separate chunks. When complete it was about 80 cm long, 50 cm wide and 20 cm thick. Its breakage isn’t a bad thing, since this allows the interior of the mass (rather than merely its edges) to be examined in detail. While virtually all of the eggshell within the accumulation is preserved as small, broken fragments, seven near-complete eggshells are included as well, as are more than 60 small bones. We term this mass of sediment, eggshell and bones the ‘Od accumulation’ after the specific outcrop where it was discovered (the Oarda de Jos site).

A whole lot of eggshell

At top: two of the seven complete eggs preserved in the Od accumulation; at bottom, one chunk of the accumulation, showing how densely packed the eggshell fragments are. Scale bars = 10 mm. From Dyke et al. (2012).

The volume of eggshell in the Od accumulation is extraordinary. About 70-80% of the accumulation as a whole is formed by eggshell, meaning that this is an eggshell-supported rock, reminiscent of the Patagonian ‘egg beds’ that incorporate vast quantities of sauropod eggshell. Averaged out, most shell fragments in the Od accumulation are about 36 mm long. They’re so closely packed that, in one 26-cm-sq section of the accumulation, more than 150 eggshell fragments are present. The several complete eggs preserved in the accumulation are 40 mm long and 25 mm wide; based on these measurements, each 100 cubic cm of the accumulation contains eggshell equivalent to about 46 whole eggs (Dyke et al. 2012). As we say in the paper, “The quantity of eggshell fragments in the Od accumulation is astonishing and beyond normal palaeontological experience”.

Adult enantiornithine humerus from the Od accumulation, from Vremir (2010). As preserved, the specimen is c. 50 mm long.

The eggshell looks avialan, and the isolated bones jumbled up within the accumulation are definitely avialan. These bones appear to belong to both adult and juvenile birds: they reveal a number of characters that allow them to be identified as those of enantiornithines, in particular a taxon close to (or congeneric with) the Late Cretaceus Argentinean form Enantiornis (Dyke et al. 2012). SEM analysis of the eggshell fragments reveals two equally thick shell layers formed of calcite crystals, as well as an absence of marked surface ornamentation. This eggshell form is typical of maniraptorans that are closer to crown-birds than to deinonychosaurs but outside of Neornithes, the crown-bird clade (Grellet-Tinner et al. 2006). Several exclusively Mesozoic bird lineages, enantiornithines among them, fit within this phylogenetic bracket. The data on eggshell microstructure is thus consistent with the osteological data from the accumulation (Dyke et al. 2012). Before anyone else says it, I should note that the phylogenetic distribution of certain eggshell characters remains the topic of debate – there are even cases where eggs confidently identified as those of maniraptoran theropods (Buffetaut et al. 2005) have turned out to be from squamates.

A waterside enantiornithine colony

Smaller version of Julio Lacerda's reconstruction.

We therefore hypothesise that the accumulation preserves the remains of an enantiornithine nesting colony, with the numerous eggshell fragments, the several complete eggs, and the various bones all belonging to the same one species.

Given the volume of eggshell, the colony must have been large. Assuming clutches similar in size to those of modern birds, we’re talking about hundreds of nests. The stacked, packed, jumbled nature of the eggshell fragments removes the possibility of this being an in-situ shared ‘mega-nest’ by the way (a bizarre possibility mooted by one reviewer!). There are no fossils of any other sort within the accumulation. It seems that a flood event swept across the colony, carrying eggs and birds (or their remains) before dumping them in a hollow area a short distance from the main channel (Dyke et al. 2012). This transport can only have occurred across a short distance (as in, several metres), since the bones are unabraded and several eggs are intact.

Waterside-nesters like plovers typically deposit their eggs in 'scrapes' - simple nests that don't involve the incorporation of vegetation or feathers. These are Ringed plover (Charadrius hiaticula) eggs. Photo by Arnoldius, licensed under Creative Commons Attribution-Share Alike 3.0 Unported license.

The form of the eggs is typical of ground-nesters that produce precocial young (Dyke et al. 2012). The absence of vegetation in the accumulation implies that the birds formed scrapes in the sediment for their eggs, as do plovers and some other waterside-nesting modern birds. [Adjacent image of Ringed plover eggs by Arnoldius.]

Given the amount of eggshell, why aren’t there more bones in the accumulation? The inner surfaces of the eggshell fragments are dull and etched, “suggesting that calcium mobilisation was advanced and that the eggs had either hatched or were destroyed late in the second half of incubation” (Dyke et al. 2012). In other words, it’s likely that many or most of the eggs present in the colony at the time of the flooding event were already hatched and lying, empty and discarded, in or close to their nests. This probably wasn’t the case for all nests, since the presence of both juvenile and adult bones implies that these birds perished when the colony flooded. Then again, it’s also conceivable that these bones belong to individuals that were already dead before the colony was swamped – it’s common to see the remains of dead chicks and even dead adults in nesting colonies.

Breeding colonies of marine birds, like these Cape gannets (Morus capensis), can be especially crowded. Photo by Octagon, licensed under Creative Commons Attribution 3.0 Unported license.

Assuming that we’re right in our overall interpretation, what does the discovery mean for enantiornithine nesting and breeding behaviour? A waterside nesting habit for an Enantiornis-like enantiornithine is not all that surprising, since there’s already morphological and stomach-content data showing that some members of this group were aquatic foragers, preying on crustaceans and other such prey (Sanz & Buscalioni 1992, Sanz et al. 1996). The fact that such a large nesting colony of this one enantiornithine species occurred in a waterside habitat strongly suggests reliance on aquatic resources, so the species concerned might have been gull-like or plover-like in ecology. [Adjacent photo of gannet colony by Octagon.]

And, like gulls, waterfowl, flamingos and other waterside, colonial nesters, it seems that these enantiornithines were sometimes unfortunate, and that local flooding events swamped and drowned their nest colonies (Dyke et al. 2012). Flooding occurs quite regularly in some modern bird colonies and is an expected hazard of nesting so close to water. There are even cases where the same colony gets flooded repeatedly during the same one breeding season (e.g., Peresbarbosa & Mellink 2001), and yet still the birds continue to nest there. Why do they nest in such dangerous places? Because they are otherwise highly suitable; more so than adjacent, well-vegetated regions in providing barren, relatively predator-free areas that are suitable for nesting, and are relatively close to the aquatic environment.

Detail of part of Julio Lacerda's reconstruction of the nesting colony. You'll note that Julio's enantiornithines look... like birds, not like the crazy feathered dragon-monsters depicted by some artists elsewhere.

On the one hand, the formation of large, waterside nesting colonies in Mesozoic birds is not all that surprising, since this behaviour is common and widespread in modern birds and has clearly evolved many times independently. On the other hand, enantiornithines are not typically imagined as being all that similar to modern, colony-nesting birds like gulls, terns, gannets, penguins and so on, and I’m not sure that I’d have guessed a colony-nesting habit for any enantiornithine prior to the discovery of the Od accumulation. For Mesozoic seabirds, like hesperornithines and Ichthyornis, sure, but – for enantiornithines – it’s a novel idea.

So we can now say that the latest Cretaceous Transylvanian Basin fauna of Romania was inhabited by peculiar, island-endemic dromaeosaurs, titanosaurian sauropods, both rhabdodontid and hadrosaurid ornithopods, azhdarchid pterosaurs, eusuchian crocodyliforms, pleurodiran turtles, and enantiornithine birds that formed enormous, waterside nesting colonies. More exciting Romanian finds are due to be announced in the near future. Special thanks to the outstanding Julio Lacerda for the excellent artwork he produced for this project. Julio has a DeviantArt page here and blogs at The Casual Paleoartist.

For previous Tet Zoo articles on Mesozoic birds, see…

• The new Crato Formation enantiornithine
• A stunning new Mesozoic bird… well, new-ish (Pengornis)
• The Mesozoic birds with weird, plastic-strip-style tail structures
• Alexornis and other ‘alexornithiforms’
• Aberratiodontus: worst paper ever?
• Luis Chiappe’s Glorified Dinosaurs: The Origin and Early Evolution of Birds
• Obscure Mesozoic birds you’ll only know about if you’re a Mesozoic bird nerd: Jibeinia luanhera

Refs – -

Buffetaut, E., Grellet-Tinner, G., Suteethorn, V., Cuny, G., Tong, H., Košir, A., Cavin, L., Chitsing, S., Griffiths, P. J., Tabouelle, J. & Le Loeuff, J. 2005. Minute theropod eggs and embryo from the Lower Cretaceous of Thailand and the dinosaur-bird transition. Naturwissenschaften 92, 477-482.

Csiki, Z., Vermir, M., Brusatte, S. L., Norell, M. A. 2010. An aberrant island-dwelling theropod dinosaur from the Late Cretaceous of Romania. Proceedings of the National Academy of Sciences 107, 15357-15361.

Dyke, G. Vremir, M. Kaiser, G. & Naish, D. 2012. A drowned Mesozoic bird breeding colony from the Late Cretaceous of Transylvania. Naturwissenschaften DOI:10.1007/s00114-012-0917-1

Elzanowski, A. 1981. Embryonic skeletons from the Late Cretaceous of Mongolia. Palaeontologica Polonica 42, 147-179.

Grellet-Tinner, G. & Norell, M. A. 2002. An avian egg from the Campanian of Bayn Dzak, Mongolia. Journal of Vertebrate Paleontology 22, 719-721.

-., Chiappe, L. M., Norell, M., Bottjer, D. 2006. Dinosaur eggs and nesting behaviours: A paleobiological study. Palaeogeography, Palaeoclimatology, Palaeoecology 232, 294-321.

Mikhailov. K. E. 1996. Bird eggs in the Upper Cretaceous of Mongolia. Palaeontology Journal 30, 114-116.

Peresbarbosa, E. & Mellink, E. 2001. Nesting waterbirds of Isla Montague, northern Gulf of California, México: loss of eggs due to predation and flooding, 1993-1994. The International Journal of Waterbird Biology 24, 265-271.

Sanz, J. L. & Buscalioni, A. D. 1992. A new bird from the Early Cretaceousof Las Hoyas, Spain, and the early radiation of birds. Palaeontology 35, 829-845.

- ., Chiappe, L. M., Pérez-Moreno, B., Buscalioni, A. D., Moratalla, J. J.,Ortega, F., Poyata-Ariza, F. J. 1996. An Early Cretaceous bird fromSpain and its implications for the evolution of avian flight. Nature 382, 442-445.

Schweitzer, M. H., Jackson, F. D., Chiappe, L. M., Schmitt, J. G., Calvo, J. O. & Rubilar, D. E. 2002. Late Cretaceous avian eggs with embryos from Argentina. Journal of Vertebrate Paleontology 22, 191-195.

Vremir, M. 2010. New faunal elements from the Late Cretaceous (Maastrichtian) continental deposits of Sebeş area (Transylvania). Acta Musei Sabesiensis 2, 635-684.

Zhou, Z. (2004). A Precocial Avian Embryo from the Lower Cretaceous of China Science, 306 (5696), 653-653 DOI: 10.1126/science.1100000

The white bag is there for a reason. Read on.

Matt Wedel kindly passed on the photos you see here. They show the Man-eater of Mfuwe, an enormous male lion Panthera leo that terrorised the small town of Mfuwe (and the surrounds) in the Luangwa River Valley of eastern Zambia. The photos were taken in Chicago’s Field Museum where the specimen has been on display since donation in 1998 (sorry about the glare – it’s impossible to take good photos without smashing into the case, and the museum generally discourages that sort of thing).

The history and behaviour of man-eating big cats is truly fascinating stuff; the stealth, cunning and occasional reckless confidence they display in getting at their victims, the level and extent of their depredations, and the stealth and cunning that human hunters have to use to put an end to these cats are frequently extraordinary and make fascinating reading. I regularly refer to the 1992 book Man-Eater: Tales of Lion and Tiger Encounters (edited by Edward Hodges-Hill). It includes articles on the Iyenpur tigress, the Jerangau, Champawat, Kempekarai and Mamandur man-eating tigers, the ‘Ogre of Bellundur’ (a tiger), the Tsavo lions, Jim Corbett’s adventures with the Thak man-eater, and many others (Hodges-Hill 1992).

Of all these killer cats, perhaps the most famous were the two male lions who killed a claimed 135 people* in what is now Tsavo National Park during the 1890s. The story of these two lions and their depredations have been the subject of three Hollywood movies, the best known of which is the 1996 The Ghost and The Darkness. An excellent and thorough book, Bruce D. Patterson’s The Lions of Tsavo: Exploring the Legacy of Africa’s Notorious Man-Eaters, discusses this episode in full, placing it within the context of modern research on lion behaviour, conservation and variation (Patterson 2004).

* This number (estimated by Colonel John Henry Patterson, the killer of the two lions) was questioned by Yeakel et al. (2009). Based on stable isotope analysis and estimates of calorific requirements, they reckoned that both lions killed about 34 people in total, a number substantially lower than Patterson’s estimate. Yeakel et al. (2009) might be right, but they might not.

It isn’t the only book on the Tsavo lions – there’s also Philip Caputo’s Ghosts of Tsavo: Stalking the Mystery Lions of East Africa (Caputo 2002). Both books discuss the Man-eater of Mfuwe; my text here is based almost entirely on their accounts.

The sign that accompanies the lion at the Field Museum. Don't read the text - it's a spoiler for the rest of this article...

The Mfuwe lion wasn’t killing and eating people back in Victorian times or anything like that. Rather, it was a modern man-eater, its attacks occurring in 1991. The first occurred as two boys were walking along a road at night. One boy fled; by the time game rangers arrived at the attack site, only pieces of clothing and part of the other boy’s skull were found. The second attack occurred at the edge of a village – this time the victim was an adult woman; the lion had broken through the door of her hut to get her. The third attack (occurring at night on a boy who ventured out to meet a friend) was foiled when a game scout fired his gun into the air during the attack, but the boy died of his severe injuries anyway.

Three more kills occurred later in 1991, the last one again involving a woman being dragged from the hut where she lived. Prior to this last kill, it was locally believed that lionesses belonging to the ‘L-pride’ were responsible for the killings, and indeed one of these lionesses was shot dead in August of 1991. The important role of an adult male lion now came to the fore. This lion entered the woman’s hut during daylight and removed a bag containing her laundry. It took the bag to the centre of the village, dropped it, and stood over the bag, roaring. Remarkably, the lion carried the bag about the countryside, leaving it at different locations and sometimes playing with it. Unsurprisingly, perhaps, people now suspected that the lion was no ordinary lion, but a demon or sorcerer in lion form.

Some sources say that people weren’t allowed to hunt and kill this lion because Zambia’s laws about game conservation didn’t allow it. That’s not wholly correct, given the killing of one of the suspected man-eating lionesses by game wardens (they also shot a juvenile male lion at the same time, but didn’t kill it). The other females in the pride were also shot dead later on.

The Mfuwe man-eater’s reign was put to an end by Californian hunter Wayne Hosek. Hosek wasn’t the first to try and kill the Mfuwe lion – a professional hunter, and a Japanese hunter and naturalist, had both tried earlier. Hosek was already in the area with permission to hunt individuals from a list of large mammal species, lion among them. There’s the whole debate here about controlled hunting and its contribution to economy and conservation versus its abuse at the hands of the unscrupulous, the selective impact it has on animal populations, and the ethics of trophy-hunting. I’m somewhat mystified by the idea that anyone can look at a lion, elephant or hippo and feel compelled to shoot it to death, but my personal perspective on this matter is that giving live wildlife an actual dollar value can only be a good thing, at least if you want live wildlife to still be present in future decades.

Anyway, Hosek and professional hunter Charl Beukes (or Buekes? Both spellings are used in the literature) sat for about three weeks in a hide, hoping that a nearby bait of hippo meat would attract the cat and allow them to kill it. Early attempts were unsuccessful – the lion even circled the bait one night, but was so stealthy that it had avoided all detection. Eventually, Hosek succeeded in killing it. On the lion’s death, people swarmed out of the village to spit at the animal and beat it with sticks.

The Mfuwe lion, today on display at the Field Museum in Chicago, is enormous: 3.2 m in total length, 1.2 m tall at the shoulder, and with a mass estimated at 249 kg (Caputo 2002). And, contrary to local descriptions of it being equipped “with a huge mane” (Caputo 2002, p. 3), it’s wholly maneless.

Manelessness in lions – the competing hypotheses

Patterson wrote that Hosek was due to write a book about the Mfuwe man-eater; so far as I can tell this hasn’t happened, but do say if you know otherwise. If you’ve read Patterson’s or Caputo’s books, or if you know a lot about lions in general, you’ll know that the Man-eater of Mfuwe was similar to the man-eating male lions of Tsavo in being maneless. In fact some witnesses did say exactly this, but their descriptions of a maneless lion were assumed to be of a lioness, hence the deliberate targeting of lionesses in efforts to eliminate the man-eater.

One of the two Tsavo lions, with Colonel John Henry Patterson (no direct relation to Bruce D. Patterson, author of The Lions of Tsavo). The image is in the public domain.

Why is manelessness prevalent among lions of this region? Actually, this is a rather controversial subject and numerous hypotheses have been entertained: for a thorough, fully referenced discussion, see Patterson (2004). Perhaps the most intriguing suggestion is that the lions concerned represent a distinct, ancient lineage, phylogenetically separate from other living lions and perhaps being relict, Pleistocene-type lions (Gnoske & Kerbis Peterhans 2000). The authors who promote this hypothesis have used the term ‘Buffalo lions’ for these animals (von Buol 2000) since they describe them as specialised predators of Cape buffalo and other especially big prey. It has also been called the ‘basal lion hypothesis’ (Patterson 2003). Molecular and morphological data does not support this idea: DNA and morphometrics both show that Tsavo-type, maneless lions are deeply nested within Panthera leo, being close to other east and south African lion populations (Patterson 2004, Dubach et al. 2005). [Maneless lion photo below by Mgiganteus.]

Maneless Tsavo lion, photographed in Tsavo East National Park, Kenya, 2007, by Mgiganteus. Licensed under the Creative Commons Attribution-Share Alike 3.0 Unported license.

The possibility that lions have lost manes because manes are physiologically costly in some environments has also been entertained, as has the idea that manes are reduced and lost in habitats where they provide a disadvantage to moving about among thorn-scrub. Then there’s the idea the maneless lions are old and poor in condition (Patterson (2004) called this the ‘down-and-out hypothesis’), and the alternative idea that their manelessness results from especially high levels of testosterone (that’s right, high levels of testosterone causes hair follicles to shrink: hair reduction and loss is the result). Given that Tsavo-type lions are evolving within a social system where single males, rather than coalitions, associate with female prides, it might be that a particular male physiology is driving manelessness. But, as usual, it seems that several factors are at play: thorn brush and a hot, dry climate might contribute to mane reduction in Tsavo-type lions (Kays & Patterson 2002, Gnoske et al. 2006, Patterson et al. 2006) as well. This issue was previously discussed in the ver 2 article Why the Lion Grew Its Mane, a book review.

Why become a man-eater?

Jim Corbett and the Champawat tigress, supposedly the killer of 436 people until shot by Corbett in 1907. This was only the first of 19 man-eating tigers and leopards that Corbett killed during his life. Photo in public domain.

As much as I’d like to continue discussing variation within lions, my time is up, and this is still a topic I plan to return to at length. One more thing is worth saying: why do some big cats become bold man-eaters in the first place?

It’s a well known bit of lore that man-eating big cats are sometimes injured or sickly individuals that take to killing humans because it’s easy, and because the animals are desperate enough to disregard normal caution or fear. The Mfuwe lion was described as looking “a little green around the gills” when seen in the field, and had sustained a major injury to the lower jaw that had resulted in pustulous lesions (Patterson 2004, pp. 76-77). The two Tsavo lions both had craniodental injuries that might have prevented normal predatory behaviour (Yeakel et al. 2009). However, this ‘infirmity hypothesis’ doesn’t explain all cases, since many problem big cats have been perfectly healthy.

Other individuals become bold and experienced following a chance encounter with an unfortunate human, and then deliberately target people in future. Others switch to humans as prey because funeral rites, atrocities resulting from slavery and so on mean that human remains left out in the open are opportunistically scavenged and hence ‘turn on’ a culture of human-eating in some individuals or groups. Finally, shortages of natural prey caused by drought, disease epidemics and extirpation by people also seem to cause lions, tigers and other big cats to switch to humans through desperation.

The two Tsavo lions, as displayed today at the Field Museum. Photo by Jeffrey Lung, licensed under Creative Commons Attribution-Share Alike 3.0 Unported, 2.5 Generic, 2.0 Generic and 1.0 Generic license.

Human encroachment on big cat habitat, the invasion of habitat by domestic livestock and by people hunting for bushmeat – in short, human disruption of ecosystems – means that big cats both become more familiar with people as potential prey items, and are increasingly pushed into a corner where humans become ever more desirable and available prey items. And some or all of these phenomena might combine, meaning that the cause of man-eating behaviour is not exactly simple. Of incidental interest is that human predators have sometimes used outbreaks of man-eating in lions and other cats to conceal their own murderous habits.  Of the several creepy stories on record, perhaps the weirdest is that concerning the ‘lion-men of Singida’. These were actually drugged young women “sewn into animal hides, and fitted with deadly prosthetic claws” (Patterson 2004).

For previous articles on lions and other cats, see…

• Belated welcome to a ‘new’ clouded leopard.. named in 1823
• Peter Hocking’s big cats: where are you now?
• So what was that mysterious black gracile felid?
• Europe, where the sabre-tooths, lions and leopards are
• Pumas of South Africa, cheetahs of France, jaguars of England
• Giraffe-killing lions exploit paved roads
• Why the Lion Grew Its Mane, a book review
• Super-size cougars
• The Pogeyan, a new mystery cat
• ‘Revising’ the Siberian tiger
• Leopard cats: exotic and (sometimes) wild in the UK
• Dissecting lions and tigers: Inside Nature’s Giants series 2, part III
• Big spotted pumas… Miracinonyx redux?
• Williams and Lang’s Australian Big Cats: do pumas, giant feral cats and mystery marsupials stalk the Australian outback?

Refs – -

Caputo, P. 2002. Ghosts of Tsavo: Stalking the Mystery Lions of East Africa. National Geographic Society, Washington, D. C.

Dubach, J., Patterson, B., Briggs, M., Venzke, K., Flamand, J., Stander, P., Scheepers, L., & Kays, R. (2005). Molecular genetic variation across the southern and eastern geographic ranges of the African lion, Panthera leo Conservation Genetics, 6 (1), 15-24 DOI: 10.1007/s10592-004-7729-6

Gnoske, T. P., Celesia, G. G. & Kerbis Peterhans, J. C. 2006. Dissociation between mane development and sexual maturity in lions (Panthera leo): solution to the Tsavo riddle?Journal of Zoology 270, 551-560.

- . & Kerbis Peterhans, J. 2000. Cave lions: the truth behind biblical myths. In The Field 71, 2-6.

Hodges-Hill, E. (ed) 1992. Man-Eater. Tales of Lion and Tiger Encounters. Cockbird Press Ltd, Heathfield.

Kays, R. W. & Patterson, B. D. 2002. Mane variation in African lions and its social correlates. Canadian Journal of Zoology 80, 471-478.

Patterson, B. D. 2004. The Lions of Tsavo: Exploring the Legacy of Africa‘s Notorious Man-Eaters. McGraw-Hill, New York.

- ., Kays, R. W., Kasiki, S. M. & Sebestyen, V. M. 2006. Developmental effects of climate on the lion’s mane (Panthera leo). Journal of Mammalogy 87, 193-200.

von Buol, P. 2000. ‘Buffalo’ lions. A feline missing link? Swara: the Magazine of the East African Wildlife Society 23 (2), 20-25.

Yeakel, J. D., Patterson, B. D., Fox-Dobbs, K., Okumura, M. M., Cerling, T. E., Moore, J. W., Koch, P. L. & Dominy, N. J. 2009. Cooperation and individuality among man-eating lions. Proceedings of the National Academy of Sciences 106, 19040-19043.

The complex structure, development and growth of feathers can, to paraphrase one expert on the subject, be seriously damaging to your mental health. Feathers are just crazy, almost certainly the most complex structures to ever grow out of any animal’s external surface.

Yet for all their marvellous complexity, for all the interest that people have displayed in their evolutionary origins and diversity, for all their role in bird behaviour and ecology, and for all their economic and cultural significance to humans, it doesn’t seem that any one book has ever been devoted to feathers and feathers alone. Thor Hanson’s 2011 Feathers is thus a rather significant book, and very nice it is too.

Hanson, a Washington State-based conservation biologist, previously wrote The Impenetrable Forest: My Gorilla Years in Uganda. He’s published technical research on such topics as the ecology of tropical trees, forest fragmentation and its impact on bird nest predation, the impact that warfare can have on biodiversity hotspots, and the behaviour of Neotropical monkeys and birds.

Hanson might not be a feather specialist, or even a dedicated ornithologist, but his many encounters with feathers, and with their structure, role, significance and uniqueness, obviously created the urge that culminated in this book. And Feathers is not the provincial view of someone only interested in ecology or conservation biology; on the contrary, this is a remarkably well-rounded review of the subject. Chapters look at the evolution and origin of feathers, their use in flight, thermoregulation and display, and their importance to humans.

Feathers from the Mesozoic

♥ Alan Feduccia.

A discussion of feathered non-avialan theropod dinosaurs and of feather origins make up early sections in the book. Hanson is very much up to date (as of 2011!), providing appropriate discussion of Richard Prum’s feather origins hypothesis (Prum & Brush 2002) and of Xu Xing and his numerous exciting feathered theropod discoveries. As per usual, we get the theropod origins model pitted against Alan Feduccia’s idea that birds just cannot be theropods, nor dinosaurs at all, but an independent derivation from non-dinosaurian diapsid reptiles of some sort. Feduccia is quoted as saying in the book that “If it has feathers, it’s a bird” (Hanson 2011, p. 57), which of course means that feathered oviraptorosaurs, deinonychosaurs and so on are now birds according to Feduccia. As Hanson notes, and as Prum and others have said before (Prum 2003), this means that Feduccia is now contradicting decades of bold assertion in which he has insisted that deinonychosaurs and other Mesozoic theropods are nothing whatsoever to do with birds. It must be understood that Feduccia’s opinion is not a valuable, informed alternative or anything like that; rather, it relies on deliberate obfuscation and misinformation and ignorance with respect to what we actually know. I cannot see that he and his colleagues have done anything but add confusion, contradiction and erroneous interpretations to our understanding of bird origins and early evolution. Hanson was able to see through this, as have previous authors who approached the topic of bird origins as apparent outsiders (e.g., Shipman 1998).

It’s often difficult to entangle the difference between the origin of birds and the origin of flight, but recently discovered small non-avialan maniraptorans and the distribution of vaned feathers in maniraptoran phylogeny may well show that flight of a sort was present in whatever taxa were ancestral to both the bird and deinonychosaur lineages. While the majority of non-avialan theropods were non-climbers, could climbing or branch-leaping, or some other way of acquiring height, help explain the earliest steps in the evolution of flight?

One of Ken Dial's WAIR diagrams. This figure shows that wing stroke is nearly invariant to gravity; the red lines represent the wing-tip trace in WAIR while the blue lines represent the wing-tip trace in level flight.

Hanson provides nice coverage of this topic, and is especially keen on Ken Dial’s Wing Assisted Incline Running (WAIR) hypothesis (Dial, Bundle & Dial 2003, Dial et al. 2006, 2008). As Hanson notes, WAIR has been embraced rather enthusiastically by palaeontologists “as the best flight evolution story to date” (p. 129). It has appeal in explaining how small, incremental steps might explain how terrestrial theropods with an early form of feathering could have evolved into scansorial animals with larger, increasingly complex feathery surfaces on the limbs. I think that WAIR is a pretty neat hypothesis. However, I note that some who work on flight origins are highly sceptical of it, stating in particular that non-ornithurine birds lacked the bony architecture required to permit the vigorous, high-amplitude flapping needed for WAIR to function.

The 1911 Trans-Saharan Ostrich Expedition, Vegas showgirls, and other feather-based tales from the human world

Frontispiece to Richard Brookes's 1790 volume The Art of Angling.

While all of these topics might be quite familiar if you’re well-read on bird and feather origins, Hanson’s other chapters cover feather growth and moulting, avian thermoregulation, the down industry, the aerodynamic and water-repellent qualities of feathers, and the use of feathers in display (both among birds and among humans), fly-fishing and writing.

The anthropological sections were the most novel. We’re all familiar with the use (or former use) of feathers in hats, dusters, cloaks and so on, but overall this subject certainly isn’t something I’ve ever had the chance to read much about. Most people interested in birds know that the feather trade was so substantial during the early decades of the 20th century that egrets and other species were being slaughtered in their millions expressly for the purpose, and indeed that conservation bodies, laws and reserves were created in direct response to the scale and nature of this trade. Feathers were so valuable at this time that the more than 40 cases of plumes lost on the Titanic would have an insurance value of more than $2.3 million in today’s currency (Hanson 2011, p. 176).

Ostriches plucked of ALL their feathers. This image shows ostriches that belonged to Shua people in Nigeria; it's from Hugo Bernatzik's 1931 book The Dark Continent; Africa, the Landscape and the People (digitized by the New York Public Library).

One of my favourite sections in the book is that on the Trans-Saharan Ostrich Expedition, whereby Russel William Thornton led a 1911 expedition from South Africa to Nigeria in order to track down, capture and exploit the fabled Barbary ostrich. The plumes of this ostrich are (or were) ‘double-flossed’ and more luxuriant than those of other ostriches. I’m somewhat uncertain of the taxonomic status of this form but, then, I think that everyone might be, since the population concerned is apparently now extinct. It has at least been suggested that ‘Barbary ostriches’ were a local form of the North African, Sudan or Red-necked ostrich Struthio camelus camelus (Freitag & Robinson 1993), but I don’t know if this has ever been demonstrated. Anyway, you might be as surprised as I was to learn that captured ostriches were plucked of all their feathers.

Stuff I saw in Las Vegas. You should be able to see pink plumes on the photo of the showgirl at top right.

Hanson’s section on the role of feathers in display talks about birds-of-paradise, sexual display, the ‘sexy son’ hypothesis, Wallace and Darwin. But he goes on to discuss Las Vegas showgirls and the tradition of using showy, enormous feather plumes in dancing costumes. Some of the largest feathery pieces used in showgirl costumes might cost “tens of thousands of dollars” should they need to be replaced (Hanson 2011, p. 172). I was actually reading the book while staying in Las Vegas, and got to walk past the famous pink-plumed dancing girls of Bally’s Hotel and Casino on a twice-daily basis. It was weird… like everything else in Las Vegas.

Hanson also covers the massive use of birds by the Aztecs. The aviaries at Tenochtitlán were described by Hernán Cortés as being enormous, housing every kind of bird and requiring the services of 300 human attendants. They were completely burnt down during the Spanish siege of Tenochtitlán in 1521.

On vultures

Vultures play an important role in the book since Hanson says in the preface that they’re what got him interested in feathers in the first place. He relates a delightful tale from field research where he had to handle a rotting zebra caecum, destined for use at a vulture feeding station. It exploded, “blowing my hair back and coating me with a spray of old blood, strands of ropy goo, and flecks of half-digested bush grass. The smell was indescribable.” (Hanson 2011, p. 250).

Thermographic image of a vulture (a cathartid, probably Cathartes aura). Image by Arno/Coen, licensed under Creative Commons Attribution-Share Alike 3.0 Unported license.

Hanson mostly includes vultures in the book because he argues that the reduced or absent feathering on their heads is an adaptation for carrion-feeding. This might be partly true, but in recent years it’s been questioned: some birds that regularly poke their heads into carcasses (giant petrels) get by fine with normal feathering, most vultures are not really naked-headed anyway, and it might be that reduced feathering on the head and neck is as much (or more) to do with thermoregulation than carrion feeding (Ward et al. 2008). Hanson also commits an oft-made mistake in writing that New World vultures are especially closely related to storks. This view was mooted a few decades ago but hasn’t been supported by any of the more recent studies published on bird phylogeny.

White-backed vulture (Gyps africanus) in flight. Image by Mark Rosen, licensed under Creative Commons Attribution-Share Alike 3.0 Unported license.

I can’t leave this mention of vultures without referring to the continuing decline of these magnificent birds due to human greed, stupidity and laziness. The die-off of Indian vultures due to use of the veterinary drug diclofenac is well known: the decline has been so rapid that, in 2008, some experts were predicting the extinction of certain vulture species within a ten-year period. Despite being banned from sale in 2005, diclofenac is still being sold illegally and some reports say that it’s continuing to have a deleterious effect. Vulture ‘restaurants’ set up by the Indian government have not been adequately funded or maintained, and an article from February of this year said that these feeding stations haven’t benefited vulture conservation at all. A symposium on the continuing decline of Asian vultures was in fact held just a few days ago (early May 2012).

The problems in India (which mostly effect the Slender-billed vulture Gyps tenuirostris, Indian vulture G. indicus and White-rumped vulture G. bengalensis) have received a reasonable amount of press interest. Less well known is that massive declines seem to have occurred in Africa as well, with some data from Mali and Niger indicating that vulture species have either disappeared entirely, or have declined by something like 98%. Here, diclofenac is less of a problem. Habitat destruction and degradation, poisoning, and killing for use in traditional and quack medicine – known as muti – have all contributed to decline. Particularly infuriating is the belief that smoking vulture brains will give people supernatural powers when gambling (yes, I said smoking vulture brains will give people supernatural powers when gambling). The demand for vulture products used in muti has soared in step with the growth of national lotteries and with events such as the 2010 World Cup, hosted by South Africa (Langley 2011). Clearly, the health of vulture populations across many areas is now a major cause for concern and urgent measures are needed.

Anyway… I digress, back to the book review. Other highlights include the discussion of the peregrine feather dropped on the surface of the moon by Commander David R. Scott in 1971, and the section on snarge. Yes, I said snarge, and will leave it at that.

Feathers is a handy, compact volume, well illustrated throughout, and with a compelling, enjoyable prose. It’s fully referenced, with footnotes (arranged at the back) providing details that would have derailed or slowed the main flow of the text. Appendices discuss and illustrate the different kinds of feathers. I appreciated and agreed with Hanson’s perspective on all the areas he discusses, I enjoyed and respected the enthusiasm and sense of wonder that he conveys when talking about the natural world, and I congratulate him on thorough research and on being up-to-date on such a fast-moving topic. In short, Feathers is required reading for anyone interested in bird biology or evolution, and I strongly recommend it.

Hanson, T. 2011. Feathers: The Evolution of a Natural Miracle. Basic Books, New York, pp. 336. ISBN 978-0-465-02013-3. Hardback, index, refs. Here on amazon. Here on amazon.co.uk.

For previous articles relevant to feathers, early bird evolution and other topics mentioned in this article, see…

• Feathers and filaments of non-avian dinosaurs, part I
• Feathers and filaments of dinosaurs, part II
• Epidexipteryx: bizarre little strap-feathered maniraptoran
• Long and Schouten’s Feathered Dinosaurs, a review
• Gary Kaiser’s The Inner Bird: Anatomy and Evolution
• Condors and vultures: their postures, their ‘bald heads’ and their sheer ecological importance
• Luis Chiappe’s Glorified Dinosaurs: The Origin and Early Evolution of Birds
• There are giant feathered tyrannosaurs now… right?

Refs – -

Bundle MW, & Dial KP (2003). Mechanics of wing-assisted incline running (WAIR). The Journal of experimental biology, 206 (Pt 24), 4553-64 PMID: 14610039

Dial, K. P. 2003. Wing-assisted incline running and the evolution of flight. Science 299, 402-404.

- ., Jackson, B. E. & Segre, P. 2008. A fundamental avian wing-stroke provides a new perspective on the evolution of flight. Nature 451, 985-989.

- ., Randall, R. J. & Dial, T. R. 2006. What use is half a wing in the ecology and evolution of birds? BioScience 56, 437-455.

Freitag, S. & Robinson, T. J. 1993. Phylogeographic patterns in mitochondrial DNA of the Ostrich (Struthio camelus). The Auk 110, 614-622.

Hanson, T. 2011. Feathers: The Evolution of a Natural Miracle. Basic Books, New York.

Langley, N. 2011. Africa’s emptying skies. World Birdwatch 33 (2), 16-18.

Prum, R. O. 2003. Are current critiques of the theropod origin of birds science? Rebuttal to Feduccia (2002). The Auk 120, 550-561.

- . & Brush, A. H. 2002. The evolutionary origin and diversification of feathers. The Quarterly Review of Biology 77, 261-295.

Shipman, P. 1998. Taking Wing. Weidenfeld & Nicolson, London.

Ward, J., McCafferty, D. J., Houston, D. C. & Ruxton, G. D. 2008. Why do vultures have bald heads? The role of postural adjustment and bare skin areas in thermoregulation. Journal of Thermal Biology 33, 168-173.

Heloderma suspectum, the Gila monster. Photo by Dave Hone.

The Gila monster Heloderma suspectum and its close relative the Mexican Beaded lizard H. horridum are the only two extant members of Helodermatidae, the gila monster clade.  It’s been agreed for a considerable time that, among living lizards, helodermatids are most closely related to monitor lizards (varanids) and to the weird Bornean earless monitor Lanthanotus borneensis. All of these lizards (and their close fossil relatives) are grouped together in Platynota. The name Varanoidea has been used for Platynota at times, but it’s been restricted specifically to the monitor lineage within Platynota at times, and hence, annoyingly, means different things to different authors.

Beaded lizard, photographed by Ltshears, from wikipedia. Released into public domain.

Helodermatids are stocky, proportionally short-limbed, short-tailed lizards with deep, robust snouts, dermal armour, osteoderms fused to the skull surface, venom grooves in the dentary teeth, and venom glands located in the lower jaw. Their teeth are formidable – check out the fantastic image below from digimorph. Gila monsters are normally 30-50 cm long in total while Beaded lizards can reach 70 cm or even nearly 1 m (Pianka & Vitt 2003). They prey on a diversity of invertebrates and small vertebrates, do a lot of digging for prey, and are sometimes regarded as specialised nest-seekers: they eat the eggs of other lizards as well as those of snakes and birds, and also prey on rodent nestlings. Gila monsters have been known to locate hen’s eggs buried 15 cm below the ground surface (Pianka & Vitt 2003). Needless to say, their olfactory and vomeronasal senses are highly developed. The venom is used for self-defense – not in subduing prey. They are surprisingly good climbers (albeit slow, cautious climbers) and may regularly forage in trees, bushes and even on cacti.

In general, these are slow, sluggish lizards that may spend about 95% of their time resting and hiding in burrows, but they are known to have one of the highest aerobic scopes of all lizards – this is higher in males than in females and is presumably a sexually selected trait related to the prolonged and metabolically costly wrestling that males engage in (Beck et al. 1995). In Beaded lizards at least, this behaviour is pretty spectacular and similar to the wrestling practised by some monitors: the animals lever off the ground with their heads and tails, forming an arch shape. The aim is then to push the opponent onto his back before biting him on the jaws (Beck & Ramírez-Bautista 1991). While we know that males engage in ‘sexual wrestling’, relatively little is known about social behaviour in general and more work is needed.

Gila monster skull, with numerous osteoderms, from digimorph (c).

Unambiguous fossil helodermatids are known from the Eocene-Oligocene of France (Eurheloderma) and the Oligocene and Miocene of the USA (Lowesaurus and H. texanum). A number of additional taxa from the Cretaceous, argued by some authors to belong together as the ‘gobidermatids’ (Lee 1997), have at times been included within Helodermatidae but are here excluded from that clade following Conrad (2008). In proportions and probably overall appearance, unambiguous fossil helodermatids seem to have been similar to the extant ones, and Eurheloderma and Lowesaurus definitely have venom grooves on their dentary teeth (Nydam 2000).

Skull of Cretaceous monstersaurian Estesia, photographed at the AMNH by Ghedoghedo. Released into public domain.

A number of additional fossil taxa group closer to helodermatids than to other platynotans, and yet don’t necessarily look helodermatid-like. Estesia mongoliensis from the Late Cretaceous of Mongolia, for example, had a shallow snout compared to gila monsters and probably looked superficially more like a monitor. Anatomical characters that link it and other species with helodermatids include tall, narrow neural spines and a distinctive form of tooth implantation.

Platynotan phylogeny plotted against time, from Conrad (2008). The clade shown at the top (the one including Heloderma) is Monstersauria. The one in the middle (including Varanus) is Goannasauria. See Conrad (2008) for a huge amount of additional detail. Mosasaurs and kin may or may not be part of Platynota.

In differentiating this ‘gila monster lineage’ from other groups within Platynota, Norell & Gao (1997) decided to name it.  They went with Monstersauria. Their original definition was node-based (Gobiderma + Heloderma), but Conrad (2008) more recently co-opted the name for the entire gila monster branch (an intention hinted at by Norell & Gao (1997), though not expressed in their node-based definition). So, Estesia, the ‘gobidermatids’, and of course helodermatids proper are monstersaurians.

Monstersauria is the sister-group to the lineage that includes monitors, Lanthanotus and a large number of fossil taxa. Perhaps surprisingly, this lineage has no name unambiguously associated with it, and one was needed. Varanoidea has been applied to it by some authors, but the use of this name for the entire gila monster + monitor clade seems more appropriate. Conrad (2008) therefore came up with a new name for the monitor branch, and opted for Goannasauria. So, Varanoidea – the entire gila monster + monitor clade within Platynota – consists of two major lineages, Monstersauria and Goannasauria.

And this, again, is why I suck at producing ‘picture of the day’ posts. Dammit.

For previous Tet Zoo articles on platynotans, see…

• Hell yes: Komodo dragons!!!
• Of giant plated lizards and rough-necked monitors
• Dinosaurs come out to play (so do turtles, and crocodilians, and Komodo dragons)
• What I saw at the zoo yesterday… (more brief comments on Komodo dragons)
• Perentie tries to swallow echidna. Echidna too spiky, Perentie gets horribly injured. Dies.

Mosasaurs – historically associated with monitors or even with snakes – have been included within Platynota by some authors (e.g., Lee 1997). This position is looking increasingly unlikely. That is, mosasaurs are probably not platynotans. Nevertheless, if you want to see some Tet Zoo articles on them, go to…

• Tongues, venom glands, and the changing face of Goronyosaurus
• Mosasaurs might have used the same microscopic streamlining tricks as sharks and dolphins
• The gigantic, shark-toothed, small-flippered, long-bodied, sea-going predatory lizard that is Hainosaurus

Refs – -

Beck, D. D., Dohm, M. R., Garland, T., Ramírez-Bautista, A. & Lowe, C. H. 1995. Locomotor performance and activity energetics of helodermatid lizards. Copeia 1995, 577-585

- . & Ramírez-Bautista, A. 1991. Combat behavior of the beaded lizard, Heloderma h. horridum, in Jalisco, Mexico. Journal of Herpetology 25, 481-484.

Conrad, J. (2008). Phylogeny And Systematics Of Squamata (Reptilia) Based On Morphology Bulletin of the American Museum of Natural History, 310, 1-182 DOI: 10.1206/310.1

Lee, M. S. Y. 1997. The phylogeny of varanoid lizards and the affinities of snakes. Philosophical Transations of the Royal Society of London B 352, 53-91.

Norell, M. A. & Gao, K. 1997. Braincase and phylogenetic relationships of Estesia mongoliensis from the Late Cretaceous of the Gobi Desert and the recognition of a new clade of lizards. American Museum Novitates 3211, 1-25.

Nydam, R. L. 2000. A new taxon of helodermatid-like lizard from the Albian-Cenomanian of Utah. Journal of Vertebrate Paleontology 20, 285-294.

Pianka, E. R. & Vitt, L. J. 2003. Lizards: Windows the Evolution of Diversity. University of California Press, Berkeley.

Is an albatross (like the Wandering albatross (Diomedea exulans) shown in the middle) more like a petrel (like the Great shearwater (Puffinus gravis) shown at left) or like a storm-petrel (like the Wilson's storm-petrel (Oceanites oceanicus) shown on the right)? Images by Patrick Coin (l and r) and JJ Harrison (centre), licensed under Creative Commons Attribution-Share Alike 3.0 Unported license and Creative Commons Attribution-Share Alike 2.5 Generic license.

Petrels (by which I mean ‘true petrels’) have conventionally been regarded as a ‘family’ (termed Procellariidae) within the ‘order’ Procellariiformes (popularly termed tubenoses). But petrels aren’t the only tubenose ‘family’. Before moving on to look at petrel phylogeny and diversity in full, glorious detail, my plan here is to look at the position of petrels within tubenose phylogeny as a whole. How are petrels related to other tubenoses? And, if we know that, what might it tell us about tubenose evolution as a whole?

The ‘four families’ system breaks down

Shy albatross (Thalassarche cauta), 'super petrel' or something else entirely? Those interested in the soft-tissue reconstruction of fossil archosaurs will already be enamored with the cheek anatomy of this (and a few other) albatross species. Photo by JJ Harrison, licensed under Creative Commons Attribution-Share Alike 3.0 Unported license.

I grew up with the idea that petrels were especially closely related to the largest, most spectacular and most famous of tubenosed seabirds, the albatrosses. In fact my impression as a rather younger person was that albatrosses were ‘super petrels’, a notion mostly based on Sibley & Ahlquist’s (1990) contention that the two groups were closely related ‘subfamilies’ that only diverged about seven million years ago. Today it seems that albatrosses are not especially close to petrels proper, and indeed views on the phylogeny of tubenoses as a whole don’t support the ‘family-level’ classification used in most 20th century texts, partly because storm-petrels (hydrobatids) seem non-monophyletic (Nunn & Stanley 1998, Kennedy & Page 2002, Hackett et al. 2008).

Interested in testing correlations between body size and rates of molecular evolution, Nunn & Stanley (1998) produced what I think was the first large-scale molecular analysis of tubenoses. As just mentioned, storm-petrels were not recovered as monophyletic, with hydrobatines (one of two storm-petrel clades) being closer to the remaining lineages than was the other storm-petrel clade. The next major divergence involved albatrosses and a diving-petrel + true petrel clade. Kennedy & Page (2002) showed via a supertree approach that studies as of that time generally supported the following topology: (hydrobatine storm-petrels + (albatrosses + (oceanitine storm-petrels + true petrels))). Diving-petrels were deeply nested within true petrels. Seeing as diving-petrels were generally regarded then as a ‘family’ (Pelecanoididae), finding them nested within ‘family’ Procellariidae (true petrels) made Procellariidae non-monophyletic as well.

Images by (l to r) Bryan Harry, Sannab, Patrick Coin, Júlio Reis and James Lloyd; images licensed under Creative Commons Attribution ShareAlike 3.0 License, Creative Commons Attribution-Share Alike 3.0 Unported license, Creative Commons Attribution-Share Alike 2.5 Generic license, and Creative Commons Attribution-Share Alike 3.0 Unported license.

Penhallurick & Wink (2004) used mitochondrial cytochrome b sequence data to look both at higher-level relationships among tubenoses, and to assess the classification of taxa at the genus, species and subspecies level. Again, storm-petrels were recovered as non-monophyletic, but albatrosses now grouped with the two storm-petrel clades, being closer to hydrobatines in their favoured tree. Shock horror. Prions, diving-petrels and true petrels formed the sister-group to the storm-petrel + albatross clade, though the position of prions (conventionally included within true petrels) was fairly labile (Penhallurick & Wink 2004). In their favoured phylogeny, diving-petrels were nested within true petrels, being especially close to gadfly petrels. Ericson et al. (2006) found albatrosses to be outside a (storm-petrel + (petrel + diving-petrel)) clade.

Click to enlarge. Attributions as above but prion image by Rosemary Tully, licensed under Creative Commons Attribution-Share Alike 3.0 Unported license.

Livezey & Zusi (2007) didn’t include many taxa in their famously large analysis, and – in contrast to most other recent studies – found true petrels and albatrosses to be sister-groups, with prions, the storm-petrel Oceanites and diving-petrels to be successively more distant to this clade. It has been argued that at least some of the character codings used by Livezey & Zusi (2007) are inaccurate, with taxa being coded for characters that they ‘should’ have, rather than what they do have (Mayr 2008). Furthermore, some of the relationships they recovered (e.g., a loon-grebe clade) are worryingly ‘traditional’.

Finally (for now), Hackett et al. (2008) found the oceanitine storm-petrel Oceanites to be outside a tubenose clade that includes albatrosses, hydrobatine storm-petrels, diving-petrels and true petrels.

Tubenose phylogeny recovered by Hackett et al. (2008). Image attribution as above.

Some implications, and what do the fossils say?

What ‘consensus’ emerges from these studies, and what does it actually mean that’s actually, you know, interesting? For starters, the tidy, apparently traditional view that Procellariiformes consists of (1) Hydrobatidae (storm-petrels), (2) Pelecanoididae (diving-petrels), (3) Procellariidae (true petrels) and (4) Diomedeidae (albatrosses) does not accurately reflect phylogeny.

Firstly, ‘Hydrobatidae’ consists of two distinct clades and is not a natural group. Secondly, while diving-petrels remain monophyletic, their classification as a ‘family’ will now confuse many given that they might be nested within true petrels. As noted above, Penhallurick & Wink (2004) advocated the view that diving-petrels are within a true petrel clade that also includes gadfly petrels, and they thus treated diving-petrels as a ‘tribal-level’ clade (termed Pelecanoidini) within a ‘subfamily-level’ clade (termed Pelecanoidinae) within ‘family’ Procellariidae.

Tubenose phylogeny from Penhallurick & Wink (2004), with clades labelled. I don't expect for a moment that you'll be able to read much of the text here, but you should be ale to see that they classified all crown-tubenoses as either within Diomedeidae, or within Procellariidae. Actually, in the tree shown here, prions are incertae sedis, but they were recovered as part of Procellariidae in other trees.

Penhallurick & Wink’s (2004) phylogeny – while not exactly a ‘last word’ on the subject – would actually make the big picture of crown-tubenose phylogeny simpler than has been conventional. This is because members of the tubenose crown-group are, in their phylogeny, either members of the albatross lineage, or members of the true petrel lineage: Penhallurick & Wink’s (2004) therefore proposed that both kinds of storm-petrel should be included within Diomedeidae alongside albatrosses (which then become the ‘subfamily-level’ clade Diomedeinae); the true petrel lineage (including diving-petrels) would obviously be termed Procellariidae.

Anyway, whichever phylogenetic hypothesis we follow, it does seem generally agreed that albatrosses are not at all close to true petrels. In fact, the relatively enormous, hyper-long-winged, soaring, often mostly white albatrosses perhaps evolved from a small, dark, storm-petrel-like ancestor given that albatrosses are nested within storm-petrels according to Penhallurick & Wink (2004), and surrounded by them according to Kennedy & Page (2002) and Hackett et al. (2008).

Are there any fossils that might shed light on the ancestral conditions of albatrosses and other crown-tubenoses? Most fossil tubenoses aren’t all that different from living ones. There are, however, a few peculiar specimens, among them the comparatively tiny albatross Murunkus subitus from the Eocene of Uzbekistan, known only from its carpometacarpus. This is seemingly from a bird about a third smaller than the smallest living albatross; so, with a wingspan of perhaps 60 cm or so. However, it’s an ‘alleged’ albatross and its identification as a member of the group awaits confirmation (Mayr & Smith 2012). Other fossil albatrosses (the oldest definite record, Tydea septentrionalis, is from the Oligocene of Belgium) are much like living ones in form, size, and probably in ecology and behaviour.

Foot skeletons of the Oligocene diomedeoidid tubenose Diomedeoides brodkorbi, from Mayr (2009b). The relatively broad, but long, toe bones and their 'closely packed' appearance is obvious, as is the blunt form of the claws. The tiny hallux is a key tubenose character.

Members of an entirely extinct tubenose group are known from the Oligocene of Europe and Iran. These are the diomedeoidids. Yeah, I know, that name is horrible – too many vowels, and too similar to Diomedeidae. Anyway, based on the small size of the dorsal supracondylar process on the distal end of the humerus and other characters, it’s been inferred that diomedeoidids are outside the crown-tubenose clade (Mayr 2009a, De Pietri et al. 2010). The suggestion that this group might be stem-tubenoses explains why I’ve sometimes made a distinction in this article between Procellariiformes as generally understood, and crown-Procellariiformes. Mayr (2009a, b) suggested, on the basis of that small supracondylar process and perhaps other features, that diomedeoidids may have been flap-gliders like oceanitine storm-petrels, and not gliders or soarers. Their very long legs recall those of oceanitines, and their peculiar wide, flattened toes (with blunt, nail-like claws) do too.

Wilson's storm-petrel (Oceanites oceanicus), practicing surface-pattering. Image by Patrick Coin, licensed under the Creative Commons Attribution-Share Alike 2.5 Generic license.

I can’t pretend that we really know all that much about diomedeoidid ecology, functional morphology or behaviour. But there are indications that these possible stem-tubenoses were storm-petrel-like ‘surface patterers’, repeatedly braking to pick up prey from the sea surface, and not soaring swiftly and efficiently on stiff wings like large petrels or albatrosses. [Adjacent image of 'surface patterning' storm-petrel by Patrick Coin].  And if this is correct, and if albatrosses evolved their stiff-winged, high-aspect-ratio soaring wings from storm-petrel-like ancestors (as indicated by some of the phylogenetic results discussed above), then any ecological, behavioural and morphological similarities shared by albatrosses and true petrels must represent convergences. When you look at the general similarity between, say, giant petrels (Macronectes) and albatrosses, that seems pretty surprising.

Much as I’d like to discuss other aspects of tubenose evolution and historical biology, I need to finish this article by briefly discussing the phylogenetic structure of true petrels (Procellariidae), since that’s where we going next.

A phylogeny for petrels

Fulmarus glacialis, a member of Fulmarini. Image by T. Müller, licensed under Creative Commons Attribution-Share Alike 3.0 Unported license.

Recent molecular phylogenetic studies indicate that true petrels (Procellariidae) consist of four major clades: Pterodromini (gadfly-petrels), Procellarinii, Fulmarini (fulmars, giant petrels and kin) and Puffinini (shearwaters). The majority of analyses have found Procellarinii and Puffinini to be sister-taxa, with Fulmarini and Pterodromini representing successively more distant clades to this pairing (Bretagnolle et al. 1998, Nunn & Stanley 1998, Kennedy & Page 2002, Penhallurick & Wink 2004).

The situation is somewhat complicated by the fact, discussed above, that diving-petrels – traditionally classified within their own ‘family’ (Pelecanoididae) – form the sister-taxon to pterodromines in some studies. If this phylogeny is followed, it might be appropriate to split Procellariidae into Procellariinae (for Procellarinii, Fulmarini and Puffinini) and Pelecanoidinae (for Pterodromini and the diving-petrels). Furthermore, prions – conventionally regarded as part of Procellarinii – were found to be outside of the diving-petrel + true petrel clade in some of the topologies recovered by Penhallurick & Wink (2004).

The positions of some species are fairly labile in analyses. Consequently, while the existence of those four major clades (Pterodromini, Procellarinii, Fulmarini and Puffinini) is generally agreed upon (Bretagnolle et al. 1998, Nunn & Stanley 1998, Kennedy & Page 2002, Penhallurick & Wink 2004), their membership varies between analyses. When we visit petrels again, we’ll be looking at pterodromines.

For previous Tet Zoo articles on petrels and other tubenosed seabirds, see…

• A symbiotic relationship between sunfish and… albatrosses? Say what?
• Because the world belongs to petrels (petrels part I)
• Living the pelagic life: of oil, enemies, giant eggs and telomeres (petrels part II)
• Petrels: some form-function ‘rules’, and pattern and pigmentation (petrels part III)
• Noel W. Cusa’s brilliant seabird drawings

And for articles about other kinds of seabirds, see…

• To the Sahara in quest of dinosaurs (living and extinct) (includes discussion of gulls and terns)
• Gannets, most awesome of seabirds
• Happy 2009, from the gulls
• Fascinated by boobies
• Mysterious channels of Alca torda
• Kleptoparasitism at Westbury Manor
• Gary Kaiser’s The Inner Bird: Anatomy and Evolution
• When bivalves attack (or: bivalves vs birds, the battle continues)

Refs – -

Bretagnolle, V., Attié, C., Pasquet, E. 1998. Cytochrome-B evidence for validity and phylogenetic relationships of Pseudobulweria and Bulweria (Procellariidae). Auk 115, 188-195.

De Pietri, V. L., Berger, J.−P., Pirkenseer, C., Scherler, L. & Mayr, G. 2010. New skeleton from the early Oligocene ofGermany indicates a stem−group position of diomedeoidid birds. Acta Palaeontologica Polonica 55, 23–34.

Ericson, P. G. P., Anderson, C. L., Britton, T., Elzanowski, A., Johansson, U. S., Källersjö, M., Ohlson, J. I., Parsons, T. J., Zuccon, D. & Mayr, G. 2006. Diversification of Neoaves: integration of molecular sequence data and fossils. Biology Letters 2, 543-547

Hackett, S., Kimball, R., Reddy, S., Bowie, R., Braun, E., Braun, M., Chojnowski, J., Cox, W., Han, K., Harshman, J., Huddleston, C., Marks, B., Miglia, K., Moore, W., Sheldon, F., Steadman, D., Witt, C., & Yuri, T. (2008). A Phylogenomic Study of Birds Reveals Their Evolutionary History Science, 320 (5884), 1763-1768 DOI: 10.1126/science.1157704

Kennedy, M. & Page R. D. M. 2002. Seabird supertrees: combining partial estimates of procellariform phylogeny. Auk 119, 88-108.

Livezey, B. C. & Zusi, R. L. 2007. Higher-order phylogeny of modern birds (Theropoda, Aves: Neornithes) based on comparative anatomy. II. Analysis and discussion. Zoological Journal of the Linnean Society 149, 1-95.

Mayr, G. 2008. Avian higher- level phylogeny: well-supported clades and what we can learn from a phylogenetic analysis of 2954 morphological characters. Journal of Zoological Systematics and Evolutionary Research 46, 63-72.

- . 2009a. Paleogene Fossil Birds. Springer, Berlin.

- . 2009b. Notes on the osteology and phylogenetic affinitiesof the Oligocene Diomedeoididae (Aves, Procellariiformes). Fossil Record 12, 133–140.

- ., & Smith, T. 2012. A fossil albatross from the Early Oligocene of the North Sea Basin. The Auk 129, 87-95.

Nunn, G. B. & Stanley, S. E. 1998. Body size effects and rates of cytochrome b evolution in tube-nosed seabirds. Molecular Biology and Evolution 15, 1360-1371.

Penhallurick, J. & Wink, M. 2004. Analysis of the taxonomy and nomenclature of the Procellariiformes based on complete nucleotide sequences of the mitochondrial cytochrome b gene. Emu 104, 125-147.

Sibley, C. G. & Ahlquist, J. A. 1990. Phylogeny and Classification of Birds. New Haven: Yale University Press.

Photo by John Sykes, provided by Jon McGowan, used with permission.

Here’s something you don’t see everyday: a female Northern (or Eurasian) sparrowhawk Accipiter nisus and male Common kestrel Falco tinnunculus, photographed together after (it seems) the hawk grabbed the kestrel as a potential prey item. The photo was taken by John Sykes in the Wick Fields area near Christchurch, Dorset, UK, and has been featured by Jon McGowan at The Natural Stuff. Whatever was going on, both birds flew off unharmed. Female sparrowhawks (larger than males) frequently prey on relatively large birds, including pigeons and magpies, and they’re not afraid to take on birds similar in size to themselves. When subduing such large prey, they grip and squeeze it with the talons, but they also start to dismember it (yes, while it’s still alive) with the bill (Fowler et al. 2009). In this case, it’s possible that the sparrowhawk made a mistake, and grabbed the kestrel thinking that it was a more easily dispatched potential prey item.

Photo © 2011 Rick Remington. Be sure to check out the whole sequence.

Or perhaps this was a bold, or experienced, sparrowhawk who thought, or even knew, that she could take on a kestrel and win. After all, remember that intraguild predation – predators killing predators – is common and ubiquitous across the natural world. There are many, many cases on record of raptors killing and eating members of other raptor species. Sparrowhawks are already documented kestrel predators, as are peregrines, goshawks, buzzards and hen harriers. Peregrines F. peregrinus have been observed killing short-toed eagles, sparrowhawks and other raptor species (Hammond & Pearson 1993). This article is not about owls, and I’m going off at a slight tangent here, but I just have to mention the incredible series of photos, taken by Rick Remington in Chicago, where a peregrine repeatedly swooped at a grounded, and clearly terrified, Snowy owl Bubo scandiacus. Some of the poses adopted by the owl were so remarkable that they’ve fooled people into thinking that they were looking at a person in an owl costume. You can see all the photos for yourself here.

Oh, and – one more thing. Unless you’re a bird phylogeny nerd, you’re probably imagining that the two species shown above are fairly close relatives. Several recent molecular studies indicate that falcons are not, in fact, close to other raptors (hawks, eagles, Old World vultures, New World vultures and secretarybirds). Rather, they belong in a clade with parrots and passerines (Ericson et al. 2006, Hackett et al. 2008, Suh et al. 2011), now called Eufalconimorphae. The morphological similarity present between falcons and the members of the hawk-eagle-vulture-secretarybird clade (now best termed Accipitriformes) is thus (presumably!) a striking case of convergence*, but it helps explain some of the otherwise weird features exhibited by falcons and not seen in those other raptors. Within Eufalconimorphae, parrots and passerines form Psittacopasserae.

* The possibility does exist that a ‘raptor-like ecomorph’ was primitive for the clade that includes Eufalconimorphae and kin, and Accipitriformes and kin.

Neornithine bird phylogeny recovered by Suh et al. (2011). (Falcons + (parrots + passerines)) = Eufalconimorphae.

Seriemas seem to be close relatives of Eufalconimorphae (Ericson et al. 2006, Hackett et al. 2008, Suh et al. 2011). You might like to think what this means for phorusrhacids.

Oh, and can everybody please stop using the word ‘raptor’ as a popular term for deinonychosaur, or dromaeosaurid? Admittedly, this rarely causes confusion, but it looks dumb and naive given that THE WORD RAPTOR IS ALREADY IN USE FOR ANOTHER GROUP OF ANIMALS. It would be like deciding to call sauropods ‘elephants’ or something.

Anyway, we’re here because I wanted to show you the photo that you’ve just seen at the very top of the article; it’s pretty exceptional and I hope you’re pleased to see it. For previous Tet Zoo articles on raptors, see…

• Yet again, poor little deer gets killed by big nasty eagle
• Life-size two-dimensional condors and teratorns
• Titan-hawks and other super-raptors
• The most amazing eagle footage ever….. faked
• Raptor makes killing in university grounds
• When eagles go bad, all over again
• Screwed-up Secretary bird
• Condors and vultures: their postures, their ‘bald heads’ and their sheer ecological importance
• Using an eagle to catch and kill a wolf
• A White-tailed eagle in southern England

And for articles on owls, see…

• Titan-hawks and other super-raptors (discusses giant Gargano barn owls)
• Why do some owls have ear tufts?
• From Morocco, with larks, babblers, gazelles, owls and GIANT DINOSAUR BONES
• Fish owls in reverse
• Chock-full of rodent bones
• Myth of the six-foot super-owl
• Why can’t my readers be dumber? Or: replica owls
• Giant owls vs solenodons
• When books die. And owls. I don’t mean “when owls die”… I just mean: owls. As in: this article is mostly about owls.

Refs – -

Ericson, P., Anderson, C., Britton, T., Elzanowski, A., Johansson, U., Kallersjo, M., Ohlson, J., Parsons, T., Zuccon, D., & Mayr, G. (2006). Diversification of Neoaves: integration of molecular sequence data and fossils Biology Letters, 2 (4), 543-547 DOI: 10.1098/rsbl.2006.0523

Fowler, D. W., Freedman, E. A. & Scannella, J. B. 2009. Predatory functional morphology in raptors: interdigital variation in talon size is related to prey restraint and immobilisation technique. PLoS ONE 4(11): e7999. doi:10.1371/journal.pone.0007999

Hackett, S. J., Kimball, R. T., Reddy, S., Bowie, R. C. K., Braun, E. L., Braun, M. J., Cjojnowski, J. L., Cox, W. A., Han, K.-L., Harshman, J., Huddleston, C. J., Marks, B., Miglia, K. J., Moore, W. S., Sheldon, F. H., Steadman, D. W., Witt, C. C. & Yuri, T. 2008. A phylogenomic study of birds reveals their evolutionary history. Science 320, 1763-1768.

Hammond, N. & Pearson, B. 1993. Birds of Prey. Hamlyn, London.

Suh, A., Paus, M., Kiefmann, M., Churakov, G., Franke, F. A., Brosius, J., Kriegs, J. O. & Schmitz, J. 2011. Mesozoic retroposons reveal parrots as the closest living relatives of passerine birds. Nature Communications Aug 23;2:443. doi: 10.1038/ncomms1448.

A tetrapod montage that you might have seen before, by Mehmet Kosemen.

Tet Zoo ver 3 – the Sci Am incarnation of this august and influential institution – has now been going for about 10 months, and a moderately respectable 78 articles have appeared on the blog so far (excluding this one). The vast majority have been lengthy, referenced, heavily illustrated articles – no brief, picture-of-the-day-style contributions or lame rehashings of press releases, thank you very much, and also comparatively little in the way of short contributions in general.

One of the major negative points of the Sci Am blog format is that navigating to older article is not at all easy. In fact it seems about impossible since there’s no date-arranged archive or useful list of categories in the sidebar or anything like that. I’ve suggested behind the scenes that a site-wide rehaul is needed, but no news yet. In fact, as many of you regular readers will know, there’s a lot not to like about the Sci Am blog platform, not only with regard to the site’s appearance and navigability, but also with respect to the commenting and login system. I don’t want to start whining though.

Terrifying tree-kangaroo rendition, from Augusto Vigna Taglianti's 1979 book The World of Mammals (Sampson Low).

If you haven’t been here from the start, what might you have missed? Well, quite a bit. Topics covered here since July 2011 include the neck posture of giant extinct Mediterranean rabbits, a (still incomplete) series of articles on east European frogs and toads, that notorious episode involving the Telegraph newspaper and the Loch Ness monster, sunbathing postures in birds, my write-up of the ZSL cryptozoology meeting (the filmed talks should be online some time soon, by the way), articles on peccary biting behaviour, entelodonts, roadkill, Neotropical jays, cattle, obscure domestic pig breeds, hummingbirds, the ‘tree-kangaroos come first’ hypothesis, vombatiform marsupials, and a fair bit about Mesozoic dinosaurs, pterosaurs and marine reptiles. The toads series is still chugging along slowly in the background (and, yeah yeah, petrels, temnospondyls etc. etc. too). Below please find a list of all articles that have been published here between July 2011 and the end of October 2011 (it takes so long to embed all the links that I ended up giving up on the idea of listing ALL the articles that have appeared here so far). If there’s something you ever wanted to say on those articles but never did, now might be a good idea to get it seen (yet another major negative point on the Sci Am blog platform – we have no ‘recent comments’ section, so any comment added anew to an old article is missed by anyone not reading that article).

Mehmet Kosemen's tongue-in-cheek suggestion that crown-Archosauria should be renamed Awesomes, as encapsulated on a (pre-Nesbitt) simplified cladogram. Share with your friends. Designed to offend those who don't work on crown-archosaurs.

Many of the topics covered below will be revisited when time allows. There’s a lot more to do on marsupials, for example… umm, oh yeah, and on just about everything else. A while back I happened to ask people on facebook what they wanted to see more of on Tet Zoo. Note to start with that this doesn’t matter in any case – as I always say, I blog for me, so you get what I’m interested in. Anyway, the suggestions were all over the place. Predictably, some people said “less dinosaurs, less pterosaurs (since they get written about a great deal already), let’s have more hoofed mammals, more rodents”. But a larger number of people said “more dinosaurs, more pterosaurs”. There were also requests for obscure Palaeozoic synapsids and such, and I can totally understand that. Anyway, I hope that the list below is useful for the purposes of navigation to some of the older stuff, if for nothing else. So, job done. Yikes, must get back to work.

• Welcome to Tetrapod Zoology ver 3
• You have your giant fossil rabbit neck all wrong
• In pursuit of Romanian frogs (part I: Bombina)
• Cryptozoology at the Zoological Society of London. Cryptozoology: time to come in from the cold? Or, Cryptozoology: avoid at all costs?
• In pursuit of Romanian frogs (part II: WESTERN PALAEARCTIC WATER FROGS!!)
• Dryosaurids 101
• Dear Telegraph: no, I did not say that about the Loch Ness monster
• Sunbathing birds
• In pursuit of Romanian frogs (part III: brown frogs)
• Obscure Mesozoic birds you’ll only know about if you’re a Mesozoic bird nerd: Jibeinia luanhera
• Big birds in the Cretaceous of Central Asia: say hello to Samrukia
• Prediction confirmed: plesiosaurs were viviparous
• A day at London’s Grant Museum of Zoology
• Why putting your hand in a peccary’s mouth is a really bad idea
• Those giant killer pigs from hell aren’t pigs
• Dead animals at the roadside
• Inside Nature’s Giants… series 3! Camel!!
• The toads series comes to SciAm: because Africa has toads too
• Obscure, extravagant tropical crows
• Frankly, cattle are awesome
• 20-chromosome toads
• Vertebrate palaeontology at Lyme Regis: of ‘All Yesterdays’, the ‘Leathery Winged Revolution’, and Planet Dinosaur
• Dinosaurs at SVPCA – no Mesozoic non-avialan theropods, thank you very much – and what about those marine reptiles?
• Mystery mammal, badly photographed
• A baby sea-serpent no more: reinterpreting Hagelund’s juvenile Cadborosaurus
• The Mulefoot and other syndactyles: not all pigs are cloven-hoofed
• Three remarkable hummingbird discoveries
• Hoatzins are no longer exclusively South American and once crossed an ocean
• The seemingly endless weirdosity of the Milu
• Artiodactyls and steep slopes, and a new banner for Tet Zoo
• The ‘Tree-Kangaroos Come First’ hypothesis
• In support of Scientific Triassicism: Sues and Fraser’s Triassic Life on Land: the Great Transition
• Of koalas and marsupial lions: the vombatiform radiation, part I
• Marsupial tapirs, diprotodontids, wombats and others: the vombatiform radiation, part II

UPDATE: courtesy of Marko ‘Lev’ Bossche, here are the others. So, this article now includes links to ALL Tet Zoo ver 3 articles. Excellent. Thanks loads, Marko (thanks to the other individuals who sent compiled lists as well, your kind help is much appreciated).

• Tet Zoo highlights 2006-2011, from a Tet Zoo superfan
• The noble tradition of military goats
• What they’re saying about The Great Dinosaur Discoveries
• The discovery and early interpretation of Spinosaurus
• The Crowing crested cobra
• Dwarf mountain toads and the ones with the doughnut-headed tadpoles
• Gerhard Maier’s African Dinosaurs Unearthed: the Tendaguru Expeditions
• Paul Brinkman’s The Second Jurassic Dinosaur Rush
• The Wealden Bible: English Wealden Fossils, 2011
• All the whales of the world, ever (part I)
• All the whales of the world, ever (part II)
• The Second International Workshop on the Biology of Sauropod Dinosaurs (part I)
• A Merry Tet Zoo Christmas
• In case you forget, softshell turtles are insanely weird
• ‘Rigid Swimmer’ and the Cretaceous Ichthyosaur Revolution (part I)
• Did dinosaurs and pterosaurs practise mutual sexual selection?
• The Second International Workshop on the Biology of Sauropod Dinosaurs (part II)
• Because giraffes are heartless creatures, and other musings
• STOP ‘feeding’ the ducks
• Happy Birthday Tetrapod Zoology: SIX YEARS of blogging
• Happy 6th Birthday, Tetrapod Zoology (part II)
• The more you know about colubrid snakes, the better a person you are
• A symbiotic relationship between sunfish and… albatrosses? Say what?
• “San Diego Demonoid”: you mean that dead opossum?
• Love for Mastigodryas, Tomodon, Sordellina and all their buddies: you know it’s right
• Williams and Lang’s Australian Big Cats: do pumas, giant feral cats and mystery marsupials stalk the Australian outback?
• Identify the Baja California mystery whale carcass!
• Greg Paul’s Dinosaurs: A Field Guide
• Grampus griseus joins the globicephalines
• A peculiar whale skeleton is included fortuitously in the sci-fi movie Hunter Prey
• The gigantic, shark-toothed, small-flippered, long-bodied, sea-going predatory lizard that is Hainosaurus
• The carcass of a large, modern-day marine reptile!
• Because the world belongs to petrels (petrels part I)
• Living the pelagic life: of oil, enemies, giant eggs and telomeres (petrels part II)
• Petrels: some form-function ‘rules’, and pattern and pigmentation (petrels part III)
• Noel W. Cusa’s brilliant seabird drawings
• Alien viruses from outer space and the great Archaeopteryx forgery
• Amphisbaenians and the origins of mammals
• There are giant feathered tyrannosaurs now… right?
• Ryan et al.’s New Perspectives on Horned Dinosaurs: a review
• Chickens, 2012
• The Cadborosaurus Wars
• Eld’s deer: endangered, persisting in fragmented populations, and morphologically weird… but it wasn’t always so
• Tet Zoo ver 3, (part of) the story so far
• Raptor vs raptor

Eld's deer stag, by Raul654, used under Creative Commons Attribution-Share Alike 3.0 Unported license.

This is Eld’s deer Cervus eldi* or the Brow-antlered deer, Thamin or Tamin, a moderately obscure, CITES-listed Old World deer discovered (by Lt. Percy Eld) in India in 1839. It was later found to occur in fragmented populations across much of south-east Asia and also in southern China. Fossils are known from Java and it seems that its original distributed was enormous, being mostly continuous across eastern India and Burma, Thailand, Laos, Cambodia and southern China.

* Many sources spell the specific name eldii but it seems that ‘eldi’ is correct. I’m putting the species in Cervus following Pitra et al. (2004).

Eld’s deer is fairly large (shoulder height is about 1.2 m, weight is 95-150 kg) with lyre-shaped antlers, the beams of which grow outwards before turning inwards. The brow tine is especially long, as hinted at by one of the common names, and appears to form a continuous curve with the beam “so that in profile the antlers appear to be bow-shaped” (Nowak 1999, p. 1108).

Eld’s deer has often been characterised as a tropical wetland specialist. However, this is not true when we look at all three recognised subspecies. In fact these different subspecies are pretty distinctive and again highlight the fact that ‘subspecies’ are not just entities invented for the purposes of bookkeeping but, rather, distinct lineages that we often need to pay attention to.

Cervus eldi eldi stag, antlers in velvet, quite possibly not photographed in the wild. Photo by Mongyamba, licensed under Creative Commons Attribution-Share Alike 3.0 Unported license.

The Manipur subspecies (C. eldi eldi) [shown here] is a true wetland deer, possessing especially large, spreading hooves and peculiar cornified skin on the rest of the digits. It lives on thick, dense mats of floating vegetation known locally as ‘phum’ or ‘phumdi’ (Geist 1999) and is critically rare, being restricted to a tiny area about 15 km square. As of 2004, there were only about 180 individuals in the wild. About another 180 animals are kept in zoos. It was actually thought extinct prior to rediscovery in 1951.

Burmese brow-antlered deer (C. e. thamin), photographed at Zurich Zoo by Albinfo, licensed under Creative Commons Attribution-Share Alike 2.5 Generic license.

Meanwhile, the Burmese or Burma brow-antlered deer C. e. thamin and Thai brow-antlered deer C. e. siamensis are rather different, being animals of dry deciduous forests. This might negate suggestions that the Eld’s deer lineage as a whole is specialised for wetlands. However, there are suggestions that the modern habitat of the Burmese and Thai forms is not natural but the result of human persecution and the removal of their ancestral habitat due to agriculture (Lekagul & McNeely 1977). High nucleotide variation in these populations – similar to that of really widespread deer like roe and sika – suggests that both forms were previously very abundant and widespread (Balakrishnan et al. 2003), an observation that agrees with the idea that their ranges have been substantially decreased in historic times.

The Eld’s deer population on Hainan is sometimes recognised as the subspecies C. e. hainanus but the validity of this form was not supported by Balakrishnan et al. (2003). These authors found the Hainan population to be close to indisputable members of the Thai form and hence recommended included it within that subspecies.

That study (based on mtDNA analysis) also found the nominate subspecies to be closer to the Burmese form than to the Thai one. That’s surprising, since the latter two look extremely similar while the Manipur form is the really distinctive one. Furthermore, all three subspecies were distinct with little evidence for intermingling, and all three exhibit their own phylogeographic structure (Balakrishnan et al. 2003). While it therefore makes sense that all three be recognised as distinct genetic entities for the purposes of conservation and viability, the problem is that some of the populations (including the Manipur one and the Hainan C. e. siamensis one) now exhibit very low amounts of genetic variation and would probably benefit from increased genetic variation. It might therefore be appropriate to start moving animals around in order to increase the genetic health of the populations (Balakrishnan et al. 2003).

Eld's deer diorama at the AMNH. Photo by ideonexus, licensed under Creative Commons Attribution-Share Alike 2.0 Generic license.

A cervine – but what sort of cervine?

Eld’s deer looks odd, and for this reason Pocock (1943) decided that it should warrant it own genus. He therefore came up with Panolia for this species [UPDATE: see comments!! I was not aware of the new taxonomy used by Groves & Grubb (2011) while writing this article]. In overall appearance, though, Eld’s deer is not all that different from the Barasingha Rucervus duvauceli and the now extinct Schomburgk’s deer R. schomburgki (though antler form is different between the three, of course). Ellerman & Morrison-Scott (1951) and Geist (1999) therefore grouped all three together in Rucervus; Geist noted that these species shared relatively high antler mass, short tails, no rump patch, large, subhypsodont teeth specialised for grazing, and legs and feet specialised for soft ground.

Geist's (1999) depiction of the three supposed close relatives placed in Rucervus. L to r: Barasingha, Eld's deer, Schomburgk's deer. Compare the Eld's deer to the stag in the AMNH diorama shown above.

Cervid phylogeny from Pitra et al. (2004). You should be able to see the Eld's deer + Milu clade near the middle of the diagram. Click to enlarge.

However, these general morphological similarities might represent convergence rather than genuine affinity. Pitra et al. (2004) used data from mitochondrial DNA to look anew at Old World deer phylogeny and were most surprised to find Eld’s deer to group closely with the Milu or Père David’s deer C. davidianus, with both forming a clade that is the sister-group to a larger cervine clade that includes sika, red deer and sambar. Similar results were reported by Gilbert et al. (2006). This discovery may or may not support the inclusion of both Eld’s deer and Père David’s deer within Cervus (as usual, that’s down to personal preference), but it does weaken the hypothesis that Eld’s deer is anything to do with Rucervus (which, following Pitra et al. (2004), I use here for the Barasingha and Schomburgk’s deer alone. Rucervus seemingly forms a clade with Axis and is outside of Cervus). The grouping of Eld’s deer with Père David’s deer also led to the curious idea that the latter might have arisen as a hybrid of Eld’s deer and a wapiti-type cervine – a subject that I covered in an article from last year devoted to Père David’s deer.

Sambar (Cervus unicolor), one of my favourite deer. Remind me to elaborate on that comment some time. Image by Wikigringo, licensed under the Creative Commons Attribution-Share Alike 2.5 Generic license.

Of course, this phylogenetic hypothesis isn’t the only one out there. Ouithavon et al. (2009) found Eld’s deer to be especially close to the Sambar C. unicolor (though note that their sample was limited, since they were only analysing taxa present in Thailand). Most recently, Zhang & Zhang (2012) found Eld’s deer to be the sister-taxon to a clade that included Père David’s deer as well as sambar, sika, red deer and so on.

There’s an important message to take from this quick look at what is now an obscure, little-known and endangered species, restricted to small populations and mostly famous for being weird. It’s that such obscure, endangered peculiarities might be the sorry relicts of what were once far more abundant, far more widespread populations. The data suggests that, just a few thousand or even few hundred years ago, Eld’s deer might have been a familiar, abundant and highly successful animal across an enormous area. Human impact on its diversity and distribution has been substantial.

Why did I write this article? Because obscure artiodactyls need more love, and I picked this species at random as one you don’t often hear about. Aiming to finish the petrel series soon, just haven’t yet found the time.

For previous Tet Zoo articles on deer, see…

• Deer oh deer, this joke gets worse every time I use it
• Britain: wildlife theme-park
• At the 56th SVPCA – hello Dublin! (Megaloceros)
• The plasticity of deer
• Pouches, pockets and sacs in the heads, necks and chests of mammals, part IV: reindeer and a whole slew of others
• The seemingly endless weirdosity of the Milu

Refs – -

Balakrishnan, C. N., Monfort, S. L., Gaur, A., Singh, L. & Sorenson, M. D. 2003. Phylogeography and conservation genetics of Eld’s deer (Cervus eldi). Molecular Ecology 12, 1-10.

Ellerman, J. R. & Morrison-Scott, T. C. S. 1951. Checklist of Palaearctic and Indian Mammals, 1758 to 1947. British Museum (Natural History) Trustees, London.

Geist, V. 1999. Deer of the World. Swan Hill Press, Shrewsbury.

Gilbert, C., Ropiquet, A. & Hassanin, A. 2006. Mitochondrial and nuclear phylogenies of Cervidae (Mammalia, Ruminantia): systematics, morphology, and biogeography. Molecular Phylogenetics and Evolution 40, 101-117.

Lekagul, B. & McNeely, J. A. 1977. Mammals of Thailand. Kurushpa Ladprao Press, Bankok, Thailand.

Nowak, R. M. 1999. Walker’s Mammals of the World, Sixth Edition. The Johns Hopkins University Press, Baltimore and London.

Ouithavon, K., Bhumpakphan, N, Denduangboripant, J., Siriaroonrat, B. & Trakulnaleamsai, S. 2009. An analysis of the phylogenetic relationship of Thai cervids inferred from nucleotide sequences of protein kinase C iota (PRKCI) intron. Kasetsart J. (Nat. Sci.) 43, 709-719.

Pitra, C., Fickel, J., Meijaard, E., & Groves, C. (2004). Evolution and phylogeny of old world deer Molecular Phylogenetics and Evolution, 33 (3), 880-895 DOI: 10.1016/j.ympev.2004.07.013

Pocock, R. I. 1943. The larger deer of British India. Part II. Journal of the Bombay Natural History Society 43, 553-572.

Zhang, W.-Q. & Zhang, M.-H. 2012. Phylogeny and evolution of Cervidae based on complete mitochondrial genomes. Genetics and Molecular Research 11, 628-635.

Two renditions of 'Cadborosaurus' (upper one by C. M. Kosemen; one at lower right by Naish) with a pipefish. Not to scale!

Over the last few months, I and two of my colleagues have been involved in an interesting dialogue in the literature. It concerns the entity dubbed ‘Cadborosaurus’ – a marine, horse-headed ‘mega-serpent’, supposedly reported by witnesses from the waters off British Columbia and elsewhere in the North Pacific. People who read my stuff (both here and in print) will know that I have more than a passing interest in cryptozoology, and especially in ‘sea monsters’; indeed, I’ve written about ‘Cadborosaurus’ quite a few times. As I always say, this interest in cryptozoology might be a dumb thing to admit, given the negative stigma attached to the field. And I’m sure that it’s based in part on adherence to the naïve and childish hope that sea monsters, relicts hominoids and such might actually be real.

Nevertheless, I remain interested in cryptozoology both because I think that some eyewitness accounts are really intriguing and difficult to explain, and because I’m interested in how people perform as observers of wildlife (see Paxton 2009). Unlike many who class themselves as sceptics, I’ve tried to understand where cryptozoologists are coming from, I’ve read and (do still read) the cryptozoological literature, and I don’t think that we should necessarily reject cryptozoological hypotheses as untenable without looking at the data (such as it is) first.

Cadborosaurus wars: Round 1

As I aim to show here, dealing with cryptozoologists can be a frustrating, even infuriating, business. As most people interested in mystery animal research will know, Michael Woodley, Cameron McCormick and myself recently argued that an alleged ‘baby Cadborosaurus’ was very likely no baby sea-serpent at all, but rather a mangled and half-remembered description of a pipefish (Woodley et al. 2011). We tabulated the various observations reported by the witness (William Hagelund), compared them to lists of characters compiled by examining numerous candidate species, and showed as clearly as possible that the pipefish identification is the one that best matches Hagelund’s observations. In other words, we did our best to examine the identity of the alleged creature in an empirical, critical fashion (Woodley et al. 2011). As I explained last time round, we have to remember that Hagelund wrote up his description of the encounter about two decades after it actually occurred, and that he did not ascribe the various ‘cadborosaur’ traits to his animal that the primary supporters of ‘Cadborosaurus’ (Edward Bousfield and Paul LeBlond) said that he did.

William Hagelund's 'baby Cadborosaurus' compared with a pipefish. Illustration by Cameron McCormick. See Woodley et al. (2011) for explanation.

Round 2

Most researchers were/are generally happy with our contention. But, perhaps unsurprisingly, I learnt from personal communication that Bousfield and LeBlond were not so happy. Essentially, they regarded our pipefish hypothesis as a non-starter hardly worthy of consideration. In a terse published response, provocatively titled ‘Pipefish or pipe dream?’, they accused us of “indulg[ing] in the common home-quarterbacking habit of insisting that anything described as different must be an erroneous description of something found in a book that vaguely looks like it” (Bousfield & LeBlond 2011, p. 779). I’ve no idea at all what a “common home-quarterbacking habit” is, and frankly the rest of the article isn’t much better. Bousfield & LeBlond (2011) cite Woodley et al. as “Naish and colleagues”, as if they weren’t paying attention, say that Hagelund’s ‘baby’ is grossly different from a pipefish and better matches ‘Cadborosaurus’, and even claim that employment of Occam’s Razor strengthens their hypothesised identification and bolsters rejection of ours (Bousfield & LeBlond 2011).

Bousfield & LeBlond's (2011) effort to show that pipefishes and Hagelund's animal are totally, totally different.

Round 3

Somewhat unhappy with the tone of Bousfield & LeBlond’s (2011) article, Michael, Cameron and I elected to produce a response (Woodley et al. 2012). Bousfield & LeBlond (2011) made three main points that require riposte. Firstly, they honestly thought that “by attempting to dismiss our pipefish identification, they had completed their task of critiquing our paper” (Woodley et al. 2012, p. 143). We find this dismissal premature and naïve, since – even if the pipefish hypothesis is deemed unsatisfactory – we (Woodley et al. 2011) drew attention to several other fish taxa that are more similar to Hagelund’s animal than is Bousfield and LeBlond’s reconstruction of ‘Cadborosaurus’.

Greater pipefish (Syngnathus acus); image from wikipedia.

Secondly, Bousfield & LeBlond (2011) argued that the Hagelund animal couldn’t be a pipefish due to the presence of an obvious neck, a horizontally aligned tail, large jaws and so on. However, they were apparently unable to appreciate or understand that the un-pipefish-like features they refer to were not reported or described by Hagelund at all. Furthermore, as I said above, Hagelund wrote about his encounter about two decades after it supposedly happened, and it again seems naïve to assume without caveat that the features he described were 100% accurate.

Thirdly, Bousfield & LeBlond’s (2011) claim that Occam’s Razor demands that their ‘baby sea serpent’ interpretation be taken more seriously than our ‘misidentified pipefish’ hypothesis is laughable. You can read our response in full to see how we dealt with this claim (Woodley et al. 2012). You might be able to guess.

Round 4

Bousfield and LeBlond will not, it seems, be responding further in print.

In case it's not obvious, this is a cartoon, and you need to understand irony to appreciate it.

But after receiving a series of emails from an apparently frustrated Ed Bousfield, I feel it’s time to say publicly that the published work on ‘Cadborosaurus’ is terrible science and more to do with the imagination and preconceptions of the protagonists than anything to do with actual biology.

Frankly, I’m tired of being told that I and my colleagues are the ones with the ridiculous ideas, the ones who aren’t thinking things through in proper scientific fashion, or the ones who are failing to recognise the true phylogenetic affinities of organisms reported by eyewitnesses. Bousfield has even – I wish I were joking – referred to us as “ivory tower scientists”, and as people who “may indeed know something about fossil vertebrate animals but know relatively little about living megaserpents in general and Cadborosaurus willsi in particular”.  I wish he would do his homework and work out for himself that only one of us is a palaeontologist. The ‘Ivory Tower’ claim is amusing, since it doesn’t take much research to appreciate that I and my colleagues are pretty much the very antithesis of that term. I offer the adjacent cartoon as a parody.

As if it’s not clear enough already, I want to state here why I think that the ‘Cadborosaurus’ work published by Bousfield and LeBlond is terrible science. Much of this has been covered elsewhere in the literature (Ellis 1994, Staude & Lambert 1995, Bauer & Russell 1996, Naish 2001, Woodley et al. 2008), but I want to repeat it so that the above discussion can be better seen in context, especially for those who are unfamiliar with this subject. And to those who have read what I’ve said about ‘Cadborosaurus’ before (in this 2006 Tet Zoo ver 1 article, for example), note that I have definitely become less sympathetic.

Interpreting Cadborosaurus: eyewitness accounts

The idea that ‘Cadborosaurus’ might exist and be real all stems, of course, from the fact that eyewitnesses have reported encounters with apparently long-bodied, horse-headed ‘mega-serpents’ in the waters of the north-east Pacific. A long-bodied, horse-headed vertebrate carcass, apparently retrieved from the stomach of a sperm whale at a Queen Charlotte Islands whaling station in 1937, sounds like the same sort of animal. Herein lies the case for ‘Cadborosaurus’. To conclude that the animal is real, you have to accept that these eyewitness reports, and the photos of that carcass, all represent descriptions or depictions of the same one species.

Needless to say, the foundations of the case for ‘Cadborosaurus’ are far from secure. Once you look at individual eyewitness accounts in detail, the hypothesis that they all represent the same animal species crumbles as clearly untenable. In fact, one of the many failings of Bousfield, LeBlond and like-minded cryptozoologists is that they’re unable or unwilling to realise that the cryptids they believe in are composite entities, constructed by combining observations of different species and/or phenomena. Some ‘Cadborosaurus’ descriptions do refer to vertically undulating, serpentine phenomena that, if they do represent animals, are difficult to reconcile with species that we know about, so please note here that I am not necessarily discounting altogether the existence of a marine vertebrate that remains unrecognised by scientists at large.

The diversity of entities believed to represent sightings of 'Cadborosaurus', from Woodley et al. (2012). Illustration by Cameron McCormick.

A 'Cadborosaurus' sighting (this one reported by fisherman David Miller in 1959). The drawing here (from Bright 1989) is actually substantially 'augmented' relative to Miller's original. Should we interpret such sightings as evidence for a new species of 'mega-serpent', or should we try to interpret them as confused accounts of seal, deer or other known animals or phenomena?

But, not only are the observations on record highly disparate, they are most parsimoniously interpreted as descriptions of many things. If you own any or all of the ‘Cadborosaurus’ literature, look at the images penned by eyewitnesses. There are various seal-shaped and deer-shaped heads, sometimes with obvious ears and short ‘horns’, and sometimes not (Heuvelmans 1965, Bousfield & LeBlond 1995, LeBlond & Bousfield 1995). Could it be, I wonder, that many (maybe all) of these sightings are actually of seal and swimming deer? The ‘camel-like’ head of ‘Cadborosaurus’, with its large, dark eyes and overhanging upper lip, is likely that of a surfacing male Northern elephant seal Mirounga angustirostris – a surprisingly large beast, the head of which might stand about a metre above the sea surface when ‘standing’ vertically in the water. I came up with this myself after seeing photos of a surfacing elephant seal and later heard it from a Canadian biologist whose father had a close encounter with a surfacing elephant seal (and immediately thought ‘Cadborosaurus’). Bauer & Russell (1996) “regard pinnipeds, especially the northern elephant seal Mirounga angustirostris, as the most likely candidates for the source of most of the [‘Cadborosaurus’] observations reported at sea” (p. 13). The Northern elephant seal is confirmed, incidentally, as a visitor of British Columbian waters, and remember that it stays down for so long, and visits the surface so briefly and intermittently that (some specialists suggest) it might be better regarded as a ‘surfacer’ than as a ‘diver’. Note that the description of the ‘Cadborosaurus’ head is also is a good match for that of a Moose Alces alces, as emphasised by the composite image below (from Cryptomundo). Moose are excellent swimmers and even dive to feed on submerged vegetation (the idea that they might explain some sea and lake monster reports is not exactly novel).

With the seal-headed and deer-headed sightings out of the picture, do the descriptions of serpentine, multi-humped animals remain as good evidence for marine ‘mega-serpents’? Elsewhere in the world, standing waves, colliding wakes, lines of swimming cetaceans, pinnipeds and even seabirds flying close to the water surface have all been misinterpreted by people as representing giant, multi-humped water monsters.

So I’m not confident that there is a large and compelling body of eyewitness data supporting the reality of ‘Cadborosaurus’. Instead we have a tainted and inconsistent pool of discordant accounts that cannot be interpreted as evidence for the existence of a single species.

Interpreting Cadborosaurus: the Naden Habour carcass

It is with the identikit image of the ‘Cadborosaurus’ construct in mind that Bousfield and LeBlond interpreted the Naden Harbour photos. The photos show a seemingly long-bodied, camel-headed vertebrate carcass.

One of several photos of the Naden Harbour 'Cadborosaurus' carcass.

Bousfield and LeBlond’s assumption that what we see in the photos represents the life appearance of a very peculiar creature is naïve. After briefly considering (and rejecting) an informal suggestion that the carcass was a “fetal baleen whale”, they adopted the hypothesis that Naden Harbour carcass = mega-serpent (Bousfield & LeBlond 1995, p. 9) without considering other hypotheses. Could it actually have been the highly decomposed, incomplete remains of a known species, like a large shark, or a bony fish of some sort? Many of us interested in ‘sea monsters’ are thinking along these lines, and in fact the idea that the carcass might be the substantially mangled remains of a shark has already been mentioned in print (Bauer & Russell 1996, Naish 2001). Bousfield & LeBlond (1995) provided a superficial and wholly amateurish description of the carcass, made some enormous and highly error-prone assumptions about its skeletal anatomy, and used one of the photographs of the carcass (yes, I said one of the photographs) as the holotype for a new species: Cadborosaurus willsi Bousfield & LeBlond, 1995.

Does any of this seem like ‘good’ science involving appropriate consideration of alternative hypotheses, caution, conservatism and best practise? I leave you to judge, but I certainly do not regard ‘Cadborosaurus’ as a valid biological entity based on the data that Bousfield & LeBlond (1995) presented. Incidentally, Bousfield and LeBlond published their description of the ‘new species’ in ‘supplement 1’ of volume 1 of Amphipacifica, a new publication stated to be devoted to invertebrate systematics. The editorial board consisted of Bousfield as Managing Editor and C. P. Staude and P. Lambert as Assistant Editors. On the ‘Cadborosaurus’ article, Staude and Lambert were “opposed to its publication as a formal species description” and felt the need to express this opinion in an editorial published at the front of the issue (Staude & Lambert 1995).

The ‘living serpentine plesiosaur’ hypothesis

Bousfield and LeBlond have consistently employed a somewhat confused approach to the identity of ‘Cadborosaurus’. The original description (Bousfield & LeBlond 1995) proposes that ‘Cadborosaurus’ is a long-bodied member of ‘Euryapsida’ (the mostly defunct and now ambiguous term once used for sauropterygians and a number of possibly allied reptile groups); specifically, they classify it as ‘Class Reptilia, Subclass Euryapsida?, Order Plesiosauria?’ (p. 8). They’ve also said that ‘Cadborosaurus’ combines reptilian and mammalian traits, and in LeBlond & Bousfield (1995) they intimated a relationship with thalattosuchian crocodyliforms. On other occasions, they’ve said that they were not/are not supporting a plesiosaurian identify for ‘Cadborosaurus’ (the abstract of Bousfield & LeBlond (1995) uses the wording “within vertebrate class Reptilia … the animal appear [sic] least unlike some plesiosaurs of Mesozoic age” (p. 3)). To be clear, they actually were and are favouring the hypothesis that ‘Cadborosaurus’ is a relict surviving plesiosaur.

'Cadborosaurus' as imagined by Bousfield and LeBlond (illustration by D. Naish).

If you’ve never heard the remarkable claim that a giant, serpentine plesiosaur might inhabit the modern waters of the north Pacific, I can hardly blame you. This work has been largely ignored by qualified biologists, mostly because they think it’s nonsense and unworthy of serious consideration. Aaron Bauer and Anthony Russell (1996), well known for their excellent work on lizards and other vertebrates, wrote a lengthy critical assessment of Bousfield & LeBlond’s (1995) description of ‘Cadborosaurus’, and I note that their conclusions and criticisms were highly similar to those of myself and my colleagues (Naish 2001, Woodley et al. 2008, 2011, 2012).

Time to say it like it is: BAD SCIENCE

Some 'Cadborosaurus' acccounts describe swimming giraffes. Just sayin' (and, no, I am not being in the least bit serious). Spanish Banks cadborosaur drawing from Bright (1989). Floating giraffe image by Don Henderson.

We’ll be coming back to ‘Cadborosaurus’ again in the future. For now, I hope that several things are clear:-

– eyewitness accounts used to support the reality of ‘Cadborosaurus’ likely represent a hodge-podge of seal and deer sightings, as well as sightings of other animals and phenomena. Maybe the north-east Pacific is home to a large, as yet unrecognised, vertebrate animal, but this is not clear from eyewitness accounts. They represent an inconsistent set of diverse descriptions that cannot be interpreted as evidence for a new species.

– the 1937 Naden Harbour carcass, believed by Bousfield and LeBlond to represent the same ‘mega-serpent’ reported as a living animal by eyewitnesses, is ambiguous and its interpretation as a modern-day serpentine plesiosaur, awarded a binomial name on the basis of old photographs, cannot be considered conservative, competent science.

– a re-interpretation of a supposed ‘baby Cadborosaurus’ account as that of a pipefish has been strenuously counter-argued by the main supporters of ‘Cadborosaurus’ on the grounds that its identification as a baby mega-serpent is more likely. Whether the ‘misidentified pipefish’ hypothesis is correct or not, it is just insulting to be told that this hypothesis is less parsimonious than the hypothesis that it represents a baby sea-serpent.

– all in all, the published research on ‘Cadborosaurus’ involves improbable conclusions, a lack of critical analysis, and a lack of the conservatism and restraint that’s normal in scientific research. It’s just bad, bad science. Furthermore, the primary supporters of the alleged reality of ‘Cadborosaurus’ have displayed a frustrating arrogance, lack of humility and stubborn attitude whenever their ideas are – quite justifiably – placed under scrutiny.

Cameron has written a series of articles about our Woodley et al. (2011) paper and additional ‘Caddy’ reports: part 1 is here, then there’s part 2a, part 2b, part 3, part 4 and part 5. For various Tet Zoo articles on ‘sea monster’ mysteries of various kinds, see…

• The amazing Hook Island sea monster photos
• Santa Cruz’s duck-billed elephant monster
• Professor Sharpe’s mysterious sea-serpent photo
• It had wool, and armour plates, a massive beak, horns, and it smelled veeeeery bad: whatever happened to the Tecolutla monster?
• Where are all the dead sea monsters?
• Skull of the Moore’s Beach monster revealed!
• The Long-necked seal, described 1751
• Statistics, seals and sea monsters in the technical literature
• A Russian sea monster carcass is claimed to be that of an ancient ‘archaeocete’ whale
• Phylogenetic roulette and the identification of sea monsters
• Sea Monsters, the CFI conference
• Won’t someone please think of the coelacanths, and other lamentations
• A sea monster poster for the 9th European Symposium of Cryptozoology
• Cryptozoology at the Zoological Society of London. Cryptozoology: time to come in from the cold? Or, Cryptozoology: avoid at all costs?
• A baby sea-serpent no more: reinterpreting Hagelund’s juvenile Cadborosaurus

Refs – -

Bauer, A. M. & Russell, A. P. 1996. A living plesiosaur?: A critical assessment of the description of Cadborosaurus willsi. Cryptozoology 12, 1-18.

Bousfield, E. L. & LeBlond, P. H. 1995. An account of Cadborosaurus willsi, new genus, new species, a large aquatic reptile from the Pacific coast of North America. Amphipacifica 1 (supplement 1), 1-25.

- . & LeBlond, P. H. 2011. Pipefish or Pipe Dream? Journal of Scientific Exploration 25, 779-780.

Bright, M. 1989. There are Giants in the Sea. Robson Books Ltd, London.

Ellis, R. 1994. Monsters of the Sea. Alfred A. Knopf, New York.

Heuvelmans, B. 1968. In the Wake of the Sea-Serpents. Hart-Davis, London.

- . & Bousfield, E. L. 1995. Cadborosaurus: Survivor From the Deep. Horsdal & Schuber Publishers Ltd., Victoria, Canada.

Naish, D. 2001. Sea serpents, seals, and coelacanths: an attempt at a holistic approach to the identity of large aquatic cryptids. Fortean Studies 7, 75-94.

Paxton, C. G. M. 2009. The plural of “anecdote” can be “data”: statistical analysis of viewing distances in reports of unidentified giant marine animals 1758-2000. Journal of Zoology 279, 381-387.

Woodley, M. A., McCormick, C. A., & Naish, D. 2012. Response to Bousfield & LeBlond: Shooting pipefish in a barrel; or sauropterygian “mega-serpents” and Occam’s razor. Journal of Scientific Exploration 26, 151-154.

- ., Naish, D. & McCormick, C. A. 2011. A baby sea-serpent no more: Reinterpreting Hagelund’s juvenile “Cadborosaur” report. Journal of Scientific Exploration 25, 497-514.

Woodley, M., Naish, D., & Shanahan, H. (2008). How many extant pinniped species remain to be described? Historical Biology, 20 (4), 225-235 DOI: 10.1080/08912960902830210

You've heard of combs and wattles. That flap posterior to the cheek region is called an ear lobe.

Transylvanian naked-necked chicken, photo by Lily15.

I don’t want to say too much about chickens as I’ll be here all day (drat, semi-failed). Must say a few brief things though. The history of chicken domestication is complex (though the Red junglefowl Gallus gallus is the primary progenitor), as is the story of how people took chickens (from a centre of origin in or near south-east Asia, perhaps in Thailand) west to Asia Minor, Europe and Africa, as well as east across the Pacific. Some Asian chicken breeds (often used primarily as fighters) were and are gigantic and strikingly long-legged, and some people have suggested that a now extinct, mysterious giant species or form of chicken – dubbed Gallus giganteus – contributed to the domestic chicken gene pool. By crossing several large Asian breeds, the American Jersey giant was created – a breed where cocks can weigh about 6 kg. Debate continues over whether Polynesians took chickens to South America prior to the time of Columbus. Polynesian chickens had key roles in entertainment (fighting) and ornamentation (on Hawaii, their feathers were used to indicate the sovereignty of kings on decorative poles termed kahilis) as well as in providing edible protein.

People have bred featherless – yes, naked – chickens, and there’s also a highly distinctive Romanian breed (sometimes unofficially dubbed a turken or churkey on account of a vague similarity with turkeys) with a naked neck. Japanese Yokohama cocks can have tail feathers measuring anything up to 10.6 m (this is the record-holder, reported in 1972 from Shikoku). Silkies are famously weird in having hair-like plumage (rendering them flightless) as well as five toes on each foot and blueish-blackish skin, flesh and bones. Black meat and bones are also present in the Indian Kadaknath and Indonesian Ayam Cemani. Birds in general typically possess white or black skin, yet many domestic chicken breeds have yellow skin. There’s a ton more that could be said, but consider this a teaser.

And, finally, for those who have never seen it before…

The accompanying paper (available as a free pdf) is available here. For more on gamebirds at Tet Zoo, see…

• How to prevent cannibalism in pheasants
• The snood of the turkey, the wires and rackets of the motmot, the face of the rook

It says something of the invigoration of dinosaur research that an enormous technical tome, 624 pages long and containing 36 chapters, could be produced on horned dinosaurs (or ceratopsians) alone. New Perspectives on Horned Dinosaurs, published in 2010 and edited by Michael Ryan, Brenda Chinnery-Allgeier and David Eberth, stems from the Royal Tyrrell Museum Ceratopsian Symposium, held at Drumheller, Alberta, in September 2007. The organisers of this meeting were inspired by the major upsurge in horned dinosaur research currently underway – it involves the description of new taxa, a renewed interest in anatomy, behaviour and functional morphology, and the application of new techniques to such areas as the mechanical behaviour of bone, niche partitioning, phylogeny, biogeography and ontogeny.

Despite the popular name, not all horned dinosaurs are horned. Ceratopsia in fact includes psittacosaurids (a group of Asian bipedal forms, the skulls of which are often superficially likened to those parrots), the deep-headed, hornless leptoceratopsids, an assemblage of small- to medium-sized, mostly quadrupedal forms that were once grouped together as protoceratopsids (they have bony frills, and some, like the eponymous Protoceratops, have short or incipient nasal horns), and the members of the large-bodied, horned Zuniceratops + ceratopsid clade*. Ceratopsidae includes both the mostly long-frilled chasmosaurines with their long supraorbital, postorbital or brow horns, and the mostly short-frilled centrosaurines with their long nasal horns and typically short (or absent) postorbital horns. Zuniceratops and several other fossils seem to show that postorbital horns are primitive for the Zuniceratops + Ceratopsidae clade. Protoceratops, ceratopsids and their frill-headed kin (that is, all ceratopsians closer to Triceratops than to Psittacosaurus) are united within Neoceratopsia, while the node-based clade that includes Protoceratops and Triceratops is termed Coronosauria (Sereno 1998). [Highly simplified cladogram below borrowed from Thomas R. Holtz's University of Maryland lecture notes. Version below is low-res, pdf here has far better resolution.]

Simplified cladogram of Ceratopsia (and other members of the ornithischian clade Marginocephalia), compiled by Thomas R. Holtz using skeletal reconstructions by Greg Paul and others. Follow link above for higher-res version.

* There’s no name for the node-based clade that includes Zuniceratops, Ceratopsidae, and all descendants of their common ancestor. However, the names Ceratopsoidea and Ceratopsomorpha both apply to the branch-based clade that includes all taxa closer to Triceratops than to Protoceratops (Sereno 1998, Wolfe & Kirkland 1998). Ceratopsoidea has been used more frequently in the literature than Ceratopsomorpha and this is the best name we have at the moment for the “Zuniceratops + Ceratopsidae clade”.

New Perspectives on Horned Dinosaurs consists of five sections: overview; systematics and new taxa; anatomy, functional biology and behaviour; phylogeny, biogeography, taphonomy and palaeoenvironment; and the history of collecting. The bulk of the book is made up of the second and third sections. The first section includes but a single article, and the last section includes just two. Virtually all contributions are very well illustrated and there is even a colour plate section. The book is large-format (c. 29 x 22 cm) and hence very different from most of the other multi-authored dinosaur-themed volumes published by Indiana University Press.

Homage to Prof. Dodson: “40 years of Ceratophilia”

Prof. Peter Dodson, professor of anatomy and paleontology at University of Pennsylvania, here holding the skull of Auroraceratops rugosus.

The quality throughout is high and most students of the Ceratopsia were evidently compelled to contribute interesting work. Some articles do stand out as especially interesting, strong, or both.

Peter Dodson’s opening article (the single contribution in section one) – ‘Forty years of Ceratophilia’ – provides an excellent, personal overview of ceratopsian studies of the past four decades. Dodson says that his apparent modern role as “the dean … of ceratopsian studies” (Dodson 2010, p. 3) is a sort of happy accident; we know, in fact, that he is deserving of this role, having led work on the group through innovation, excellent research and brilliant writing (Dodson 1996). The ‘genealogy of Dodson ceratopsian students’ is a really nice touch, paralleling the efforts of other palaeontologists to produce phylogenies of academic ‘relatedness’. [Adjacent image of Dodson from his blog.]

Among other articles that really stand out, Dodson, You and Tanoue’s article on the basicranium and palate anatomy of psittacosaurids and non-ceratopsid neoceratopsians includes some wholly new anatomical information. Scott Sampson and Mark Loewen’s chapter is an excellent review of ceratopsid biogeography and phylogeny. One of their main conclusions – perhaps surprising to those who think that there might now be too many ceratopsids in Late Cretaceous North America! – is that “diversity will increase greatly once less explored geographic regions and temporal intervals are subject to greater sampling” (Sampson and Loewen 2010, p. 423). Indeed, as anyone who’s even vaguely aware of horned dinosaur diversity will know, a surprising number of new (mostly North American) taxa have been named within the last few years.

Those many, many new taxa

The volume itself includes the names of no fewer than six new taxa (one of which represents a renaming, not a wholly new taxon).

Awesome life restoration of Coahuilaceratops magnacuerna, by Lukas Panzarin. Lukas's horned dinosaurs (and other ornithischians) are easily among the best and most accurate dinosaur life restorations ever produced.

Coahuilaceratops magnacuerna Loewen et al., 2010 is a new chasmosaurine from the Cerro del Pueblo Formation of Mexico, notable for its gigantic, massively robust postorbital horncores, among the largest of any ceratopsian. Phylogenetic analysis recovers it as a close relative of Anchiceratops and Arrhinoceratops (Loewen et al. 2010) (though see Sampson et al. 2010). The second new taxon, Diabloceratops eatoni Kirkland and Deblieux, 2010, is a new brow-horned centrosaurine from the Wahweap Formation of Utah, named for a spectacular skull. Its nasal horn is tiny, the epijugal processes on the cheeks are especially long, and slender, outwardly curved epoccipitals are present at the posterior edges of the parietals. As noted above, we would predict that centrosaurines began their history with short nasal horns and long brow horns: nevertheless, it is good to have this confirmed in the form of both Diabloceratops and the also recently described Albertaceratops. The authors report a second specimen that seems to represent a second, as-yet-unnamed Diabloceratops species.

Wonderful life restoration of Rubeosaurus by Lukas Panzarin, licensed under Creative Commons Attribution 2.5 Generic license.

Rubeosaurus McDonald & Horner, 2010 is the new generic name for the centrosaurine previously known as Styracosaurus ovatus. While the inclusion of this Upper Two Medicine Formation taxon in Styracosaurus had never previously been challenged (see Ryan et al. 2007), it is not, in fact, a close relative of Styracosaurus proper (S. albertensis) if new phylogenetic studies are to be believed. In fact it is closer to Einiosaurus within the pachyrhinosaur lineage. Today it seems all too easy to forget that, just 20 years ago, Pachyrhinosaurus canadensis was regarded as the sole representative of a peculiar and otherwise enigmatic centrosaurine lineage.

One of the first papers to report new pachyrhinosaurine taxa – Horner et al. (1992) – made news at the time by proposing that Styracosaurus, Rubeosaurus, Einiosaurus, Achelousaurus and Pachyrhinosaurus belonged to an anagenetic lineage, the evolution of which was ‘forced’ by a marine transgression that caused ‘habitat bottlenecking’ and consequent intense selection pressure and rapid evolution. McDonald & Horner (2010) note that this hypothesis is not redundant altogether since rapid evolution and the displacement effects of the marine transgression both seem to have occurred; nevertheless, new species and new studies seem to show that cladogenesis was occurring instead or in addition. Indeed, yet another member of the pachyrhinosaur lineage is described in another of the volume’s chapters. It’s an unnamed form from the Upper Dinosaur Park Formation. Given that it likely represents a new species of Pachyrhinosaurus, this clade alone (that is, Pachyrhinosaurus) now includes four species.

Skull reconstruction of Ojoceratops fowleri, incorporating both holotype squamosal and referred elements, produced by Robert Sullivan. Available under Creative Commons CC0 1.0 Universal Public Domain Dedication, from wikipedia.

Ojoceratops fowleri Sullivan & Lucas, 2010 is named for a left squamosal from the Ojo Alamo Formation of New Mexico’s San Juan Basin, identified as that of a Triceratops-like chasmosaurine. Already it has been suggested that O. fowleri may in fact be synonymous with Triceratops (Longrich 2011). One peculiarity of Sullivan & Lucas’s (2010) terminology is their use of ‘crown chasmosaurine’ (p. 177). Another new chasmosaurine – Medusaceratops lokii Ryan et al., 2010 – is named for parietal fragments from the Judith River Formation of Montana, originally referred to the centrosaurine Albertaceratops. Large, robust postorbital horns from the same bonebed are inferred to belong to it. Medusaceratops is the oldest reported chasmosaurine.

Skull reconstruction of Tatankaceratops sacrisonorum (from Ott & Larson 2010), with (B) showing it superimposed on the skull of Triceratops horridus.

Then there’s Tatankaceratops sacrisonorum Ott & Larson, 2010 from the Hell Creek Formation of South Dakota. The existence of a new chasmosaurine in the late Maastrichtian of western North America is interesting in view of debates about dinosaur diversity at this time and place, and it is inevitable that some will immediately identify the type and only specimen as a juvenile Triceratops (as anyone interested in dinosaurs will know all too well, debate continues as to whether some or all late Maastrichtian chasmosaurines represent distinct taxa, or growth stages of the same species). However, Tatankaceratops has proportionally short, subvertical postorbital horns that look different from both the far longer, posteriorly curving horns present in juvenile Triceratops and the even longer, anteriorly curving horns present in adult Triceratops (Horner & Goodwin 2006). The possibility that it might represent a dwarf form of Triceratops (Longrich 2011) appears plausible, as does the suggestion that it could be an aberrant individual. These ideas could perhaps be tested via histological analysis.

A new species of Archaeoceratops, A. yujingziensis, is also named in the volume. A redescription of the Montanoceratops cerorhynchus holotype and a new report of Prenoceratops from the Oldman Formation add new data points to the growing body of literature on leptoceratopsids.

Spinops sternbergorum, restored by Dmitry Bogdanov.

I should finish this discussion of new taxa by noting that descriptions of new horned dinosaur taxa have not exactly been restricted to this volume. Since this book has appeared, North America has yielded the additional new centrosaurine Spinops sternbergorum Farke et al., 2011 as well as the new chasmosaurines Utahceratops gettyi Sampson et al., 2010 and Kosmoceratops richarsoni Sampson et al., 2010. The new name Vagaceratops Sampson et al., 2010 has been given to the taxon originally described as Chasmosaurus irvinensis, and the somewhat controversial Mojoceratops perifania Longrich, 2010 and Titanoceratops ouranos Longrich, 2011 have also been named. Then there’s the Asian centrosaurine Sinoceratops zhuchengensis Xu et al., 2010, the new leptoceratopsids Zhuchengceratops inexpectus Xu et al, 2010, Gryphoceratops morrisoni Ryan et al., 2012 and Unescoceratops koppelhusae Ryan et al., 2012 and the protoceratopsid-grade taxa Ajkaceratops kozmai Ősi et al., 2010 and Koreaceratops hwaseongensis Lee et al., 2011.

Chasmosaurine phylogeny (showing positions of the 2010 taxa Utahceratops and Kosmoceratops) from Sampson et al. (2010). From PLoS ONE, licensed under Creative Commons Attribution License.

CMN 8547, mounted with a replica Anchiceratops skull. Photo courtesy of ReBecca Hunt-Foster.

One of my favourite chapters is Jordan Mallon and Robert Holmes’s description of CMN 8547, a near-complete chasmosaurine specimen (mounted at Ottawa’s Canadian Museum of Nature with a replica Anchiceratops skull), conventionally assigned to Anchiceratops due to the association of some supposedly diagnostic frill fragments. It turns out that those fragments aren’t all that diagnostic and that the taxonomic status of the specimen is uncertain. This is unfortunate (and needs resolving) since CMN 8547 is such a beautifully preserved, near-complete specimen. Its robust build, massive limbs and short tail are peculiar features that might suggest a hippo-like lifestyle.

Horns, frills, herding and lifestyle… a few unorthodox ideas

What of behaviour and ecology? The diverse and often spectacular horns, frills and other cranial structures present in the group have of course invited substantial speculation on social behaviour, sexual selection, the need to deal with contemporaneous theropods, thermoregulation and even acoustics. Needless to say, there is a huge amount of information here that will be of interest to those investigating horned dinosaur ecology, behaviour and palaeoenviromental preferences. Chapters include David Krauss and colleagues’ on the correlation between horn and frill morphology in chasmosaurines (parietal fenestrae are positioned just outside horn reach), Donald Henderson’s on niche partitioning as indicating by skull shape, and Rebecca Hunt and Andrew Farke’s on behavioural information as inferred from bonebeds. They conclude that, while there is good evidence for herding behaviour in some taxa, the evidence is not so good that we should make assumptions about herding across the clade.

Skull of Protoceratops andrewsi (though this one is without the sclerotic rings you need to determine eye size). Photo by Jordi Payà, from wikipedia.

Two behaviour-themed articles in particular stand out as unusual, and indeed the editors note in the preface that they are “sure to spark debate” (Ryan et al. 2010, p. xiii). Nick Longrich uses data from sclerotic ring size to propose that the unusually large-eyed Protoceratops might have been scotopic (or nocturnal). Given the abundance of specimens and quality of preservation, surprisingly little has been published on the palaeobiology of Protoceratops. Most work has concentrated on the supposed presence of sexual dimorphism and on the nests and eggs associated (sometimes incorrectly) with this dinosaur; its probable diet and habits have not been well explored. Schmitz & Motani (2011) used a similar technique to Longrich (2010) to analyse possible activity patterns in dinosaurs and other Mesozoic archosaurs, though did not report the same conclusions for Protoceratops. While this angle of analysis looks promising, already there are uncertainties given that Microraptor – inferred from eye form to be scotopic – is known to possess glossy feathers, a feature never present in extant scotopic species (Li et al. 2012). [Adjacent photo by Jordi Payà, licensed under Creative Commons Attribution-Share Alike 2.0 Generic license.]

Psittacosaurus preserved alongside the turtle Manchurochelys - proof of aquatic habits in a psittacosaurid! (irony). Image by Christopher, Tania and Isabelle Luna, from wikipedia (uploaded by FunkMonk). Released under Creative Commons Attribution 2.0 Generic license.

In the second ‘unusual’ chapter, Tracy Ford and Larry Martin put forward the surprising (but not wholly novel) proposal that Psittacosaurus was amphibious. The evidence used to support this idea is underwhelming; none of the features they point to are reliable indicators of aquatic habits. They make vague and unconvincing comparisons between the psittacosaurid forelimb and that of sea-turtles, cetaceans and other swimmers, imply that flexible hindlimb joints provide evidence for a swimming habit, suggest that gastroliths may have been present because of a role in buoyancy control, and draw attention to the presence of a laterally compressed tail skeleton and dorsally placed nostrils and orbits. [Adjacent photo by Christopher, Tania and Isabelle Luna]. Most or all of these features either do not link consistently with swimming habits, or are clearly present in other animals that do not, or did not, regularly swim. Their suggestion that the long bristle-like tail structures seen in one psittacosaurid specimen “may have supported a caudal fin that was somewhat analogous to the caudal fin in modern amphibians, such as the Hellbender … and tadpoles” is surprising (Ford & Martin 2010, p. 335). I’m not sure that it’s completely untenable, but the very slender, often overlapping fibres on the psittacosaurid tail clearly seem to be external to the epidermis, not sandwiched within it (as they’d have to be if they were within a continuous skin frill or fin). And, so… we have this vision (below, courtesy of Zach Miller of When Pigs Fly Returns).

Imaginatively restored aquatic psittacosaurid, by Zach Miller. Used with permission.

That famous psittacosaurid with the quilly tail - so far the only one known to have these structures (from Mayr et al. 2002). Did I ever say that Naish & Martill (2001) were just about the first to illustrate this specimen? Yeah, like that means something. And I do use the term "illustrate" rather loosely...

As David Eberth points out in his review of palaeoenvironmental associations and taphonomy, the unusually high number of fully articulated psittacosaurid specimens suggests that their carcasses did not endure a lengthy ‘bloat-and-float’ phase prior to burial, an observation apparently at odds with Ford and Martin’s model. Nevertheless, the idea is not so bizarre that it can be dismissed entirely without consideration. As the authors note, the proposal may not apply to all psittacosaurid species, and we have to keep in mind the ecological and behavioural diversity seen in certain extant ‘genera’. Furthermore, amphibious habits should not be ruled out entirely for all ceratopsians. The good news is that we have so many psittacosaurid specimens that testing this hypothesis would be very simple, should someone feel it worthy of proper investigation.

An accompanying CD ROM includes two lengthy contributions that would not have worked well as published articles but are worth having for completists: Tracy Ford’s stratigraphically arranged specimen list, and Darren Tanke’s substantial, date-arranged compilation of discoveries, events and biographies relevant to horned dinosaur research in Alberta.

All in all, New Perspectives on Horned Dinosaurs is arguably the most significant dinosaur book to appear in recent years, and this is against a lot of competition. It demonstrates – just in case there was any doubt – that feathered maniraptorans and tyrannosaurids are not the only sections of the dinosaur tree where exciting research and discoveries are happening. In term of density, significance, novelty and sheer volume of content, and in the quality of the text and illustrations, it’s outstanding – certainly head and shoulders above the many other dinosaur-themed volumes published by Indiana University Press. It does a superb job of capturing the status of horned dinosaur research at it was in the first decade of the present century. Surely, with this much exciting research and investigation underway, there are many surprises yet to come.

Ryan, M. J., Chinnery-Allgeier, B. J. & Eberth, D. A. (eds) New Perspectives on Horned Dinosaurs: The Royal Tyrrell Museum Ceratopsian Symposium. Indiana University Press, Bloomington. Hardback, index, refs, pp. 624. ISBN 978-0-253-35358-0. Here on Amazon, here on Amazon.co.uk.

There’s now quite a lot on Tet Zoo about horned dinosaurs. For previous articles, see…

• Tet Zoo picture of the day # 11 (or, speculations on omnivory in ceratopsids)
• Pentaceratops: that’s quite the skull
• SVPCA 2007: dinosaurs attack (includes stuff on Psittacosaurus)
• A month in dinosaurs (and pterosaurs): 3, Minotaurasaurus and giant chasmosaurines
• Udanoceratops tschizhovi, the basics
• No-one talks about Anchiceratops, boo hoo
• Zuniceratops and the early acquisition and alleged dimorphism of ceratopsian brow horns
• A very alternative view of horned dinosaur anatomy
• Ceratopsian dinosaurs: cheeky or beaky?
• Greek-nosed first-horned face and the ‘bagaceratopids’
• What they’re saying about The Great Dinosaur Discoveries
• Did dinosaurs and pterosaurs practise mutual sexual selection?
• Greg Paul’s Dinosaurs: A Field Guide

Refs – -

Dodson, P. 1996. The Horned Dinosaurs. Princeton University Press, Princeton, NJ.

- . 2010. Forty years of Ceratophilia. In Ryan, M. J., Chinnery-Allgeier, B. J. & Eberth, D. A. (eds) New Perspectives on Horned Dinosaurs: The Royal Tyrrell Museum Ceratopsian Symposium. Indiana University Press, Bloomington, pp. 3-13.

Ford, T. & Martin, L. D. 2010. A semi-aquatic life habit for Psittacosaurus. In Ryan, M. J., Chinnery-Allgeier, B. J. & Eberth, D. A. (eds) New Perspectives on Horned Dinosaurs: The Royal Tyrrell Museum Ceratopsian Symposium. Indiana University Press, Bloomington, pp. 328-339.

Horner, J. R. & Goodwin, M. B. 2006. Major cranial changes during Triceratops ontogeny. Proceedings of the Royal Society, London B 273, 2757-2761

- ., Varricchio, D. J. & Goodwin, M. B. 1992. Marine transgressions and the evolution of Cretaceous dinosaurs. Nature 358, 59-61.

Li, Q., Gao, K.-Q., Meng, Q., Clarke, J. A., Shawkey, M. D., D’Alba, L., Pei, R., Ellison, M., Norell, M. A. & Vinther, J. 2012. Reconstruction of Microraptor and the evolution of iridescent plumage. Science 335, 1215-1219.

Loewen, M. A., Sampson. S. D., Lund, E. K., Farke, A. A., Aguillón-Martínez, M. C., De Leon, C. A., Rodríguez-De La Rosa, R. A., Getty, M. A. & Eberth, D. A. 2010. Horned dinosaurs (Ornithischia: Ceratopsidae) from the Upper Cretaceous (Campanian) Cerro del Pueblo Formation, Coahuila, Mexico. In Ryan, M. J., Chinnery-Allgeier, B. J. & Eberth, D. A. (eds) New Perspectives on Horned Dinosaurs: The Royal Tyrrell Museum Ceratopsian Symposium. Indiana University Press, Bloomington, pp. 99-116.

Longrich, N. R. 2010. The function of large eyes in Protoceratops: a nocturnal ceratopsian? In Ryan, M. J., Chinnery-Allgeier, B. J. & Eberth, D. A. (eds) New Perspectives on Horned Dinosaurs: The Royal Tyrrell Museum Ceratopsian Symposium. Indiana University Press, Bloomington, pp. 308-327.

- . 2011. Titanoceratops ouranos, a giant horned dinosaur from the Late Campanian of New Mexico. Cretaceous Research 32, 264-276.

Mayr, G., Peters, D. S. & Plodowski, G. 2002. Bristle-like integumentary structures at the tail of the horned dinosaur Psittacosaurus. Naturwissenschaften 89, 361-365.

McDonald, A. T. & Horner, J. R. 2010. New material of “Styracosaurus” ovatus from the Two Medicine Formation of Montana. In Ryan, M. J., Chinnery-Allgeier, B. J. & Eberth, D. A. (eds) New Perspectives on Horned Dinosaurs: The Royal Tyrrell Museum Ceratopsian Symposium. Indiana University Press, Bloomington, pp. 156-168.

Naish, D. & Martill, D. M. 2001. Boneheads and horned dinosaurs. In Martill, D. M. & Naish, D. (eds) Dinosaurs of the Isle of Wight. The Palaeontological Association (London), pp. 133-146.

Ryan, M. J., Holmes, R. & Russell, A. P. 2007. A revision of the late Campanian centrosaurine ceratopsid genus Styracosaurus from the Western Interior of North America. Journal of Vertebrate Paleontology 27, 944-962.

- ., Chinnery-Allgeier, B. J. & Eberth, D. A. 2010. Preface. In Ryan, M. J., Chinnery-Allgeier, B. J. & Eberth, D. A. (eds) New Perspectives on Horned Dinosaurs: The Royal Tyrrell Museum Ceratopsian Symposium. Indiana University Press, Bloomington, pp. xiii-xiv.

Sampson, S. D. & Loewen, M. A. 2010. Unravelling a radiation: a review of the diversity, stratigraphic distribution, and evolution of horned dinosaurs (Ornithischia: Ceratopsidae). In Ryan, M. J., Chinnery-Allgeier, B. J. & Eberth, D. A. (eds) New Perspectives on Horned Dinosaurs: The Royal Tyrrell Museum Ceratopsian Symposium. Indiana University Press, Bloomington, pp. 405-427.

- ., Loewen, M. A., Farke, A. A., Roberts, E. M., Forster, C. A., Smith, J. A. & Titus, A. L. 2010. New horned dinosaurs from Utah provide evidence for intracontinental dinosaur endemism. PLoS ONE 5(9): e12292. doi:10.1371/journal.pone.0012292

Schmitz, L., & Motani, R. (2011). Nocturnality in Dinosaurs Inferred from Scleral Ring and Orbit Morphology Science, 332 (6030), 705-708 DOI: 10.1126/science.1200043

Sereno, P. C. 1998. A rationale for phylogenetic definitions, with application to the higher-level taxonomy of Dinosauria. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 20, 41-83.

Sullivan, R. M. & Lucas, S. G. 2010. A new chasmosaurine (Ceratopsidae, Dinosauria) from the Upper Cretaceous Ojo Alamo Formation (Naashoibito Member), San Juan Basin, New Mexico. ). In Ryan, M. J., Chinnery-Allgeier, B. J. & Eberth, D. A. (eds) New Perspectives on Horned Dinosaurs: The Royal Tyrrell Museum Ceratopsian Symposium. Indiana University Press, Bloomington, pp. 169-180.

Wolfe, D. G. & Kirkland, J. I. 1998. Zuniceratops christopheri n. gen. & n. sp., a ceratopsian dinosaur from the Moreno Hill Formation (Cretaceous, Turonian) of west-central Montana. New Mexico Museum of Natural History and Science Bulletin 14, 303-317.

Regular readers might have noticed that I’m not all that keen on covering stories that get massive, global exposure across the blogosphere. Consequently, sexy dinosaur news is mostly ignored here. Sometimes, though, I suppose I have to make an exception. Maybe I have a duty to, since the Tet Zoo audience includes more than an average number of dinosaur specialists (meaning that comments and discussions can often be pretty in-depth; more in-depth than they are elsewhere on the web). Furthermore, Tet Zoo is unlike many other sites that cover Mesozoic dinosaurs in that it appeals to many readers who, while zoologically informed, aren’t Mesozoic specialists and hence don’t necessarily get barraged by the same amount of ‘new dinosaur’ announcements that I (and other dinosaur workers) do. Anyway, enough with the preamble…

Brilliant life restoration of four Yutyrannus individuals, by Brian Choo. Is that snow on the ground? You know, I think it is.

Today sees the publication of Yutyrannus huali Xu et al., 2012 in the hallowed pages of Nature. Known from three well-preserved specimens, it’s yet another feathered theropod dinosaur from the Lower Cretaceous Yixian Formation of Liaoning Province, China. But it isn’t a dromaeosaurid, early bird or other maniraptoran – it’s a giant tyrannosauroid.

Substantially simplified phylogeny of tyrannosaurs, in this case from Benson et al. (2010). Note difference between Tyrannosauridae and the far more inclusive Tyrannosauroidea.

That is, a member of the same major group of coelurosaurian theropods as Tyrannosaurus and its close kin, but not a member of Tyrannosauridae (the highly modified, short-armed, two-fingered tyrannosauroid clade that includes Tyrannosaurus, Albertosaurus and so on). Tyrannosaurids are mostly big to very big, but non-tyrannosaurid tyrannosauroids were often small to medium-sized (1-4 m being a very approximate range). Anyone who follows me on twitter will know that I spent a significant portion of my life within recent months thinking and writing about tyrannosauroids and not much else.

Skull of Yutyrannus in right lateral view. You should be able to see the rugose dorsal crest along the length of the snout.

Anyway, as a non-tyrannosaurid tyrannosauroid, Yutyrannus is typical in having three-fingered hands, proportionally large bony nostril openings, and distal hindlimb segments that are not especially elongate for the size of the animal (Xu et al. 2012). Diagnostic (= unique) characters include the presence of a peculiar rugose crest along the dorsal midline of the snout and a bony boss on the postorbital bone. We’ll get back to those features in a minute, since there’s something very familiar about them.

The feathers!! The feathers!!

As usual, the main gee-whiz points about Yutyrannus are already being widely discussed. We’ve known for a while (since the publication of Dilong paradoxus in 2004) that at least some tyrannosauroids possess ‘stage 1 feathers’ (Xu et al. 2004). That is, filamentous integumentary structures that seem to be evolutionary precursors to the true, complex feathers that evolved elsewhere within coelurosaurian theropods. Yutyrannus is another feathery/filamenty tyrannosauroid, but it’s remarkable in being huge – it’s about 9 m long, meaning that here is the first GIANT feathery/filamenty tyrannosauroid.

A now historic representation of Eotyrannus, from an online 2001 article by Erik Stokstad. OMG, is that scaly skin?? Oooh, the regret... The moron artist should have known better (ps - irony. Artist = some guy called Naish).

As Xu et al. (2012) note, there are other giant coelurosaurs that would also have been feathered (examples: the huge oviraptorosaur Gigantoraptor, the therizinosaur Therizinosaurus, and perhaps the gargantuan ostrich dinosaur Deinocheirus), but Yutyrannus is the first really big one to actually have the structures preserved. If feathers of this sort were present in Dilong and Yutyrannus, it stands to reason that tyrannosauroids close to these two in the phylogeny would have been feathered as well. The tyrannosauroid I know best – Eotyrannus lengi – is thus depicted as fully fuzzed-up by Xu et al. (2012). I can live with that.

The structures in Yutyrannus are mightily impressive, being tightly massed together and as much as 15 cm long. While none of the three Yutyrannus specimens preserves a complete integumentary covering (if you think that’s weird or suspicious or something, go look at rotting animal carcasses some time), their filamentous structures are preserved in association with the neck, hip region, tail and elsewhere. I thus find it totally reasonable to imagine that the covering was complete across much of the body. These were big, shaggy tyrannosaurs in feathery coats – exactly the sort of thing that artists have been drawing and painting for years now. Huh, stupid artists.

Brian Choo's life restoration again, this time without the pretty background.

If you’re wondering, claims that the integumentary structures present in coelurosaurian theropods might actually be decayed collagen fibres (yes, collagen fibres – - I’m not kidding!!) are not likely and never have been.

Cold adapted tyrannosaurs?

Maniraptorans playing in the snow. An, err, somewhat speculative scene by Luis V. Rey.

Regular readers will perhaps know that I (and others) have been saying for a while that Mesozoic Earth was not the global hothouse that many have long assumed. There’s evidence for cool continental interiors and poles during parts of the Jurassic and Cretaceous (Barron & Washington 1982, , Sloan & Barron 1990, Sellwood et al. 1994), sea surface temperatures that were similar to those of the cold, modern north Pacific and Atlantic (Van de Schootbrugge et al. 2000), and some researchers think that there were glaciation events – that’s right, I said glaciation events – late in the Jurassic and early in the Cretaceous (Dromart et al. 2003, McArthur et al. 2007).

Recently, data has been compiled indicating that the Liaoning region was cool during the Early Cretaceous – its average temperature being about 10 degrees C (Amiot et al. 2011). If you insist that dinosaurs were scaly and ‘cold-blooded’, this idea of insulated big theropods running around in cool climates must seem silly, but if you disregard preconceptions and biases, I think we can now take very seriously the idea that insulated, endothermic dinosaurs were inhabiting cool, or cold, habitats.

Xu et al. (2012) suggest that Yutyrannus was ‘cold adapted’, and that the tyrannosauroids of warmer climes were scaly-skinned and not insulated in the same way. That might be correct, but it’s worth pointing out that we actually know little or nothing about the integumentary covering of tyrannosauroids outside of the Liaoning taxa. There are a few patches of scaly skin alluded to here and there, but nothing substantial, so far. I’m with Andrea Cau (and others) on the idea that ‘fuzziness’ was a gradational thing in archosaurs, with fuzz and feathering mostly (but not wholly) grading out as body size increased. That’s nicely illustrated in Andrea’s diagram below: the redder parts of the diagram are the more scaly/less fuzzy ones, while the purple parts are fuzziest/featheryest (the skeletal reconstructions were produced by Greg Paul, Jaime Headden, Scott Hartman, Marco Auditore and Lukas Panzarin). For a sharper version go here on Theropoda. And I have no idea if “featheryest” is a word.

Yutyrannus the… carcharodontosaurian?

I must confess to being somewhat sceptical of the tyrannosauroid identification for Yutyrannus. I reviewed this paper (to those who don’t know: I did my PhD thesis on basal tyrannosauroids), and noted immediately that Yutyrannus actually resembles carcharodontosaurian allosauroids in some respects. Those of you who know theropods, or who have very good memories, will recall the brouhaha back in 2010 when the carcharodontosaurian Concavenator was described from Lower Cretaceous rocks of Spain (Ortega et al. 2010). Concavenator is an awesome fossil, but what really guaranteed its appearance in the top-tier glamour mags is the claim that raised bumps on its ulnae are quill node homologues – that is, evidence for proto-feathers on its arms. I was sceptical of this at the time (the ‘quill node homologues’ look like bumps on an intermuscular line to me), and am sceptical of it still.

Life restoration of Concavenator by Raul Martin. With feathery, filamenty things on the arms.

Anyway, it’s somewhat ironic – given its alleged featheryness – that Concavenator should raise its head, so to speak, in the discussion here. But I think that Concavenator and Yutyrannus are somewhat alike. The skulls of both animals are particularly similar. Both possess a rugose dorsal crest along each nasal bone, decorated on its lateral surface by a series of subcircular (?pneumatic) recesses, both have a rounded posterodorsal boss (possibly with a concavity on its lateral surface) on the postorbital bone, and, in both, a projection from the postorbital bar invades the orbit.

Other elements of these taxa are also similar. Concavenator is superficially tyrannosauroid-like in possessing a small anterodorsal concavity on the anterior lobe of the ilium (also present in Yutyrannus) and the shaft of its ischium appears more slender than the shaft of the pubis (a feature noted in the Yutyrannus paper to be a tyrannosauroid-like character).

While the authors took account of the carcharodontosaurian-like features of Yutyrannus and modified their phylogenetic analysis accordingly, they stuck to their guns about it being a tyrannosauroid. I agree that Yutyrannus is tyrannosauroid-like in many features but, I dunno, I think it needs to be included within a larger and more comprehensive data set. Why didn’t I do this myself when I was reviewing the paper? Several reasons. One being that – so long as a given proposal isn’t fatally flawed or shot full of holes – you can’t stop authors from coming up with their own favoured phylogenetic hypothesis, even if you disgree with it. I should note that my suggestion here is just a suggestion, not an outright challenge, and I’m not all that confident about being right. I’m just sceptical.

Mark Witton's fuzzy styracosaur again. Hey, weirder stuff has happened. Oh yeah, and it's eating a dead tyrannosauroid.

Anyway, none of this stops Yutyrannus from being an awesome dinosaur discovery that will get lots of attention in future publications. And it raises lots of questions. Most evidence indicates that the tyrannosauroids of the Early Cretaceous were generally small and ecologically ‘inferior’ to contemporaneous megalosauroids and allosauroids. If Yutyrannus is a tyrannosauroid, does it show that things were more complex, and that big tyrannosauroids lived alongside big megalosauroids and/or allosauroids in some or many places? Or is it that tyrannosauroids were ‘controlling’ regions (like – the cooler regions?) where those other groups were absent? Were tyrannosaurids perhaps better adapted for cool/cold climates than other big-bodied theropod lineages, and could this have been key to tyrannosauroid success? Were the cranial bossess and nasal crest we see in Yutyrannus sexually selected characters, and is their presence in more than one individual an indication of mutual sexual selection? (Hone et al. 2012). Perhaps most interestingly of all, will more big, fuzzy dinosaurs be announced in future? Place your bets.

For previous Tet Zoo articles on some of the subjects mentioned here, see…

• Feathers and filaments of non-avian dinosaurs, part I
• Feathers and filaments of dinosaurs, part II
• 100 years of Tyrannosaurus rex
• Tyrant dinosaurs were not a Northern Hemisphere speciality: they also colonised Australia!
• Concavenator: an incredible allosauroid with a weird sail (or hump)… and proto-feathers?
• Did dinosaurs and pterosaurs practise mutual sexual selection?

Refs – -

Amiot, R., Wang, X., Zhou, Z., Xiaolin Wang, X., Buffetaut, E., Lécuyer, C., Ding, Z., Fluteau, F., Hibino, T., Kusuhashi, N., Mo, J., Suteethorn, V., Yuanqing Wang, Y., Xu, X. & Zhang, F. 2011. Oxygen isotopes of East Asian dinosaurs reveal exceptionally cold Early Cretaceous climates. Proceedings of the National Academy of Sciences 108, 5179-5183.

Barron, E. J. & Washington, W. M. 1982. Cretaceous climate: a comparison of atmospheric simulations with the geological record. Palaeogeography, Palaeoclimatology, Palaeoecology 40, 103-133.

Benson, R. B. J., Barrett, P. M., Rich, T. H. & Vickers-Rich, P. 2010. A southern tyrant reptile. Science 327, 1613.

Dromart, G., Garcia, J.-P., Picard, S., Atrops, F., Lécuyer, C. & Sheppard, S,. M. F. 2003. Ice age at the Middle-Late Jurassic transition? Earth and Planetary Science Letters 213, 205-220.

Hone, D. W. E., Naish, D. & Cuthill, I. C. 2012. Does mutual sexual selection explain the evolution of head crests in pterosaurs and dinosaurs? Lethaia 45, 139-156

McArthur, J. M., Janssen, N. M. M., Reboulet, S., Leng, M. J., Thirlwall, M. F. & van de Schootbrugge B. 2007. Palaeotemperatures, polar ice-volume, and isotope stratigraphy (Mg/Ca, d18O, d13C, 87Sr/86Sr): the Early Cretaceous (Berriasian, Valanginian, Hauterivian). Palaeogeography, Palaeoclimatology, Palaeoecology 248, 391-430.

Ortega, F., Escaso, F. & Sanz, J. L. 2010. A bizarre, humped Carcharodontosauria (Theropoda) from the Lower Cretaceous of Spain. Nature 467, 203-206.

Sellwood, B. W., Price, G. D. & Valdes, P. J. 1994. Cooler estimates of Cretaceous temperatures. Nature 370, 453-455.

Sloan, L. C. & Barron, E. J. 1990. Equable climates during Earth history. Geology 18, 489-492.

Van de Schootbrugge, B., Föllmi, K. B., Bulot, L. G. & Burns, S. J. 2000. Paleoceanographic changes during the early Cretaceous (Valanginian-Hauterivian): evidence from oxygen and carbon stable isotopes. Earth and Planetary Science Letters 181, 15-31.

Xu, X., Norell, M., Kuang, X., Wang, X., Zhao, Q., & Jia, C. (2004). Basal tyrannosauroids from China and evidence for protofeathers in tyrannosauroids Nature, 431 (7009), 680-684 DOI: 10.1038/nature02855

- ., Wang, K., Zhang, K., Ma, Q., Xing, L., Sullivan, C., Hu, D., Cheng, S. & Wang, S. 2012. A gigantic feathered dinosaur from the Lower Cretaceous of China. Nature 484, 92-95.

Among the most controversial and remarkable of living tetrapods are the bizarre amphisbaenians: a group of fossorial, long-bodied carnivorous animals with reduced or absent limbs, spade-shaped or bullet-shaped skulls strongly modified for burrowing, and an annulated body where distinct, regularly arranged transverse segments give the animals a worm-like appearance. [Adjacent image combines diagram from Gans (1974) with photograph of Amphisbaena bakeri by Father Alejandro Sanchez, used with permission]. Until recently it was generally thought that amphisbaenians are reptiles, and part of Squamata (the reptile group that includes snakes and lizards). But, in a fascinating case of multi-disciplinary co-operation involving genetics, neophenetics, and good old-fashioned critical thinking, intuition and balls, a daring group of feisty young zoologists have challenged the old guard of the ‘Mammals are Derived Synapsids, y’all’ (or MADsy) school of thought, and have demonstrated that these are not mere squirmy reptiles. They are, in fact, the true ancestors of mammals.

The first authors to formally suggest a non-reptilian affinity for amphisbaenians published their observations in the 1940s and 50s (Zangerl 1945, Kesteven 1957). In a classic case of textbook orthodoxy triumphing over the brilliantly shining light of massive truthfulness, their work was all but ignored in what amounts to a conspiracy of some sort, and the more media-friendly idea that mammals descended from Palaeozoic synapsids like Dimetrodon took centre stage (e.g., Remor 1979, 1980). However, animals like Dimetrodon clearly come loaded with too much baggage to serve as mammal ancestors! They are way too big (Dimetrodon was about the size of a man!), and the two or three characters they share with mammals must have arisen by way of similar lifestyles! We are therefore left searching for the true mammal ancestor! And this is where amphisbaenians come in!!

Mongolian death-worm? No, the amphisbaenid amphisbaenian Amphisbaena alba (photo by Diogo B. Provete). Click to enlarge.

After being all but ignored in the literature on mammal origins for several decades, an impressive list of unassailable genetic papers (Koch & Wyman 2007 and references therein) have now shown that mammals are actually nested within the amphisbaenian radiation (Koch & Wyman 2007 and references therein). [Adjacent photo by Diogo B. Provete, from wikipedia.] Initially this might sound remarkable but, after thinking about it a bit, a number of workers have reassessed some of the older literature – much of which shows that the character evidence linking mammals to amphisbaenians is compelling, well documented, well established, meticulously detailed, and deserving of other such terms (Koch & Wyman 2007 and references therein). We don’t need to worry about looking critically at the genetic data (Koch & Wyman 2007 and references therein), as few people really understand genetic stuff anyway, and no one really reads the methods sections of papers anymore (Koch & Wyman 2007 and references therein).

The evidence from behaviour

Relatively little is known about amphisbaenian biology and behaviour. What we do know, however, shows that these animals share more in common with mammals than with other amniotes. In one of the few taxa for which a large amount of data on population structure has been collected, Bipes (the Mexican limbed amphisbaenians), Papenfuss (1982) concluded that the animals were K-selected, exhibiting delayed maturity, small clutch size, and non-annual reproduction. Does this sound like typical reptile behaviour to you? Of course not, it is part of the overwhelming body of evidence linking amphisbaenians to mammals.

Mexican limbed amphisbaenian: Ajolote (Bipes)

Among mammals, an incredible plethora of tunnel-excavating habits continue to be practised even by more recently evolved lineages, and are manifested by such activities as trench warfare, the Channel Tunnel, the London underground, the salt-mining culture of the elephants of Mount Elgon, the propensity for humans to dig tunnels on sandy beaches, and that dude who lived in a hole for years. Is it a coincidence that people seek out caves and tunnels to explore and wonder at while on holiday? Many people admit to psychiatrists that they dream of burrowing and tunnelling. And let us not forget that millions of people travel to and from work on a daily basis via subterranean tunnels, showing a statistical preference for this mode of travel rather than for the supra-terrestrial environment shunned by our amphisbaenian ancestors. In all this data, we have irrefutable evidence pointing to our subterranean, amphisbaenian ancestry.

The evidence from soft-tissue anatomy

In their often pinkish colour, amphisbaenians share an important synapomorphy with mammals, most of which are pinkish when shaved. This character is best expressed in Bipes, and in the desert sharks and allghoi-khorkhoi (more on these taxa in a minute), but it is also widespread within amphisbaenid amphisbaenians. Some amphisbaenian taxa are capable of autotomy (the defensive shedding of the tail when it’s grabbed by a predator), another similarity shared with mammals. An often overlooked fact concerning the amphisbaenian body is that, despite the simple, worm-like shape, these animals have a highly complex musculature, as was noted by Camp (1923) and Gans (1978). It has often been stated that mammals owe their complex musculature to the fact they went through a small-bodied cryptic phase, in which they had to clamber over stones, roots, dinosaur toes etc., but in reality mammalian musculature has been derived directly from the condition present in amphisbaenians.

Heterocephalus: without doubt, an amphisbaenian-like mammal.

A loose skin that is mobile relative to the underlying tissues is also shared by amphisbaenians and fossorial mammals like naked mole-rats Heterocephalus glaber and pet hamsters. Indeed, amphisbaenian-like Heterocephalus [shown in adjacent image] is one of the most basal of mammals, its poikilothermic physiology (Buffenstein & Yahav 1991) demonstrating that poikilothermy was retained by ancestral mammals and only later modified as mammals began to take to the supra-terrestrial environment. All of this is depicted in the phylogram shown below – truly, unassailable science (sensu Olson 2002).

The evidence from skeletal anatomy

The short, rigid skulls of amphisbaenians are shockingly mammal-like. Indeed Carl Gans, the great expert on amphisbaenians, has repeatedly noted the strong similarity evident between amphisbaenian and mammal skulls (Gans 1969, 1974), drawing special attention to this in his 1974 volume Biomechanics: An Approach to Vertebrate Biology. Here, Gans noted the strong similarity between the skull of the trogonophid Agamodon and that of a cat (Gans 1974, p. 134, Fig. 4-10), describing how “a comparison between the skull of a cat and that of the trogonophid Agamodon documents the … similarity which … [shows] that the skulls of amphisbaenians most [resemble] those of mammals”. Later, Gans showed how the jaw motion of Agamodon is strikingly like that of elephants and other placental mammals (Gans 1974, p. 178, Fig. 4-40: the diagram shown at the top of the article).

The cranial bones of amphibaenians overlap one another extensively, meeting at strongly interdigitated sutures, again as they do in mammals. A whopping big coronoid process on the amphisbaenian mandible is again a shared character present in mammals, though admittedly this character is labile and has coincidentally popped up in various reptile lineages. Labouring under the mistaken assumption of a reptilian identity for amphisbaenians, zoologists have interpreted one of the most unusual features of the amphisbaenian skull – the extracolumella, a structure that links the side of the jaw to the ear – as a character of reptilian flavour, and perhaps as a unique evolutionary solution to the problem of sound conduction. However, Wever & Gans (1973) noted that the extracolumella is “not homologous to the structure of the same name in lizards” (p. 189), and Gans (1978) reiterated that it “is not directly homologous with the structure of that name seen in the Sauria and Sphenodon” (p. 377).

Amphisbaenian skull from digimorph, extracolumella shown in red.

Given that the ear bones of mammals are really complicated and are known to have somehow been modified from bits and pieces that used to be at the back of the lower jaw, it is unarguably logical to conclude that the freaky stuff going on in the amphisbaenian lower jaw and ear region offers us the real answer to mammalian ear evolution. By employing OOA (obfuscatory ocular analysis, or squinting), Ratsarse (2006) was able to show that amphisbaenian skulls are highly similar to those of mammals, the extracolumella looking like some sort of magic streak showing the viewer where all the funky evolutionary pazazz is occurring, especially when it’s highlighted in red as shown here. [Adjacent image of Geocalamus acutus from here on digimorph.]

Gans (1974). He knew.

Gans (1960, 1974) noted that, by depicting amphisbaenian skulls within transformation grids, the long and low, superficially squamate-like skulls of trogonophid amphisbaenians could be ‘morphed’ into the short, mammal-like skulls of taxa like Agamodon (the latter one of the key taxa demonstrating the true affinities of the Amphisbaenia). In fact, Gans simply got it the wrong way round, as we now know that the mammal-like Agamodon skull is primitive, and it is the squamate-like skull of Trogonophis and similar taxa that are derived. Any idiot can see this, it’s soooooooo obvious.

Another important feature that amphisbaenians share with mammals is the mysterious reduction of the pelvis. Whereas animals with reduced limbs typically lose their forelimbs and retain their hindlimbs (look at pythons and reduced-limbs lizards like some cordylids), some amphisbaenians retain forelimbs and a pectoral girdle, yet have completely lost the hindlimbs and only possess a relictual pelvis consisting of the ilium alone (Kearney 2002). Where else do we see this unusual pattern? In reptiles? Heavens, no, in mammals: just look at whales! Again this is an important shared character demonstrating the amphisbaenian heritage of Mammalia. This self-evident evidence is so evident that it has been ignored by many so-called experts; they have brushed it under the carpet, donned blinkers, thrown up their arms, and uttered ‘harrumph!’. In short, anything other than consider the obvious TRUTH.

A transition lost in time, ignored by orthodoxy

Psammonarus, from Dixon (1988).

Elsewhere in classic zoological literature, we can see that Dixon (1981) knew full well that amphisbaenians gave rise to mammals, and that the earliest mammals were fossorial amphisbaenian-like creatures: an assertion demonstrated by the existence within Dixon’s work of the desert sharks Psammonarus, sausage-shaped predators that were perfect evolutionary intermediates between the Amphisbaenia and Mammalia.

While at least some of the creatures described by Dixon (1981) are generally accepted as hypothetical, recent work has demonstrated that a surprising number of genuine creatures were included as well. We can assume that, eschewing the hidebound, tediously slow, reactionary work of the peer-reviewed journal system, Dixon snuck genuine zoological discoveries into his work, knowing full well that smart people would spot them for the real creatures they obviously are. The swimming ‘monkey’ Natopithecus ranapes Dixon, 1981, for example, has since been shown to be a pre-emptive description of the aquatic primates later described by Coleman & Huyghe (1999), the Flooer Florifacies mirabila was an accurate description of the flower-faced snouters Cephalanthus described by Stümpke (1957)*, while the insectivorous tree drummers (Proboscicuncus) were evidently based on somewhat garbled descriptions of the remarkable Peruvian murid Rhagomys longilingua (Luna & Patterson 2002).

* Unfortunately, Dixon was unaware of Stümpke’s prior description of this form (D. Dixon, pers. comm.). Rhinogradentians were covered on Tet Zoo ver 2 here and here.

We can therefore assume that Dixon learnt of Psammonarus at one of the many palaeontology conferences he attends. Admittedly, no published abstract or article attests to the description of such a creature, and we are forced to conclude that it was deleted from the place of publication and its authors assassinated; possibly, those in control of the MADsy conspiracy somehow travelled back in time and erased Psammonarus from the technical literature, as this is the sort of thing they do.

Accurate portrait of Dinovermis mackerlei.

Evidently closely related to Psammonarus, and even closer to Mammalia than are other amphisbaenians, is the specialised relict form of Mongolia and Kazakhstan, the Allghoi-khorkhoi or Mongolian death-worm Dinovermis mackerlei, a controversial animal suggested to be mythical or semi-fictional by some, but known to be real by those who care. Originally suggested to be an annelid or burrowing snake (Shuker 2003), the truncated tail, pinkish hue, annulated body, amphisbaenian-style demeanour, and DNA of the Allghoi-khorkhoi have since demonstrated its amphisbaenian identity (Koch & Wyman 2007 and references therein).

Dinovermis is of course not called the death-worm for nothing. It is venomous, and here we find the key synapomorphy linking this animal with mammals. Not only are living primitive mammals – like monotremes, shrews and solenodons – venomous, we now know that venomosity was widespread in early mammals (Hurum et al. 2006). Only in the light of the amphisbaenian origin model does mammalian venomosity make any sort of sense.

Giant otter shrew (Potamogale velox), as illustrated by Paul Belloni Du Chaillu in 1868.

Dinovermis is also electrogenic (that is, able to generate an electric field), a fact unnervingly similar to the fact that electroreceptive abilities are present in those primitive mammals, the monotremes (and perhaps also in moles and other mammals). Electrogenic fish usually have electroreceptive abilities, so we can safely infer that electrogenic and electroreceptive abilities were primitive for mammals, but that the electrogenic ability was mostly lost. It wasn’t entirely lost, as some African people think that the Giant otter-shrew Potamogale velox is electrogenic. This will doubtless prove correct, and provide yet more affirmation of the amphisbaenian origin model.

Given that textbook dogma stakes matter-of-fact that mammals supposedly evolved somehow from quadrupedal synapsids of the Palaeozoic, we have to wonder why so many zoologists have fooled themselves into relying on cold, stony fossils, and not on the obvious living evidence that we can glean from the creatures around us today. We have learnt that we need not concern ourselves with Sineoamphisbaena from Upper Cretaceous Mongolia, and other fossils allegedly linked to the amphisbaenians. You might be surprised to hear this coming from a palaeontologist but, well, that’s just how it is. So fossils are vastly over-rated for this sort of thing and can be safely ignored. In fact, they may as well not exist.

The presence within amphisbaenians of retractile hemipenes, a squamate-style kidney, a transerve cloacal slit, keratinised scales, a telencephalic roof divided into three cortices, and a squamate-like Jacobson’s organ, are all clearly convergences with squamates. Some scientists, probably looking down from ivory towers or hiding behind the thick hedges of ivy that covers the walls of their colleges, and generally hoping to maintain the MADsy model, continue to deny this, as they have for a while now. But, like Saruman in the second Lord of the Rings movie, it is only a matter of time before the ents of justice arrive, and demonstrate the true, amphisbaenian origin of Mammalia. Then we will be free!

The digging style employed by Bipes, from Gans (1974). Clear evidence of mammalian affinity.

Apologies to long-term readers – have had no time to prepare anything new.

Refs – -

Buffenstein, R. & Yahav, S. 1991. Is the naked mole-rat, Heterocephalus glaber, an endothermic yet poikilothermic mammal? Journal of Thermal Biology 16, 227–232.

Camp, C. L. (1923). Classification of the lizards Bulletin of the American Museum of Natural History, 48, 289-481

Coleman, L. & Huyghe, P. 1999. The Field Guide to Bigfoot, Yeti, and Other Mystery Primates Worldwide. Avon Books, New York.

Dixon, D. 1981. After Man: A Zoology of the Future. Granada, London.

Gans, C. 1969. Amphisbaenians – reptiles specialized for a burrowing existence. Endeavour 28, 146-151.

- . 1974. Biomechanics: An Approach to Vertebrate Biology. J. B. Lippincott Company, Philadelphia, Toronto.

- . 1978. The characteristics and affinities of the Amphisbaenia. Transactions of the Zoological Society of London 34, 347-416.

Hurum, J. H., Luo, Z.-X. & Kielan-Jaworowska, Z. 2006. Were mammals originally venomous? Acta Palaeontologica Polonica 51, 1-11.

Kearney, M. 2002. Appendicular skeleton in amphisbaenians (Reptilia: Squamata). Copeia 2002, 719-738.

Kesteven, H. L. 1957. Notes on the skull and cephalic muscles of Amphisbaenia. Proceedings of the Linnean Society of New South Wales 82, 109-116.

Koch, A. & Wyman, J. 2007. Limblessness in amniotes shared by worm-lizards and basal mammals: the data from ribosomal ISBN and ILN genes. Yay Genes, Today! 1, 20-31.

Luna, L. & Patterson, B. D. 2003. A remarkable new mouse (Muridae: Sigmodontinae) from southeastern Peru: with comments on the affinities of Rhagomys rufescens (Thomas, 1886). Fieldiana: Zoology, New Series 101, 1-24.

Olson, S. L. 2002. Review of: New Perspectives on the Origin and Early Evolution of Birds. Auk 119, 1202-1204.

Papenfuss, T. J. (1982). The ecology and systematics of the amphisbaenian genus Bipes Occasional Papers of the California Academy of Sciences, 136, 1-42

Ratsarse, M. 2006. OOA as applied to the vexing problem of mammal baramins. Creation Science ‘Journal’ 3, 2-3.

Remor, S. A. 1979. Why Dimetrodon is just way cooler than an amphisbaenian as a mammal ancestor. Big Science 101, 7-10.

- . 1980. Amphisbaenia: you blow! Great Plain Science Musings 75, 556-566.

Shuker, K. P. N. 2003. The Beasts That Hide From Man. Paraview Press, New York.

Stümpke, H. 1967. The Snouters: Form and Life of the Rhinogrades. The Natural History Press, Garden City, New York.

Wever, E. G. & Gans, C. 1973. The ear in Amphisbaenia (Reptilia); further anatomical observations. Journal of Zoology 171, 189-206.

Zangerl, R. 1945. Contributions to the osteology of the post-cranial skeleton of the Amphisbaenidae. American Midland Naturalist 33, 764-780.

The 'London specimen' of Archaeopteryx (in this case, a cast), now the holotype of A. lithographica. Image by H. Raab, from wikipedia.

Today I want to talk about something completely different. During the 1980s astronomer Sir Fred Hoyle proposed (with his colleagues Chandra Wickramasinghe, Lee Spetner and R. S. Watkins) that the London Archaeopteryx specimen [shown here: image by H. Raab] was a forgery, made by pressing chicken feathers into plaster laid about the skeleton of the small predatory dinosaur Compsognathus.

This much is well known; also well known is Hoyle et al.’s claim that the specimen was originally acquired by Richard Owen “as a known fraud, with the intent of trapping Darwin and Huxley into claiming it in support of the evolutionary theory” (Hoyle & Wickramasinghe 1986, p. 112). As Gould (1987) argued, Hoyle and Wickramasinghe’s idea that Owen was a creationist and that it would have been to his advantage to discredit Archaeopteryx is a gross and thoroughly naïve misunderstanding of just about every aspect of Victorian palaeontology and sociopolitics, let alone Owen’s personal motivations and philosophy. Hoyle and Wickramasinghe seemed not to credit or even realise how Owen sunk time, effort, work and money into the reality of Archaeopteryx, nor what impact the unveiling of involvement in a hypothetical forgery would have on Owen’s personal reputation, or on that of the British Museum.

Perhaps not as well known as it should be is that Hoyle et al.’s ideas were published in the British Journal of Photography. I have no wish to cast aspersions, or to say anything negative about that venue, but it certainly cannot be considered a normal outlet for scientific results and I somehow doubt that it meets (or met) the standards of peer-review or any conditions usual for scientific results. The bulk of what Hoyle and Wickramasinghe had to say went into a book, Archaeopteryx The Primordial Bird: A Case of Fossil Forgery (Hoyle & Wickramasinghe 1986). They explained how Lee Spetner (a physicist based in Rehovot, Israel) had initially proposed the forgery idea in correspondence, and how R. S. Watkins both photographed the specimen on behalf of the group, and suggested British Journal of Photography as a publication venue.

This rather unfair cartoon of Fred Hoyle is credited to 'Paul S' and appeared in Moreton (1988).

We know from published articles, footnotes in papers, and Hoyle and Wickramasinghe’s book that a few ‘formal’ meetings were held between Hoyle et al. and research and curation staff at London’s Natural History Museum (at the time known as the British Museum (Natural History)). I wonder how friendly these meetings were (minutes do survive). Numerous to-ings and fro-ings in the scientific press were published, most appearing rightly critical of the Hoyle et al. idea, and some being very critical of Hoyle himself (e.g., Charig 1985, Williams 1985, Benton 1987, Connor 1987, Moreton 1988).

The museum’s Alan Charig and colleagues published an authoritative response in Science* (Charig et al. 1986). Stating at the outset that they “reject this forgery hypothesis unequivocally” (p. 623), they refuted in detail all of the evidence alleged to support the claim. They pointed to the many methodological and philosophical problems inherent to it, and showed time and again how the supposedly suspicious details raised by Hoyle and Wickramasinghe couldn’t be taken as evidence of forgery, but were instead genuine geological features or artefacts resulting from decades of preparation (Charig et al. 1986).

* Why Science and not Nature? Hmm.

The ‘Archaeopteryx is a forgery’ idea remains popular among creationists and others on the lunatic fringe, but even they fail to appreciate the bizarre logic behind Hoyle and Wickramasinghe’s argument. As explained in their book, Hoyle & Wickramasinghe (1986) sought to show that Archaeopteryx was a forgery because it proved an obstacle to their idea that dinosaurs and other Mesozoic vertebrates had been transmogrified by viral and/or bacterial storms that had rained down on the Cretaceous world from outer space, grafting new genetic information onto the animals and causing them to change into the birds and mammals of the Cenozoic (Hoyle & Wickramasinghe 1986). In other words, they were seriously proposing this sort of thing…

[‘Transmogrification event caused by incorporation of alien bacteria’ should really say ‘Transmogrification event caused by incorporation of alien VIRUSES’. My bad, sorry.]

As a pre-Cenozoic bird, Archaeopteryx did not fit and had to be explained away (Hoyle and Wickramasinghe were generally unaware of other pre-Cenozoic birds, and ignored Mesozoic mammals entirely). Had this entertaining scenario been presented to the public at the same time as the ‘Archaeopteryx is a forgery’ claim, it is doubtful if it would have been taken as seriously as it was in some circles.

Part of this text is adapted from the biography of Alan Charig that Richard Moody and I published a few years ago (Moody & Naish 2010). I’ve been saying for ages that I wanted to bring the true weirdness of Hoyle and Wickramasinghe’s evolutionary model to the fore, and what better way than to do it than with cartoons.

PS – this article was not intended as a full critique or evaluation of the ‘forged Archaeopteryx’ claim. I don’t feel the need to address this because it’s a total non-starter: there are now more than ten archaeopterygid specimens (most of which have nothing whatsoever to do with Victorian sociopolitics), and – even without the feathers – the skeletal anatomy of Archaeopteryx demonstrates an affinity with Cenozoic birds (Hoyle et al. very wrongly assumed that Archaeopteryx was skeletally identical to Compsognathus), as do a huge number of other Mesozoic maniraptoran fossils (many of which are feathered).

For previous Tet Zoo articles relevant to Archaeopteryx and some of the other issues touched on here, see…

• Tet Zoo picture of the day # 24 (on archaeopterygids)
• Long and Schouten’s Feathered Dinosaurs, a review
• Cyril Walker
• Gary Kaiser’s The Inner Bird: Anatomy and Evolution
• Dinosaurs and Other Extinct Saurians: A Historical Perspective, the book
• Luis Chiappe’s Glorified Dinosaurs: The Origin and Early Evolution of Birds

Refs – -

Benton, M. J. 1987. Why Archaeopteryx is not a fake but suffers from too much publicity. Geology Today 3, 118-121.

Charig, A. J. 1985. Is Archaeopteryx a forgery? Biologist 32 (3), 122-123.

CHARIG, A., GREENAWAY, F., MILNER, A., WALKER, C., & WHYBROW, P. (1986). Archaeopteryx Is Not a Forgery Science, 232 (4750), 622-626 DOI: 10.1126/science.232.4750.622

Connor, S. 1987. Riddle of missing rock resurrects Archaeopteryx controversy. New Scientist 115 (1573), 27.

Gould, S. J. 1987. The fossil fraud that never was. New Scientist 113 (1553), 32-36.

Hoyle, F. & Wickramasinghe, C. 1986. Archaeopteryx The Primordial Bird: A Case of Fossil Forgery. Christopher Davies, Swansea.

Moody, R. T. J. & Naish, D. 2010. Alan Jack Charig (1927-1997): an overview of his academic accomplishments and role in the world of fossil reptile research. In Moody, R. T. J., Buffetaut, E., Naish, D. & Martill, D. M. (eds) Dinosaurs and Other Extinct Saurians: A Historical Perspective. Geological Society, London, Special Publications 343, pp. 89-109.

Moreton, S. 1988. Is Archaeopteryx the only old, cracked fossil? Geology Today 4 (3), 83-84.

Williams, N. 1985. Fraudulent feathers? Nature 314, 210.

Storm-petrels are small tubenoses, found in oceans worldwide and famous for the ‘surface pattering’ behaviour they practise while foraging at the sea surface. Some storm-petrels (the mostly southern Oceanites, Garrodia, Pelagadroma, Fregatta and Nesofregatta, grouped together in Oceanitinae) have long legs and proportionally short, rounded wings. Others (the mostly northern taxa grouped together in Hydrobatinae: Hydrobates, Halocyptena and [the paraphyletic] Oceanodroma) have short legs and longer, more pointed wings. The two groups may not in fact form a clade: some studies find hydrobatines closer to albatrosses than to oceanitines (Penhallurick & Wink 2004). Anyway, one of the things I really like about Lockley’s Flight of the Storm Petrel is the artwork – in particular, Noel W. Cusa’s brilliant, black-and-white-drawings. I previously featured one of them here. In the interests of bringing them to further attention, I’m sharing some (but certainly not all) of them here. They’re interesting for several reasons.

Many of Cusa’s drawings in Flight of the Storm Petrel show storm-petrels being killed, eaten or dismembered by predators. In fact, as you’re about to see, there appears to be an unusually high number of pictures where storm-petrels are depicted as unfortunate victims, not as heroes or victors. There are a few ways to interpret this. Maybe Cusa was making the point that life as a storm-petrel is dangerous and nasty, and that they live a risky life where nefarious predators lurk around every corner: owls, raptors, gulls, skuas… This is only partly true – many tubenoses (storm-petrels included) are long-lived for their size and deaths at the hands or bills of predators are comparatively rare (major caveat: feral cats and rats have devastating impacts on many breeding tubenoses, and humans are doing their best to rid the world of albatrosses and other tubenoses via plastic pollution and the fishing industry). Another possible interpretation of Cusa’s many drawings showing storm-petrel death and dismemberment is that he was illustrating especially interesting vignettes of storm-petrel natural history – after all, there are only so many times you can illustrate seabirds foraging over the surface of the open ocean. Another interpretation is that he didn’t like storm-petrels much. I kid, of course.

In the above drawing, a British or European storm-petrel Hydrobates pelagicus is being beheaded by a Little owl Athene noctua. This scene is inspired by descriptions of this exact interaction, witnessed occurring on Skokholm (off Pembrokeshire, Wales). Lockley (1983) writes that “one day in July 1936 we found a [Little owl] nest … in a rabbit burrow, with two tiny owl chicks and a larder of nearly 200 corpses of storm petrels, the majority with only the head removed!” (p. 62). Lockley and colleagues actually took the step of removing these owls and of “shooting or banishing” any additional ones from Skokholm, “preferring to protect our petrels rather than encourage this acclimatised importation from Europe” (p. 62). In 1954 the step was taken to remove Little owls from Skokholm entirely (Hayden 2004), and they’ve since been blamed for a decline in storm-petrels on Skomer. Little owls are native to much of continental Europe but they were introduced to the British Isles during the 1840s, and from here they colonised Skokholm and Skomer during the 1920s. Moving on…

This illustration (above) shows Matsudaira’s storm-petrel Oceanodroma matsudairae. The one thing that books always say about this hydrobatine species is that it nests on volcanic islands south of Japan, and you’ll note the very obvious Japanese references (art-memes?) in Cusa’s drawing – I mean, come on, I don’t need to even mention them, do I? Matsudaira’s storm-petrel migrates to the Indian Ocean outside of the breeding season, moving all the way west to the coast of Somalia and Kenya. Observations suggest that they migrate through the channel that separates Australia from Indonesia (Harrison 1983). Oceanodroma as conventionally imagined is paraphyletic with respect to the other hydrobatine taxa Hydrobates and Halocyptena. One solution is to lump them all into the same genus (Hydrobates Boie, 1822 has priority); another is to propose a revised taxonomy – and this is at least consistent with the apparently substantial divergence dates inferred from amino acid distance data. According to the revised taxonomy, Matsudaira’s storm-petrel belongs to the same lineage as the Least storm-petrel Halocyptena microsoma, and thus becomes another member of Halocyptena (Penhallurick & Wink 2004).

Here’s more art that might be deemed offensive to storm-petrels. In the above illustration, a Peregrine Falco peregrinus stands over a White-faced or Frigate storm-petrel Pelagodroma marina. If you know Cusa’s artwork, you might recognise a similar-looking peregrine featured elsewhere, though this time in colour. I don’t know if the illustration is based on a specific observation, but Lockley (1983) does refer to the use of ships as hunting platforms for peregrines that will “strike and kill small birds, including storm petrels, fluttering and feeding in the ship’s wake” (p. 154). Some peregrine populations really can be considered specialist predators of seabirds. Incidentally, we also now know that some Gyrfalcons F. rusticolus live at sea for months, resting on sea-ice and subsisting entirely on auks and other seabirds.

Below, skuas (one of the Catharacta species, perhaps C. antarctica) are the bad guys. Skuas are awesome predators – more awesome than you might think, since they can and do kill seabirds similar in size to themselves – but they tend not to bother with storm-petrels since they’re too nimble in flight. Lockley (1983) says that a dead or disabled storm-petrel, however, will of course be taken by a skua. The storm-petrel shown here is Wilson’s storm-petrel Oceanites oceanicus.

But don’t get me wrong, there are about 60 of Cusa’s drawings in Lockley (1983), and the ones featuring predation, dismemberment etc. are actually very much in the minority. We end with a flock of Dove prions Pachyptila desolata flying over a stormy ocean (below)…

I hope you like Cusa’a drawings as much as I do and that you enjoyed my showcasing of them here.

For previous Tet Zoo articles on tubenosed seabirds, see…

• A symbiotic relationship between sunfish and… albatrosses? Say what?
• Because the world belongs to petrels (petrels part I)
• Living the pelagic life: of oil, enemies, giant eggs and telomeres (petrels part II)
• Petrels: some form-function ‘rules’, and pattern and pigmentation (petrels part III)

And for articles about other kinds of seabirds, see…

• To the Sahara in quest of dinosaurs (living and extinct) (includes discussion of gulls and terns)
• Gannets, most awesome of seabirds
• Happy 2009, from the gulls
• Fascinated by boobies
• Mysterious channels of Alca torda
• Kleptoparasitism at Westbury Manor
• Gary Kaiser’s The Inner Bird: Anatomy and Evolution
• When bivalves attack (or: bivalves vs birds, the battle continues)

Refs – -

Harrison, P. 1988. Seabirds: an Identification Guide. Houghton Mifflin Company, Boston.

Hayden, J. E. 2004. The diet of the little owl on Skomer 1998-2003. CCW Contract Science Report 673.

Lockley, R. M. 1983. Flight of the Storm Petrel. David & Charles, Newton Abbott & London.

Penhallurick, J., & Wink, M. (2004). Analysis of the taxonomy and nomenclature of the Procellariiformes based on complete nucleotide sequences of the mitochondrial cytochrome b gene Emu, 104, 125-147

Time for more petrels. This article is another introduction, this time to generalities of behaviour and form-function. The previous petrel articles are here and here, and see the list of links below as well.

Several distinct foraging styles are employed by petrels and they’re more diverse in feeding behaviour than most accounts imply. The majority of species are flexible feeders, their diet mostly being determined by the abundance and availability of fish, crustacean and cephalopod prey species.

Aquaflying shearwaters in pursuit of fish. A still from Blue Planet, BBC (c).

Many feed by alighting on the surface to feed on plankton, fish, squid or floating carrion. Surface seizing of prey is fairly common (where the birds reach down to grab prey from the sea surface while in flight), but some species practise aerial pursuit of other seabirds (that is, they’re aerial pirates or kleptoparasites: more on this in a later article). Others dive into the water in pursuit of prey, practising either short, surface dives from a few metres up (a technique termed surface plunging), or longer, more extensive dives in which they pursue prey well beneath the surface (a technique termed pursuit diving). Indeed, some species (some shearwaters and possibly some others) even ‘fly’ down in the water to depths of 10 or even 20 metres and are capable aquaflyers (Habib 2010). I’ll be coming back to aquaflying later on as well. It’s a fairly well known bit of behaviour nowadays, having famously been featured in the 2001 BBC series Blue Planet.

Prions move in a peculiar way across the sea surface, paddling with their feet, holding their bodies lightly against the water, their wings out to the side, and their heads disappearing and reappearing into the water as they scoop up plankton with the bill (Nelson 1980). This form of feeding has been termed hydroplaning.

Ashmole's (1971) classic diagram of seabird feeding techniques (illustrator: Jon Ahlquist). Note the hydroplaning prion and 'pursuit plunging' shearwater. Today we know that a few shearwaters should be included within the 'pursuit diving with wings' (= aquaflying) category.

In one of the neatest papers ever published in ornithology (that’s just my opinion of course), Spear & Ainley (1998) showed how the morphological characters of petrels correlated with their diet and foraging style. Tropical species, they showed, tended to have longer and deeper bills, longer tails, and longer wings with greater surface areas than polar species (Spear & Ainley 1998). Presumably these differences have arisen because tropical species have to cover much larger expanses of ocean in order to find food; furthermore, tropical prey items (like flying fish and flying squid) are typically fast-moving, agile and capable of flight, whereas polar prey are less mobile and tend to be concentrated near the water surface. The presence of longer bills in tropical species might relate to thermoregulatory constraints (it’s better to have small appendages in cold climates).

A polar specialist: the all-white Snow petrel (Pagodroma nivea). Photo by Samuel Blanc, from wikipedia.

Polar species tend to have a relatively low overlap in morphological characters compared to their tropical relatives, though it’s not immediately clear why this is so given that more resources are available to polar seabirds than to tropical ones. [Adjacent photo of Snow petrel Pagodroma nivea by Samuel Blanc © http://www.sblanc.com/]  Possibly, polar petrels have evolved alongside a greater diversity of other seabird groups than have tropical ones, and as a consequence the number of niches available to polar petrels may have been relatively low (Spear & Ainley 1998). On the other hand, polar environments may provide seabirds with more niches, and hence more opportunities for specialisation. In contrast, tropical regions are comparatively sparse over large areas: consequently, seabirds that evolve here may have to be generalists.

Mostly blacks, whites and greys, but browns as well: patterns and pigmentation

Tahiti petrel (Pseudobulweria rostrata), image by Aviceda, from wikipedia.

Petrels are mostly patterned in greys, white and blacks. White undersides and dark dorsal surfaces are common and dark wing-tip markings, pale rump bands and facial masks and caps are seen in various species [Tahiti petrel Pseudobulweria rostrata shown here; image by Aviceda]. Some species – gadfly-petrels and prions in particular – have complex pigmentation pattern. The Snow petrel has entirely white plumage [see image above].

Bretagnolle's (1993) diagram of plumage variation seen within tubenoses. This paper is a must-read if you're properly interested in seabird colouration and its adaptive significance.

It may be somewhat surprising to those who know little about seabirds and expect all species to be patterned in grey, white and black to learn that quite a few petrels are solidly black or dark brown. There are black-brown species of gadfly-petrels, shearwaters and Procellaria petrels, and the Bulweria and Pseudobulweria species are wholly dark as well. Some populations of Macronectes (the giant petrels) are wholly dark brown and the Northern fulmar Fulmarus glacialis – generally familiar as a mostly white bird with a grey back and wings – also includes a mostly brown form in parts of its range.

Is there any consistency to this diversity of colours and patterns? That’s hard to answer. It might be ideal, for purposes of camouflage from prey and predators, for a seabird to be pale ventrally and greyish or bluish dorsally. However, numerous sometimes incompatible selection pressures mean that organisms do not necessarily evolve in an ‘optimal’ direction when it comes to pigmentation (Endler 1978, Burtt 1981). Seabird colouration seemingly incorporates sexually selected display traits, so there are evolutionary pressures to be showy and distinctive.

Masked shrike, by Claudia Becher, from wikipedia.

Plumage colours may also reflect the need for social cohesion (species that feed in groups need to be able to see conspecifics from a long way off), they may play a role in thermoregulation, and birds may sometimes use feather colour to reflect or absorb sunlight. Dark masks, for example, may help reduce glare and enable predators to see better in strongly lit environments. That last idea is mentioned in the ornithological literature with respect to the dark masks of peregrines and other raptors, and it’s even been suggested that raccoons and other mammals might benefit from their masks in the same way. A recent experimental study on shrikes provides support for the hypothesis, since Masked shrikes Lanius nubicus with dark facial masks were more effective hunters (being far better able to hunt facing the sun) than ones with artificial pale masks (Yosef et al. 2012). [Masked shrike image by Claudia Becher.]

Differential erosion of feathers according to melanin content. At top: the unpigmented primary tips of a Herring gull (Larus argentatus) are worn away while the dark remainder persists. At bottom: brown feather segments in the Snow bunting (Plectrophenax nivalis) are worn away during the year, meaning that these feathers appear to change from brown to black. From Strauch (1991).

The role of melanin in protecting feathers from abrasion and UV damage may also mean that feathers, or their tips, are dark for reasons unrelated to ecology or behaviour. Some petrel (and other seabird) species may also be coloured to mimic others (more on this later). Feeding style and prey type may also have some control over pigmentation, and diurnal and nocturnal species are obviously going to be under different pressures when it comes to camouflage. And do the selection pressures that act on nesting (often burrow-dwelling) petrels have as much, or even more, influence on their pigmentation than the pressures they experience while foraging and avoiding predators at sea? Remember that some petrels may be visiting their nest burrows for more than seven months out of the year. Species are also sometimes patterned or coloured the way they are because that’s what their ancestors were like – that is, their patterns and colours aren’t necessarily adaptive. Some biologists argue that exaptation is a very dodgy idea since it fails to account for (often under-studied) adaptational explanations for structures and behaviours. There may be some truth in this, but a historical, phylogenetic perspective demonstrates that exaptation must have occurred in many lineages.

Some of the graphs from Bretagnolle (1993), here showing the results of correspondence analyses between colouration, diet and feeding style in petrels and other tubenoses.

Bretagnolle (1993) looked specifically at the adaptive significance of tubenose pigmentation and concluded that no one factor had a dominant effect. However, foraging style and group size seemed to be the most important controlling factors. Small species that feed in groups (like prions and some gadfly-petrels) seemingly tend to be cryptically coloured in whites and greys, but dark upperparts may also aid concealment against the sea surface when aerial predators (like larger tubenoses and skuas) need to be avoided. Prominent countershading (like that present in many shearwaters) was suggested to correlate with the underwater pursuit of fish and the avoidance of aquatic predators.

These proposed explanations are somewhat speculative and, as I said above, the incompatibility of various of the selection pressures acting on pigmentation may mean that some (or many, or most) of these birds don’t really possess optimal coloration for their lifestyle. Bretagnolle (1993) also suggested that tubenoses might differ from the majority of other seabird groups in pigmentation since they tend to practise continuous feeding (that is, they forage constantly across the whole duration of their time at sea, rather than commuting to an area of prey abundance). In other words, they might not follow the same rules as other seabird groups.

And, yes, more on petrels to come. For previous Tet Zoo articles on seabirds, see…

• To the Sahara in quest of dinosaurs (living and extinct) (includes discussion of gulls and terns)
• Gannets, most awesome of seabirds
• Happy 2009, from the gulls
• Fascinated by boobies
• Mysterious channels of Alca torda
• Kleptoparasitism at Westbury Manor
• Gary Kaiser’s The Inner Bird: Anatomy and Evolution
• When bivalves attack (or: bivalves vs birds, the battle continues)
• A symbiotic relationship between sunfish and… albatrosses? Say what?
• Because the world belongs to petrels (petrels part I)
• Living the pelagic life: of oil, enemies, giant eggs and telomeres (petrels part II)

Refs – -

Ashmole, N. P. 1971. Avian Biology Volume 1. Academic Press, London.

Bretagnolle, V. (1993). Adaptive Significance of Seabird Coloration: The Case of Procellariiforms The American Naturalist, 142 (1) DOI: 10.1086/285532

Burtt, E. H. 1981. The adaptiveness of animal colors. BioScience 31, 81-102.

Endler, J. A. 1978. A predator’s view of animal color patterns. In Hecht, M. K., Steere, W. C. & Wallace, B. (eds) Evolutionary Biology, Volume 11. Plenum, New York, pp. 319-364.

Habib, M. 2010. The structural mechanics and evolution of aquaflying birds. Biological Journal of the Linnean Society 99, 687-698.

Nelson, B. 1980. Seabird: Their Biology and Ecology. Hamlyn, London.

Spear, L. B. & Ainley, D. G. 1998. Morphological differences relative to ecological segregation in petrels (family: Procellariidae) of the Southern Ocean and tropical Pacific. The Auk 115, 1017-1033.

Strauch, 1991. Feathers. In Brooke, M. & Birkhead, T. (eds) The Cambridge Encyclopedia of Ornithology. Cambridge University Press (Cambridge), pp. 20-26.

Yosef, R., Zduniak, P. & Tryjanowski, P. 2012. Unmasking Zorro: functional importance of the facial mask in the Masked Shrike (Lanius nubicus). Behavioral Ecology doi: 10.1093/beheco/ars005

Cape petrel (Daption capense) - one of my favourite petrels. Photo by JJ Harrison, from wikipedia.

In the previous article we looked very briefly at a few basic aspects of petrel diversity, focusing in particular on their distinctive, tube-nosed bills (though, note: not unique to petrels but present across Procellariiformes). Here, I want to look at these birds in a bit more depth. We focus here on oil production and storage and also on reproduction, longevity, mortality and some related aspects of natural history. [Adjacent photo by JJ Harrison. Annoying fuzziness of these images due to scrunching - I have to reduce them as much as possible via the ‘save for web’ function.]

Oil, squirting oil, using oil, and why contain oil in the first place?

It used to be thought that the petrel nasal tube acted as a nozzle allowing the birds to squirt stomach oil as a defensive measure. In fact, the birds squirt oil defensively through the mouth (though, when handled, they do sometimes drip oil from the nasal tube). This oil-squirting behaviour is most famously practised by fulmars (Fulmarus) but other members of the petrel group Fulmarini (all of which nest out in the open, rather than in burrows like many other petrels) do it as well. [Fulmar image below by T. Müller.]

Northern fulmar pair, here not squirting oil. Photo by T. Müller, from wikipedia.

Fulmars in fact are able to squirt with quite some accuracy up to a distance of a metre or two, and young fulmar chicks seem to squirt at just about anything that comes within reach, including their parents (the chicks later learn not to do this). The oil mats feathers together and destroys their water-proofing abilities, so soiled birds generally die from chilling and/or drowning (fulmars seem ok when soiled by other fulmars, however, being able to preen and wash the oil out). There are cases where sea-eagles have died after being squirted by fulmars (Dennis 1970) and about another 20 species are known to have been killed as well, including herons, gulls, owls, falcons, crows and a few unfortunate small passerines (Broad 1974, Booth 1976, Warham 1977). A fulmar kept in captivity with gulls and auks managed to kill five of them by soaking them with oil. There aren’t many images of oil-spitting petrels out there, but I managed to find the one below. It’s from anneleeuw’s flickr site.

Brilliant image of oil-spitting fulmar by anneleeuw (c).

Where does this oil come from? A peculiarity of tubenose biology is that adults and young contain large quantities of oil in the proventriculus (e.g., Weimerskirch & Cherel 1998, Cherel et al. 2002).

This oil varies from clear to deep reddish-brown and solidifies to form a wax when cool. It has been used medicinally by humans as well as in the lubrication of machinery and of course in providing illumination. Oil from Short-tailed shearwaters Puffinus tenuirostris has been used in the manufacture of sun-tan lotion and as a coat shine for horses (Warham 1977).

It used to be thought that the birds manufactured the oil themselves. In fact, its chemical composition shows that they obtain it by rupturing their animal prey, differentially digesting the proteins and lipids, and concentrating huge quantities of energy in oil form (Warham 1977). This serves as an energy reserve for adults on long foraging trips but also allows them to provision their chicks with a condensed, lightweight (specific gravity 0.88) and highly nutritious substance in between periods of absence. Adult petrels tend to ‘overstock’ their chicks with oil, perhaps as a precaution against unexpectedly long trips away from the nest, and the oil-squirting abilities of chicks show that they contain a seemingly large amount of oil at any one time. They couldn’t do this if they were only being given the bare minimum to stay alive. Adults actually need to store the energy they gain on their long foraging trips in order that they can afford the many short trips required to feed the chick (Weimerskirch et al. 2003). In other words, the long foraging trips seem to allow parents to recoup their energetic losses.

Pelagic life

As a generalisation, petrels are stiff-winged soaring birds with low wing loadings. Wingspans range from about 60 cm in some of the prions (Pachyptila) to 2 m in the awesome, vulturine giant petrels (Macronectes). Petrels are pelagic, covering substantial distances across the open ocean. Some species are known to travel distances of 15,000 km on single foraging trips. This particular factoid concerns the Short-tailed shearwaters Puffinus tenuirostris that breed on the southern coast of Australia yet were shown by satellite tracking to forage at the limits of Antarctic shelf-ice (Klomp & Schultz 2000).

Wedge-tailed shearwater. Photo by Bryan Harry, from wikipedia.

After breeding, adults disperse. Some species (like prions, some gadfly-petrels and shearwaters like the Wedge-tailed shearwater Pu. pacificus [shown here] in the tropic Pacific and Indian oceans and Townsend’s shearwater Pu. auricularis in the eastern Pacific) are sedentary and only move to adjacent waters, while other range far and wide, even travelling to other oceans. Most Manx shearwaters Pu. puffinus, for example, breed in the north-east Atlantic (some breed off Cape Cod in the west as well*) and then disperse to the south-west before returning north, but it seems that some travel round Cape Horn and end up going north on the ‘wrong’ side of the Americas (Harrison 1988).

* This was only discovered in 1973 (Bierregaard et al. 1975). There are indications that Manx shearwaters bred off the east coast of North America in prehistoric and historic times but became extirpated relatively recently.

Similarly, Cory’s shearwater Calonectris diomedea breeds in the Mediterranean and then disperses right across the Atlantic with many ending up around South Africa and Natal. However, some go round Africa into the western Indian Ocean. These normally (it is thought) move back into the Atlantic before heading north, but the presence of individuals in the northern Red Sea suggests that some of these birds get back into the Mediterranean by migrating north along the east side of the Africa, not the west side.

Petrel predators

Great black-backed gull (Larus marinus) about to start dismembering Manx shearwater. Illustration by Noel Cusa, from Ronald Lockley's Flight of the Storm Petrel. More on Cusa's artwork at a later date.

We saw a moment ago that petrel chicks on nests need to defend themselves from raptors and also from predatory gulls and skuas. Asio owls, caracaras and Buteo hawks are also documented predators of petrels. On Codfish Island, New Zealand, Weka Gallirallus australis were taking such a major toll on Mottled petrels Pterodroma inexpectata and Cook’s petrel Pt. cooki that the decision was eventually made to remove them (Imber et al. 2003).

More unusual predators includes crabs, snakes, tuatara and skinks, all of which are on record as eating petrel eggs and/or nestlings. Foxes predate on some petrel populations and stoats introduced to New Zealand are known to have a significant impact on breeding Hutton’s shearwaters Pu. huttoni (Cuthbert & Davis 2002). Red deer Cervus elephus on the Scottish island of Rhum famously took to biting the heads, wings and legs off Manx shearwater chicks, apparently because of calcium deficiency (Furness 1988). Petrels are not great movers on the ground – their legs are weak, they mostly move with a shuffling gait, and they generally need to climb inclined surfaces (slopes, cliffs or tree trunks) for a takeoff. Needless to say, the altricial chicks are even more helpless, so they can fall prey to just about any animal capable of finding and overpowering or dismembering them.

Domestic cats – introduced to various islands used by nesting petrels – represent a serious threat and seem to have caused massive declines or even the local extinction of some species. To give some idea of how serious cat predation can be, note that about 5000 Cook’s petrels seem to have been killed by cats on a single New Zealand island in a single breeding season (Imber et al. 2003). A colony of Bulwer’s petrel Bulweria bulweria that previously nested on the southwest coast of Gran Canaria was seemingly eradicated by cats (Luzardo et al. 2008). On Marion Island in the southern Indian Ocean, Soft-plumaged petrels Pt. mollis had extremely poor rates of breeding success due to cat predation (just 7.9% of pairs succeeded in raising chicks) while the Common diving-petrels Pelecanoides urinatrix there became extinct some time round about 1952, apparently as a direct result of cat predation. Eradication of cats on Marion Island in 1991 resulted in improved breeding success in some (but not all) of the affected species (Cooper et al. 1995) and cat eradication elsewhere has resulted in petrel recovery (Keitt & Tershy 2003).

This diagram - from Rayner et al. (2007) - shows that, when cats are present (far left), petrel breeding success was higher than when cats were absent (but rats were still present). Needless to say, things only really improved once rats and cats were removed (far right).

However, while eradicating cats might seem like a great move, this isn’t necessarily always the case. People haven’t just introduced cats to islands – they’ve also introduced rats and mice. Freed from cat predation, rats on some islands have experienced so-called ‘mesopredator release’, increasing in number and becoming more significant predators of petrels (Rayner et al. 2007) [Laelaps blogged about this research soon after it was published]. A factor that may complicate attempts to assess the impact of rat predation is that adult petrels may persist for a long time (due to their longevity: read on) even when they are totally failing to raise chicks due to increased rat predation. Conversely, while cat predation may result in greater losses of adults, it may not necessarily have the same devastating impact on nestlings that rat predation does (though… it might). Accordingly, some workers urge that we need to be careful when deciding how to deal with alien predators (Le Corre 2008).

Breeding, brooding, ageing and claims of immortality

From Rahn et al. (1975). Obviously, some petrels produce proportionally large eggs for their body size.

Petrels are long-lived, slow-breeding birds that put a massive amount of effort into raising their chicks. Individuals of some species (the Manx shearwater being the classic example) might be at their nesting grounds from February all the way to September, October or even November. They produce a single, proportionally large egg – somewhere round about 21-25% of the adult’s mass (and thus within the upper range limit for birds)* – and have an incubation period that is round about, and sometimes exceeds, 50 days (Warham 1983). Cases where abandoned, second chicks have been adopted by breeding pairs suggest that parents aren’t able to provision two chick, and furthermore that chicks exhibit high levels of aggression towards one other (Archuby et al. 2010).

"Above is an x-ray of a pregnant kiwi. I think it's pretty clear that the bird can't eat, drink, poop, or even breathe while that monster is in the pipe. It's science." From the pen of Dr Mathew Wedel.

* Caveat # 1: after I said on twitter that some tubenoses exceed kiwi in relative egg mass (in the Little spotted kiwi Apteryx owenii, egg mass can be 22% of body mass), the brilliant Mike Dickison reminded me that allometry needs to be considered. To paraphrase Mike, avian egg mass scales to body mass at about the two-thirds power, so a 200 g bird that produces an egg 25% of body mass is much less remarkable than a 1000 g bird that does likewise. Caveat # 2: while it’s relatively easy to find data on egg mass as a percentage of adult female body mass (Rahn et al. 1975), most birds produce clutches of several or many eggs, and finding clutch size expressed as a percentage of body mass is less easy. Anyway, viewed within the context of both allometry and of total clutch mass, the single eggs of tubenoses and kiwi are not necessarily that remarkable in terms of parental investment, scaling linearly for body size with those of their relatives. It’s the fact that just a single enormous egg is produced that’s so unusual: a risky strategy that relies on low egg/juvenile mortality.

Southern giant petrel with chick. Image by Brocken Inaglory.

Chicks grow slowly. Cory’s shearwater, for example, spend about 90 days in the nest prior to fledging. There are indications that this slow development is related to previously unappreciated peculiarities of tubenose biology. Maternal antibodies, for example, persist for an unusual length of time (20 days or so) in Cory’s shearwater (Garnier et al. 2011).

I said that petrels are generally ‘long-lived’. By this I mean that individuals of some species (like the giant petrels Macronectes) can live for over 50 years (and there are suggestions that some albatrosses can exceed 70 years). [Image above by TheBrockenInaGlory, use licensed under the Creative Commons Attribution-Share Alike 3.0 Unported, 2.5 Generic, 2.0 Generic and 1.0 Generic license. I apologise profusely for not correctly attributing information earlier: this was an unfortunate oversight]. Longevity (though not necessarily of exactly this scale) is not unique to giant species. An individual of Bulwer’s petrel Bulweria bulwerii, for example, was at least 24 years old when last recorded in 1992 on Johnston Atoll in the Pacific.

Some tubenoses (including giant petrels and also storm-petrels) have been at the centre of argument and interest over the role and function of their telomeres. Telomeres are DNA fragments that ‘cap’ the ends of chromosomes, shortening at each cell division event, and also shortening as oxidative stress takes its toll on an animal over time. Older individuals thus have shorter telomeres than younger ones. Sexual differences in telomere length have been reported in some insect, squamate and mammal species (including Homo sapiens).

While the database we have on telomere length is not (so I understand) all that comprehensive, tubenoses are especially interesting because telomere length does not change all that much in adults (Foote et al. 2010). Furthermore, giant petrels were the first birds in which a sexual difference in telomere length was reported, with males have shorter telomeres (Foote et al. 2010). It may or may not be coincidental that giant petrels are unusual among petrels in exhibiting strong sexual dimorphism – perhaps the strongest dimorphism reported in any seabird. What, if any, impact this has on the biology and behaviour of these birds, or whether it skews survival rates between the sexes, is not entirely clear… or, it’s not clear to me, anyway. Do say if you know otherwise.

Leach's storm-petrel. Note: a storm-petrel, NOT a petrel proper. Photo by USFWS, from wikipedia.

Haussmann & Mauck (2008) reported that telomeres appeared to lengthen during growth in Leach’s storm-petrel Oceanodroma leucorhoa. This probably doesn’t mean that storm-petrels are immortal (as is intimated in some popular re-tellings of this research) but it is probably related to the fact that these birds live about four times longer than expected for their size. One explanation for this pattern is that hatchlings are variable in telomere length and that individuals with long telomeres are the ones that make it to old age. In other words, it isn’t that telomere length is really increasing with age; rather, old individuals are the ones that made it to old age because they’re the ones with the long telomeres!

MUCH more on petrels still to come. For previous Tet Zoo articles on seabirds, see…

• To the Sahara in quest of dinosaurs (living and extinct) (includes discussion of gulls and terns)
• Gannets, most awesome of seabirds
• Happy 2009, from the gulls
• Fascinated by boobies
• Mysterious channels of Alca torda
• Kleptoparasitism at Westbury Manor
• Gary Kaiser’s The Inner Bird: Anatomy and Evolution
• When bivalves attack (or: bivalves vs birds, the battle continues)
• A symbiotic relationship between sunfish and… albatrosses? Say what?
• Because the world belongs to petrels (petrels part I)

Refs – -

Archuby, D. I., Coria, N. R., Harrington, A., Fusaro, B., Montalti, D. & Favero, M. 2010. Is it possible for a procellariiform to raise two chicks? A case of chick adoption in southern giant petrels Macronectes giganteus in the South Shetland Islands, Antarctica. Marine Ornithology 38, 125-127.

Bierregaard, R. O., David, A. B., Baird, T. D. & Woodruff, R. E. 1975. First northwestern Atlantic breeding record of the Manx shearwater. The Auk 92, 145-147.

Booth, C. L. 1976. Peregrine and Raven possibly contaminated by Fulmar oil. British Birds 69, 61.

Broad, R. A. 1974. Contamination of birds with fulmar oil. British Birds 67, 297-301.

Cherel, Y., Bocher, P., De Broyer, C. & Hobson, K. A. 2002. Food and feeding ecology of the sympatric thin-billed Pachyptila belcheri and Antarctic P. desolata prions at Iles Kerguelen, Southern Indian Ocean. Marine Ecology Progress Series 228, 263-281.

Cooper, J., Marais, A. v. N., Bloomer, J. P. & Bester, M. N. 1995. A success story: breeding of burrowing petrels (Procellariidae) before and after the eradication of feral cats Felis catus at subantarctic Marion Island. Marine Ornithology 23, 33-37.

Cuthbert, R. & Davis, L.L.S. 2002. The impact of predation by introduced stoats on Hutton’s Shearwaters, New Zealand. Biological Conservation 108, 79-92.

Dennis, R. H. 1970. The oiling of large raptors by fulmars. Scottish Birds 6, 198-199.

Foote, C., Daunt, F., Gonzalez-Solis, J., Nasir, L., Phillips, R., & Monaghan, P. (2010). Individual state and survival prospects: age, sex, and telomere length in a long-lived seabird Behavioral Ecology, 22 (1), 156-161 DOI: 10.1093/beheco/arq178

Furness, R. W. 1988. Predation on ground-nesting seabirds by island populations of red deer Cervus elaphus and sheep Ovis. Journal of Zoology 216, 565-573.

Garnier, R., Ramos, R., Staszewski, V., Militã, T., Lobato, E., González-Solis, J. & Boulinier, T. 2011. Maternal antibody persistence: a neglected life-history trait with implications from albatross conservation to comparative immunology. Proceedings of the Royal Society B doi:10.1098/rspb.2011.2277

Harrison, P. 1988. Seabirds: an Identification Guide. Houghton Mifflin Company, Boston.

Haussmann, M. F & Mauck, R. A. 2008. Telomeres and longevity: testing an evolutionary hypothesis. Molecular Biology and Evolution 25, 220-228.

Imber, M. J., West, J. A. & Cooper, W. J. 2003. Cook’s petrel (Pterodroma cookii): historic distribution, breeding biology and effects of predators. Notornis 50, 221-230.

Keitt, B. S. & Tershy, B. R. 2003. Cat eradication significantly decreases shearwater mortality. Animal Conservation 6, 307-308.

Klomp, N. I. & Schultz, M. A. 2000. Short-tailed shearwaters breeding in Australia forage in Antarctic waters. Marine Ecology Progress Series 194, 307-310.

Le Corre, M. 2008. Cats, rats and seabirds. Nature 451, 134-135.

Luzardo, J., López-Darias, M., Suárez, V., Calabuig, P., García, E. A. & Martín, C. 2008. First breeding population of Bulwer’s Petrel Bulweria bulwerii recorded on Gran Canaria (Canary Islands) – population size and morphometric data. Marine Ornithology 36, 159-162.

Rahn, H., Paganelli, C. V. & Ar, A. 1975. Relation of avian egg weight to body weight. Auk 92, 750-765.

Rayner M. J., Hauber M. E., Imber M. J., Stamp R. K. & Clout M. N. 2007. Spatial heterogeneity of mesopredator release within an oceanic island system. Proceedings of the National Academy of Sciences of the United States of America 104, 20862-20865.

Warham, J. 1977. The incidence, functions and ecological significance of petrel stomach oils. Proceedings of the New Zealand Ecological Society 24, 84-93.

- . 1983. The composition of petrel eggs. Condor 85, 194-199.

Weimerskirch, H., Ancel, A., Calouin, M., Zahariev, A., Spagiari, J., Kersten, M. & Chastel, O., 2003. Foraging efficiency and adjustment of energy expenditure in a pelagic seabird provisioning its chick. Journal of Animal Ecology 72, 500-508.

- . & Cherel, Y. 1998. Feeding ecology of short-tailed shearwaters: breeding in Tasmania and foraging in the Antarctic? Marine Ecology Progress Series 167, 261-274.

Short-tailed shearwater (Puffinus tenuirostris) megaflock, photographed in Unimak Pass, Alaska. Photo by USGS.

Seabirds are fascinating (as I often say: hey, just like all the other tetrapods). To me, they’ve always seemed to be one of those groups that requires a lifetime of immersion and specialisation should you hope to become properly acquainted with them. I just don’t feel that you can get to know them via books and published articles – you have to actually be out there, at sea, in diverse latitudes, armed perpetually with binoculars and a telescope. My chances to become immersed in the ‘seabird experience’ have been few and far between, but I have done my fair share of watching gulls, terns, auks, gannets and shearwaters as and when possible. [Awesome photo above from USGS.]

Seabirds of many species are phenomenally abundant, and data shows that they play major roles as predators, planktivores and scavengers in various marine ecosystems. But as everyone interested in the natural world should know, seabird populations today are beleaguered by such problems as plastic and oil pollution, climate change, human disturbance, alien predators that exploit them on their nesting grounds, declines in sea ice, and depletion of fish and plankton stocks. There’s therefore a rush to better understand their ecology in an effort to minimise the population losses and the general degradation of the marine ecosystems that these birds are part of.

Introducing petrels

Montage showing representatives of the five main petrel clades. Top left: Great shearwater (Puffinus gravis). Top right: Black-capped petrel (Petrodoma hasitata), both images by Patrick Coin. Below, top to bottom: Southern giant petrel (Macronectes giganteus) by Brocken Inaglory; Broad-billed prion (Pachyptila vittata), by Rosemary Tully; Westland Petrel (Procellaria westlandica), by Mark Jobling.

Here I want to talk about one of my favourite seabird groups, the petrels, also known as true petrels or procellariids. [Adjacent photos by Patrick Coin, BrockenInaGlory*, Rosemary Tully and Mark Jobling, all from wikipedia.]

* Licensed under the Creative Commons Attribution-Share Alike 3.0 Unported license.

For people who like the same sort of thing that I do – recently discovered species, species known from just a few (or even single) specimens, species about which just about nothing is known, species with remarkable life histories, and species capable of really surprising bits of behaviour – petrels are where it’s at. Numerous short articles have been published reporting sightings of petrel species outside of their normal ranges and some species have been reported on so few occasions that any additional sightings are worthy of publication. Other species have disappeared and reappeared throughout history. And major contributions have been made by people able to identify and differentiate species at sea and hence report new, distinguishing characters. There are some excellent field guides to the petrels of the world. I especially recommend Tuck & Heinzel (1978) and Harrison (1983).

The main aim here – and, by “here” I mean “over the next several articles” – is to cover petrel diversity (there are about 75 living species, grouped into five main clades), to overview their biology, ecology and behaviour, and also to discuss how certain aspects of petrel behaviour and ecology match with morphology. I tried hard not to get bogged down in tedious discussions of phylogeny and systematics. As usual, however, you need to understand at least a bit about the phylogeny and systematics of the group before you can hope to realistically understand the evolutionary patterns within the group. In the end I’ve tried to combine everything in the hope that it presents a coherent picture of petrel diversity, ecology and form-function correlation. Completing these petrel-themed articles has not been easy – I started writing them in 2009 or earlier but they’ve been on the backburner.

Petrels (Procellariidae) are tubenosed seabirds, or tubenoses: part of the same neornithine clade as albatrosses (Diomedeidae), storm-petrels (Hydrobatidae) and diving-petrels (Pelecanoididae). The way I’ve just listed those groups – I mean, with all four being recognised as distinct taxonomic ‘families’ – represents the way tubenoses have been classified in mainstream 20th (and 21st) century literature. However, this ‘four family system’ does not match the phylogeny of the group as recovered by recent studies. More on this later.

Wilson's storm-petrel (Oceanites oceanicus) doing the 'water walking' thing. Image by Patrick Coin, from wikipedia.

I should note that ‘petrel’ is supposed to be pronounced in similar fashion to the name Peter, since it apparently originated as a diminutive form of that name (and hence started out as ‘peter-el’). The books say that petrels are named after St. Peter since he “walked (somewhat fearfully) on the stormy sea of Galilee at Christ’s invitation” (Lockley 1983, p. 8). The irony here is that the ‘water walking’ behaviour referred to here is practised by storm-petrels, not by petrels proper (storm-petrels don’t really walk on the water; rather, they patter across the surface with their large webbed feet while flying) [Adjacent image, showing this behaviour, by Patrick Coin]. So, petrels – most of which are speedy soarers – are named after the idiosyncratic behaviour practised by storm-petrels.

Totally tubular (external) nostrils

Labelled section of a tubenose compound rhamphotheca, from Hieronymus & Witmer (2010).

Tubenoses are united by such distinctive morphological features as their tubular nostrils and strongly reduced hallux (it consists just of a single phalanx, or is absent altogether). They also share a terminal hook on the bill. The tubenose bill is composed of a series of distinct plates rather than a continuous rhamphothecal covering and hence is referred to as a ‘compound rhamphotheca’. These different bill plates all have names (Coues 1866). The naricorn, latericorn, culminicorn and maxillary or premaxillary unguis (or nail) are all on the upper jaw; the ramicorn and mandibular unguis (or nail) are on the lower.

The persistence of prominent grooves between these plates might be due to their role in helping to drain unwanted saline fluid from the salt glands. As is typical for seabirds, the glands are located in bony depression over the eye sockets, but the fluid that drains from them is discharged through the nostrils and drips away from the end of the bill (Schmidt-Nielsen 1960). If you’re wondering, in the seabirds that have sealed external nostrils (gannets, cormorants etc.), the fluid is discharged via the internal, palatal nostrils and then runs along the dorsal surface of the palate before being released at the bill tip.

Short-tailed shearwater (Puffinus tenuirostris), photo by J. J. Harrison.

In petrels, the nostrils are united in a single, dorsally positioned tube, and this is also the case in diving-petrels and storm-petrels [see adjacent photo of Short-tailed shearwater, photo by J. J. Harrison]. Albatrosses, however, differ in having two tubes, one on either side of the bill. Given that (virtually all) phylogenies nest albatrosses somewhere within the clade that includes all other tubenoses, you might like to wonder what sort of transformation occurred during the evolution of these birds. Did albatrosses have a single, dorsally placed tube ancestrally, or have albatrosses retained a primitive condition while members of the other lineages convergently evolved the single, dorsally-placed condition?

Incidentally, there are no clear indications from the tubenose skull that tubular nostrils – let alone a single, dorsally located tube – would be present in life. In a tubenose skull, the bar located between the long, oval bony nostrils is raised relative to the sides and more anterior parts of the rostrum, but I can’t see that anyone would link this with the presence of a united tube in the absence of other information. As I write this, I have Northern fulmar Fulmarus glacialis skulls right in front of me.

Northern fulmar skulls - would you think from these skulls that a single, dorsally located nasal tube was present in place of paired external nostrils? Note also the depressions for the salt glands and the hooked premaxillary tip.

The nostril tube may help in channelling scent and it’s well known that storm-petrels and true petrels at least are attracted to dimethyl sulphide (DMS) (Nevitt et al. 1995), a compound released by phytoplankton when it’s subjected to grazing by zooplankton. The tubenoses that have so far exhibited high sensitivity to DMS are nocturnal foragers, so it seems logical that they might respond to olfactory cues more than visual ones when foraging (Nevitt et al. 1995). However, in view of their nocturnal habits, it’s not surprising that many petrels have highly acute night-time vision, with a flattish cornea and a lens that does most of the work in focusing light being among several specialisations for improved vision in the dark (Martin 1990).

I just mentioned that the nostril tube might help to channel scent. It has also been suggested that the tubular (laterally placed) nostrils of albatrosses contain pressure receptors that help them negotiate and exploit shifting air masses over waves (Kaiser 2007).

Next: Oil, squirting oil, using oil, and why contain oil in the first place?

For previous Tet Zoo articles on seabirds, see…

• To the Sahara in quest of dinosaurs (living and extinct) (includes discussion of gulls and terns)
• Gannets, most awesome of seabirds
• Happy 2009, from the gulls
• Fascinated by boobies
• Mysterious channels of Alca torda
• Kleptoparasitism at Westbury Manor
• Gary Kaiser’s The Inner Bird: Anatomy and Evolution
• When bivalves attack (or: bivalves vs birds, the battle continues)

Refs – -

Coues, E. 1866. Critical review of the family Procellariidae: Part V; embracing the Diomedeinae and the Halodrominae. With a general supplement. Proceedings of the Academy of Natural Sciences of Philadelphia 18, 172-197.

Harrison, P. 1988. Seabirds: an Identification Guide. Houghton Mifflin Company, Boston.

Hieronymus, T., & Witmer, L. (2010). Homology and Evolution of Avian Compound Rhamphothecae The Auk, 127 (3), 590-604 DOI: 10.1525/auk.2010.09122

Lockley, R. M. 1983. Flight of the Storm Petrel. David & Charles, Newton Abbott & London.

Kaiser, G. W. 2007. The Inner Bird: Anatomy and Evolution. University of British Columbia, Vancouver.

Martin, G. 1990. Designer eyes for seabirds of the night. New Scientist 128 (1741), 46-48.

Nevitt, G. A., Veit, R. R. & Kareiva, P. 1995. Dimethyl sulphide as a foraging cue for Antarctic procellariiform seabirds. Nature 376, 680-682.

Schmidt-Nielsen, K. 1960. The salt-secreting gland of marine birds. Circulation 21, 955-996.

Tuck, G. & Heinzel, H. 1978. A Field Guide to the Seabirds of Britain and the World. Collins, London.

Amazing stuff. Good to see that some marine reptiles have evolved a rather novel way to deal with the problems of discharging waste and procreating (do you know what I’m getting at? Say so in the comments). And – - scaly skin? Thanks to Bruce Schumacher.

In other news, lots of seabirds to come real soon. Just have to finish this Aquatic Ape Hypothesis article (and a bunch of other stuff) first.

Life restoration of Hainosaurus bernardi, by Dmitry Bogdanov.

Long-time readers will know that I’ve been involved in a great long list of failed book projects. Once upon a time I made significant progress on a book about MESOZOIC MARINE REPTILES; alas, it exploded on the launch pad and everybody died. Unable to find time to do anything else (I’m working, when time allows, on a set of articles that I’ve been trying to finish since 2007), I thought I may as well re-post one of the sections of text I completed: that on the mosasaur Hainosaurus. I’ve added citations and a few small updates. [Life restoration above by Dmitry Bogdanov, from wikipedia.]

Hainosaurus is one of the largest and most spectacular of the mosasaurs. Reaching a possible length of 13 m, Hainosaurus was clearly an awesome predator, essentially capable of tackling any other animal it encountered. Direct evidence for its diet comes from the stomach contents of a Belgian specimen that had swallowed at least part of a gigantic turtle, though whether it had eaten the entire turtle or simply ripped the paddles from a floating carcass is unknown.

Holotype skull (IRSNB R23) of Hainosaurus bernardi, photographed at the IRSNB in Brussels.

Compared to those of many other mosasaurs, the teeth that lined the sides of the jaws in Hainosaurus were strongly compressed from side to side and possessed finely serrated keels running the length of the crown (Lindgren 2005). They are thus superficially like those of some predatory sharks, though still with much stouter crowns [see comment below from Mike Everhart]. The teeth on the palate also had finely serrated keels. These features suggest that Hainosaurus was better able to slice through flesh and cut prey into pieces than were its relatives. Wear facets on the sides of its teeth indicate that it was frequently contacting bone when it was biting prey.

French specimen (MNHN 1896-15) suggested to be referable to Hainosaurus but more recently argued to be a species of Tylosaurus. Image by FunkMonk, from wikipedia.

Hainosaurus was originally named by Belgian palaeontologist Louis Dollo in 1885 for a near-complete skeleton discovered in the latest Cretaceous Belgian chalk (the holotype of the species Hainosaurus bernardi). Hainosaurus fossils were later reported from the Pierre Shale of Manitoba in Canada (Nicholls 1988) and additional specimens from Sweden, France, England and Poland were recognised more recently (Bardet 1990, Jagt et al. 2005, Lindgren 2005) [adjacent image by FunkMonk]. English teeth belonging to Hainosaurus had been described as early as 1845 but were not recognised as those of Hainosaurus until 1993 (Lingham-Soliar 1993). The Canadian and French specimens proved controversial and have since been reassigned to Tylosaurus (Lindgren 2005, Bullard & Caldwell 2010) but a new species, H. neumilleri, was named in 2007 from South Dakota (Martin 2007). Hainosaurus thus inhabited the shallow seas of both Europe and North America during the Santonian, Campanian and Maastrichtian ages of the Late Cretaceous.

Another view of the fantastic Hainosaurus holotype skull (IRSNB R23) on display at the IRSNB, Brussels. Click to enlarge.

Several features make Hainosaurus unusual, the most notable of which is that its limb girdles are, proportionally, the smallest of any mosasaur. Hainosaurus was previously thought to possess a particularly long tail with a very high number of vertebrae, and when reconstructed in this way it was estimated to be as much as 15-17 m long (Russell 1967, Lingham-Soliar 1995, 1998). If correct, this would make it the longest mosasaur. However, articulated specimens show that this is incorrect and that a more modest 12-13 m is correct (Lindgren 2005).

In several features Hainosaurus was quite similar to Tylosaurus and as a result most experts have regarded them as close relatives within the mosasaur clade Tylosaurinae. Their paddles are similar: the forefins are elongate and narrow while the hindfins are broader due to a diverging fifth toe that has particularly large bones at its base. Like Tylosaurus, Hainosaurus has a bony ‘prow’ projecting from the tip of its upper jaw. Some scientists have described this as consisting of solid bone and have suggested that it might have functioned as a ramming device used for stunning prey, but others doubt that it was strong enough to allow such activities. Tylosaurus and Hainosaurus are also similar in possessing very elongate, slit-like bony nostril openings, though exactly what this means for the form of the external fleshy nostrils is unknown.

This is one of those brief, hurriedly-published ‘filler’ articles. Some far more substantial stuff to come very soon…

For previous Tet Zoo articles on mosasaurs, see…

• Tongues, venom glands, and the changing face of Goronyosaurus
• Mosasaurs might have used the same microscopic streamlining tricks as sharks and dolphins

Refs – -

Bardet, N. 1990. First report of the genus Hainosaurus (Squamata, Mosasauridae) in France. Comptes Rendus de l’Academie des Sciences, Paris, Serie II 311, 751-756.

Bullard, T. S. & Caldwell, M. W. 2010. Redescription and rediagnosis of the tylosaurine mosasaur Hainosaurus pembinensis Nicholls, 1988, as Tylosaurus pembinensis (Nicholls, 1988). Journal of Vertebrate Paleontology 32, 416-426.

Jagt, J. W. M., Lindgren, J., Machalski, M. & Radwakski, A. 2005. New records of the tylosaurine mosasaur Hainosaurus from the Campanian-Maastrichtian (Late Cretaceous) of central Poland. Netherlands Journal of Geosciences – Geologie en Mijnbouw 84, 303-306.

Lindgren, J. 2005. The first record of Hainosaurus (Reptilia: Mosasauridae) from Sweden. Journal of Paleontology 79, 1157-1165.

Lingham-Soliar, T. 1993. The mosasaur Leiodon bares its teeth. In Sarjeant, W. A. S. (ed) Vertebrate Fossils and the Evolution of Scientific Concepts. Gordon and Breach Publishers, pp. 483-498.

Lingham-Soliar, T. (1995). Anatomy and Functional Morphology of the Largest Marine Reptile Known, Mosasaurus hoffmanni (Mosasauridae, Reptilia) from the Upper Cretaceous, Upper Maastrichtian of the Netherlands Philosophical Transactions of the Royal Society B: Biological Sciences, 347 (1320), 155-172 DOI: 10.1098/rstb.1995.0019

- . 1998. Unusual death of a Cretaceous giant. Lethaia 31, 308-310.

Martin, J. 2007. A North American Hainosaurus (Squamata: Mosasauridae) from the Late Cretaceous of southern South Dakota. The Geological Society of America Special Paper 427, 199-208.

Nicholls, E. L. 1988. The first record of the mosasaur Hainosaurus (Reptilia: Lacertilia) from North America. Canadian Journal of Earth Sciences 25, 1564-1570.

Russell, D. A. 1967. Systematics and morphology of American mosasaurs (Reptilia, Sauria). Peabody Museum of Natural History Yale University, Bulletin 23 1-240.

It’s funny how things work out. We looked recently at a ‘mystery’ whale carcass from Baja California. As explained here, it turned out to be a Risso’s dolphin Grampus griseus. I recently watched a 2009 sci-fi movie called Hunter Prey. Should you wish to know more about it, the wikipedia article is pretty good. Anyway, at a few points in the movie, we see the skeleton of what is evidently a living species of cetacean (and not, as I suppose we’re meant to think, that of some weird alien beast). In best hi-tech fashion, I took a few photos of my TV screen, and the images you see here are the result.

There are quite a few places in the world – mostly coastal deserts – where you can go and see sun-bleached cetacean skeletons, lying in the same places where they were deposited in decades or even centuries past. Hunter Prey includes a reference to the ‘Baja dunes’ in the end credits, and wikipedia says that it was filmed in Mexico. Anyway – your task: identify the species we see in the movie. I include a cropped image of its skull.

Photo by Douglas Eernisse, University of California (Santa Cruz).

Thanks to everyone who had a go at identifying the Baja California whale carcass. Some suggestions were, err, a little far of the mark (fin whale, mesoplodont beaked whale); others were pretty reasonable (pilot whale). So, what do we have? Answer: a toothed cetacean (that is, an odontocete), total length about 3 m, with a large, subtriangular dorsal fin, a blunt head, and a strongly reduced dentition. The photos clearly showed that dentition is absent in the upper jaws, and that the lower jaws have just two or three short, subconical teeth on each side. The carcass is greyish, but it’s difficult to know if this really reflected the colour in life.

One cetacean – and one cetacean alone – matches these listed characters: Risso’s dolphin Grampus griseus (sometimes known as the Grey dolphin or Grey grampus). Well done Cameron McCormick, suferable, vdinets, leecris, Hai-Ren, Dartian, Mark Evans and Phoca, all of whom correctly said Risso’s dolphin. Regarding other possible matches… pilot whales (Globicephala), False killer whales Pseudorca crassidens, Pygmy killer whales Feresa attenuata and Melon-headed whales Peponocephala electra all have both a higher numbers of teeth overall and emergent teeth in the upper jaw, and all are much darker in living condition.

Life-size Risso's dolphin model, on display at the Natural History Museum, London.

So, the carcass is definitely that of a Risso’s dolphin. A few features normally present in life are not obvious on the carcass. These include a distinctive white, anchor-shaped marking on its chest, and a vertical crease on the forehead. The latter feature is highly distinctive and not seen in other blunt-headed delphinids. Phillips et al. (2003) suggested that it allows the dolphin to create a uniquely angled sonar beam. Unusual specimens with emergent teeth in the upper jaws have been reported on occasion (Kruse et al. 1999) but the typical condition is to have two to seven pairs of teeth in the lower jaws and apparently one or two pairs of unerupted upper jaw teeth.

This is a widespread dolphin, found in tropical, subtropical and temperate oceans and seas worldwide. In contrast to several other globally widespread dolphin ‘species’, molecular variation between populations is low, being about 0.2% (LeDuc et al. 1999). This is consistent with the fact that – in contrast to other globally distributed delphinid ‘species’ – there have not yet been recent proposals to split G. griseus into more than one species.

Risso's dolphin photographed off California by Mike Baird.

Risso’s dolphin seems to show a preference for oceanic canyons and continental slope environments and there are indications that it takes advantage of upwelling phenomena and the movement of oceanic fronts up and down continental slopes (Baumgartner 1997, Bearzi et al. 2011) [adjacent photo by Mike Baird]. It often feeds at night. While its ecology and natural history haven’t really been well studied, reliable data show that it’s a cephalopod specialist (Clarke 1986), mostly eating mesopelagic squid and also octopuses. Stomach content data indicates that fish and tunicates are eaten as well, sometimes, in some places. In prey requirements, morphology and habitat choice, Risso’s dolphin is very similar to the Short-finned pilot whale Globicephala macrorhynchus and there are indications that the two avoid competing by segregating (Shane 1995). When they do meet, interactions are aggressive.

More life-size cetacean models at London's NHM, with the Risso's dolphin down at the bottom. Note the pale markings on the underside of the pilot whale at top left - we'll be coming back to this later. Click to enlarge.

Risso’s dolphin is one of the largest delphinids, with some individuals reaching 4 m and 500 kg. Males and females are similar in size and adults of both sexes are always conspicuously marked with numerous white scars across their bodies and tails. Some of these might result from grappling with cephalopods, but the majority are tooth marks made by other individuals. The ubiquity and number of these score marks show that biting or – at least – ‘scoring’ with the mandibular dentition is a commonplace activity in these dolphins, perhaps occurring as part of play and courtship as well as combat. Some individuals are so scarred that they’re white across much of their dorsal surface.

Risso’s dolphin was first recognised scientifically in 1812 when George Cuvier described a skin and skull discovered at Brest in France. He recognised that it represented a new species and named it Delphinus griseus. At about the same time, Cuvier also obtained a description (with drawing) of another distinctive dolphin that stranded near Nice. This report came from a M. Risso, so Cuvier took to referring to the animal as ‘dauphin de Risso’. He named it D. aries, apparently “because the drawing showed that this dolphin had a creased forehead something like the horn boss on a ram’s head” (Watson 1988, p. 254). It’s been near-universally agreed for some decades now that both specimens belonged to the same species, so the vernacular name ‘Risso’s dolphin’ became transferred to D. griseus.

Skull and mandible of Grampus griseus, specimen BMNH SW1938, photo courtesy Colin McHenry (c) NHM. Note that this specimen has an emergent tooth in the upper jaw. Click to enlarge.

In 1828, John Gray decided that D. griseus and a number of other blunt-headed delphinids should be given their own ‘subgenus’, Grampus. This name – once widely used as the vernacular term for various blunt-headed delphinids – seems to derive from a corruption of the French ‘grand poisson’. Gray’s version of Grampus at times included pilot whales, false killer whales and killer whales as well as species later moved to Lagenorhynchus and Cephalorhynchus, and in addition to G. griseus he and others also named the species G. sakamata Gray, 1846, G. richardsoni Gray, 1850 (based on a lower jaw of unknown origin), G. chinensis Gray, 1866 and G. stearnsii Dall, 1873 (also based on a lower jaw). All of these names (and others) are now included in the synonymy of G. griseus (Hershkovitz 1966).

There have been a couple of misguided attempts to come up with new generic names for Grampus. Grayius Scott, 1873 was proposed since Grampus was deemed inappropriate; Iredale and Troughton (1933), similarly, decided that the name Grampus should be applied specifically to Orcinus orca, the Killer whale, and they therefore proposed the new generic name Grampidelphis for Risso’s dolphin. Their arguments were erroneous; Hershkovitz (1961) said “All this and more too painful to recite is sheer fantasy” (p. 549).

Long-finned pilot whale (Globicephala melas) showing characteric features associated with globicephalines (blackness, bulbous melon, absent rostrum). Image from wikipedia.

Risso’s dolphin is very obviously a true dolphin – that is, part of the delphinoid clade Delphinidae. But where might it belong within this large and morphologically diverse group? Despite its blunt head and lack of a long beak or rostrum*, it has – ignoring confusion from before the 20th century – traditionally not been included within the ‘globicephaline’ group often recognised for pilot whales and other ‘blackfish’. Some authors have included killer whales (Orcinus) within Globicephalinae, and have therefore used the name Orcininae for this group (the latter is Orcininae Wagner, 1846; the former Globicephalinae Gray, 1850). Obviously, globicephalines are mostly black [adjacent pilot whale illustration from here]. Risso’s dolphin isn’t, and I think this partly explains why it hasn’t typically been regarded as a member of the group. But it’s said to exhibit several features of palatal sinus anatomy that are shared with Delphinus and kin (Muizon 1988). Until recently, Risso’s dolphin has, therefore, been included within Delphininae, typically being hypothesised to be closer to Delphinus and Stenella than to the ‘lags’ (= lissodelphinines) or to the ‘stenines’.

* The rostrum is not truly absent in Risso’s dolphin, it’s just very short and not clearly demarcated from the rest of the head.

Delphinoid section of Geisler et al.'s (2011) cladogram. Note the position of Grampus.

However, recent molecular studies do not support a delphinine position for Grampus. In the earliest comprehensive analyses of molecular phylogeny within delphinids, LeDuc et al. (1999) proposed several novel, surprising hypotheses, one being that Grampus is not close to the long-beaked dolphins (Delphinus, Stenella and kin*), but is instead a globicephaline, most closely allied with Pseudorca (false killer whale), Feresa (pygmy killer whale), Peponocephala (melon-headed whale) and Globicephala (pilot whales). [Caveat: I’m using the name Globicephalinae here as if it’s branch-based, and thus that any delphinid closer to Globicephala than to Delphinus or Lagenorhynchus is a globicephaline. Some authors seem to use the name as if it’s unique to the ‘blackfish’ clade. Given that there are no explicit phylogenetic definitions anywhere in the literature (so far as I can tell), it’s difficult to know exactly what to do.]

* All of which (Delphinus, Tursiops, Stenella and Lagenodelphis) are potentially ripe for synonymisation according to these authors (LeDuc et al. 1999).

Rough-toothed dolphin (Steno bredanensis), photo by NOAA, from wikipedia. Sure doesn't look like the other globicephalines... Richard Ellis said that it looks like an ichthyosaur.

McGowen et al. (2009) recovered evidence for a similar clade, though Orcaella (snubfin dolphins) was now recovered as the sister-taxon to the others and, surprisingly, Steno (the peculiar rough-toothed dolphin) was positioned in between Orcaella and the remainder recovered as being closer to Grampus and the ‘blackfish’ than was Orcaella (McGowen et al. 2009). Both the long-beaked delphinines and the lissodelphinines were closer to this clade than was Orcinus, the killer whales. Extremely similar results have been reported in other studies (Caballero et al. 2008, Cunha et al. 2011, McGowen 2011, Geisler et al. 2011). In fact we can now say that the position of Grampus within Globicephalinae is well supported and represents the present consensus. Incidentally, Steno was taken away from the globicephalines and put close to the long-beaked delphinines by Cunha et al. (2011).

Time-calibrated cetacean phylogeny, from Cunha et al. (2011). Hopefully you can see the divergence dates for Grampus and other members of the globicephaline lineage.

McGowen et al. (2009) estimated that Grampus diverged from other globicephalines round about 5 million yeas ago (early on in the Pliocene) while Cunha et al. (2011) similarly suggested (based on relaxed molecular clock calibration) a divergence date of 5.5 million years ago. Fossil specimens of Grampus are rare but there’s supposed to be one from the Early Pliocene of Japan and a Pleistocene record from Madagascar.

Some speculations about sinus form, snout shape and pigmentation

The pterygoid sinus patterns of (a) Globicephala, (b) Steno, (c) Lagenorhynchus (sensu lato), (d) Grampus, (e) Tursiops, (f) Stenella, and (g) Delphinus. From Muizon (1988), itself based on the figure in Fraser & Purves (1960). Click to enlarge.

What does this mean for delphinid evolution? It suggests several interesting things. One is that pterygoid sinus morphology – the sole anatomical character complex used to ally Grampus with long-beaked delphinines – is seemingly less reliable than classically thought. That might not be surprising given evidence from elsewhere in the tree of life that the development of sinuses and air-sacs is labile and opportunistic. I think in any case that the complex sinus system of Grampus is only superficially similar to that of long-beaked delphinines: when I look at the sinus configurations illustrated by Fraser & Purves (1960), the Grampus condition looks like an elaborate version of the simpler condition present in pilot whales, and not especially like that of Tursiops, Stenella and Delphinus.

Another interesting thing concerns snout shape (and note here that, by ‘snout shape’, I refer to the condition in the living animal: that is, where the melon and its associated soft tissues envelop much of the length of the bony rostrum). Consider that porpoises and monodontids (narwhals and belugas) are close relatives of delphinids and, as we’ve just seen, most recent phylogenetic studies find killer whales and lissodelphinines (Cephalorhynchus and the ‘lags’ and so on) to be outside the globicephaline + long-beaked delphinine clade.

Delphinid rostra as seen in dorsal view: (a) Globicephala, (b) Pseudorca, (c) Peponocephala, (d) Orcaella, (e) Orcinus. From Muizon (1988). Globicephala (pilot whales) is strangest in having such anteriorly broad premaxillae. And remember that molecular phylogenies do not recover Orcinus (killer whales) as a close relative of the others shown here. Click to enlarge.

Notably, all of these groups are short-snouted, or at best ‘medium-snouted’ (though, long-beaked fossil porpoises do confuse the picture somewhat). Could it be, therefore, that short snouts were ancestral for delphinids and widespread during most of their history, that globicephalines were primitively short-snouted (that is, they did not evolve from long-snouted ancestors), and that long snouts were only evolved at the base of the delphinine lineage? On balance, I think it’s safest to assume that modest, mid-length snouts were the norm for most of delphinid history. Globicephalines might not be as unusual (in being blunt-headed) as people used to think, but they’re still modified relative to the ancestral condition in having an especially wide bony rostrum (this is most evident in pilot whales and false killer whales) and an enlarged, anteriorly prominent melon.

Finally, what about pigmentation? Morphological character sets on extant species often neglect to include or code pigmentation data, but – while undeniably labile – it clearly carries a phylogenetic signal (and tests involving coat patterns in cats and other groups often produce encouraging results). As is obvious from the vernacular term ‘blackfish’, globicephalines are unusual among delphinids in being mostly black. If Grampus and Orcaella are part of Globicephalinae [see caveat above], but outside the ‘blackfish’ clade (Pseudorca, Feresa, Peponocephala and Globicephala), then we might infer that the greyish Grampus and Orcaella represent the primitive condition. In fact (time for some rampant speculation), Grampus is born dark and becomes paler with age, only retaining the darker pigmentation on its fins, flukes and dorsal fin (and sometimes on the chin as well). Could the dark overall colouration in the blackfish perhaps be a paedomorphic feature?

Spyhopping pilot whale, showing pale chest patch similar to that present in Risso's dolphin. Photo by Barney Moss, from wikipedia.

Also interesting is that Grampus has a wide, white area on its chest, joined by a thin ‘stem’ to another wide, white area on its belly. The same, ‘anchor-shaped’ configuration is present in pilot whales, false killer whales, pygmy killer whales and melon-headed whales: in these taxa the ‘stem’ is longer, the abdominal field is smaller, and the ‘anchor’ is sometimes greyish or off-white rather than pure white [adjacent pilot whale image by Barney Moss]. Nevertheless it’s conceivable that this is a shared derived character.

As usual, what was meant to be a quick, couple-hundred-words-article saying “Yes, it was a Risso’s dolphin” evolved in something with a bit more substance. I hope you learnt something!

For links to ALL previous Tet Zoo cetacean articles (there are quite a few of them), please see…

• All the whales of the world, ever (part I)
• All the whales of the world, ever (part II)

Refs – -

Baumgartner, M. F. 1997. The distribution of Risso’s dolphin (Grampus griseus) with respect to physiography in the northern Gulf of Mexico. Marine Mammal Science 13, 614-638.

Bearzi, G., Reeves, R. R., Remonato, E., Pierantonio, N. & Airoldi, S. 2011. Risso’s dolphin Grampus griseus in the Mediterranean Sea. Mammal Biology 76, 385-400.

Caballero, S., Jackson, J., Mignucci-Giannoni, A. A., Barrios-Garrido, H., Beltrán-Pedreros, S., Montiel-Villalobos, M. G., Robertson, K. M., Baker, C. S. 2008. Molecular systematics of South American dolphins Sotalia: sister taxa determination and phylogenetic relationships, with insights into a multilocus phylogeny of the Delphinidae. Molecular Phylogenetics and Evolution 46, 252-268.

Clarke, M. R. 1986. Cephalopods in the diet of odontocetes. In: Bryden, M. M. & Harrison, R. (eds.) Research on Dolphins. Claredon, Oxford, pp. 281-321.

Cunha HA, Moraes LC, Medeiros BV, Lailson-Brito J Jr, da Silva VM, Solé-Cava AM, & Schrago CG (2011). Phylogenetic status and timescale for the diversification of Steno and Sotalia dolphins. PloS one, 6 (12) PMID: 22163290

Fraser, F. C. & Purves, P. E. 1960. Hearing in cetaceans: evolution of the accessory air sacs and the structure and function of the outer and middle ear in recent cetaceans. Bulletin of the British Museum (Natural History) 7, 1-140.

Geisler, J. H., McGowen, M. R., Yang, G. & Gatesy, J. 2011. A supermatrix analysis of genomic, morphological, and paleontological data from crown Cetacea. BMC Evolutionary Biology 2011, 11:112 http://www.biomedcentral.com/1471-2148/11/112

Hershkovitz, P. 1961. On the nomenclature of certain whales. Fieldiana : Zoology 39, 547-565.

- . 1966. Catalog of living whales. Smithsonian Institution United States National Museum, Bulletin 246, 1-259.

Iredale, T., & Troughton, E. Le G. 1933. The correct generic names for the Grampus or Killer Whale, and the so called Grampus or Risso’s Dolphin. Records of the Australian Museum 19, 28-36.

Kruse, S., Caldwell, D. K. & Caldwell, M. C. 1999. Risso’s dolphin Grampus griseus (G. Cuvier, 1812). In Ridgway, S. H. & Harrison, R. (eds.) Handbook of Marine Mammals, vol. 6, The Second Book of Dolphins and Porpoises. Academic Press, San Diego, pp.183-212.

LeDuc, R. G., Perrin, W. F. & Dizon, A. E. 1999. Phylogenetic relationships among delphinid cetaceans based on full cytochrome b sequences. Marine Mammal Science 15, 619-648.

McGowen, M. R. 2011. Toward the resolution of an explosive radiation – a multilocus phylogeny of oceanic dolphins (Delphinidae). Molecular Phylogenetics and Evolution 60, 345-357.

- ., Spaulding, M., Gatesy, J. 2009. Divergence date estimation and a comprehensive molecular tree of extant cetaceans. Molecular Phylogenetics and Evolution 53, 891-906.

Muizon, C. de 1988. Les relations phylogénétiques des Delphinida (Cetacea, Mammalia). Annales de Paléontologie (Vert.-Invert.) 74, 159-227.

Philips, J. D., Nachtigall, P. E., Au, W. W., Pawloski, J. L. & Roitblat, H. L. 2003. Echolocation in the Risso’s dolphin, Grampus griseus. Journal of the Acoustical Society of America 113, 605-616.

Shane, S. H. 1995. Relatioship between pilot whales and Risso’s dolphins at Santa Catalina Island, California, USA. Marine Ecology Progress Series 123, 5-11.

Watson, L. 1988. Whales of the World. Hutchinson, London.

Paul (2010): cover of the A & C Black edition.

Greg Paul is an independent researcher who specialises on dinosaurs; he’s well known for his popular articles and books and his technical papers, but in particular for his hugely influential artwork. Paul’s most recent book – the 2010 Dinosaurs: A Field Guide (aka The Princeton Field Guide to Dinosaurs) – is, simply put, the ultimate Greg Paul book. It’s a large (320 pp.), heavily illustrated catalogue of over 400 reconstructed skeletons, accompanied throughout with life restorations and brief chunks of text that present data on the world’s Mesozoic dinosaur species. The idea that this book might function as a “field guide” is of course fanciful, and indeed it’s stated early on that the book is intended to be “in the style of a field guide”. A lengthy introductory section reviews dinosaur anatomy, biology, evolution, behaviour, and the climate, atmosphere and palaeogeography of the Mesozoic.

Most basic pieces of biographical information about Paul are already well known. Studying informally under the famously iconoclastic Robert Bakker at Johns Hopkins University during the 1970s and 80s, Paul became interested both in the idea that most dinosaurs were metabolically similar to extant mammals and birds, and that dinosaurs (and other fossil archosaurs) were being portrayed inaccurately by other artists. Under Bakker’s guidance, and inspired by the work of Charles Knight and other artists interested in anatomy, Paul honed a distinctive visual style for both the high-fidelity skeletal reconstructions he has become famous for, and for his reconstructions of dinosaurs in their environments.

One of Greg Paul's most famous scenes: the giant Tendaguru brachiosaur Giraffatitan brancai, with the diplodocoid Dicraeosaurus at left. Image (c) Greg Paul.

It is difficult to overstate the impact and significance of Paul’s work. Like it or not, when most of us think about dinosaurs (or other Mesozoic archosaurs), the images we have in our minds are generally “Greg Paul dinosaurs”. Paul wasn’t the first to depict slim-limbed, fully terrestrial sauropods, galloping ornithischians, or theropods with horizontal bodies and tails, drumstick-like shank muscles, or feathery pelts – Bakker did all of this in his articles from the 1960s and 70s. But the fact that Paul did this consistently, produced both black and white illustrations and colour paintings, and became published across popular mainstream sources, soon made him a dominant force in the world of dinosaur art. While many palaeontologists dislike or disagree with what Paul says about dinosaurs and their biology and evolution, I think that his role in the dinosaur renaissance, and especially in the way dinosaurs are portrayed in art and the media, should always be credited (Naish 2009).

Greg Paul's skeletal reconstructions of the iguanodontians Dollodon bampingi (top) and Mantellisaurus atherfieldensis (below). The status of the former is controversial. Images (c) Greg Paul, used with permission.

Furthermore, Paul’s massive influence mostly comes from the fact that his reconstructions have always been based on an underlying, apparently empirical effort to depict anatomy. In an ideal world, all attempts to reconstruct fossil animals would proceed this way; alas, most illustrators of prehistoric life have done their work by looking at mounted skeletons, guessing the limits of the surrounding soft tissue, and producing the final product with little or no recourse to the anatomy of extant animals. Historically, even those who knew anatomy well – Charles Knight is a classic example – thought it ok to imagine dinosaurs with massive, flabby bodies and (paradoxically) small, lizard-like muscles, despite substantial skeletal evidence to the contrary. In articles, papers and books, Paul argued that one should strive to produce multi-view skeletal reconstructions of fossil archosaurs, and that a good understanding of the overlying musculature should result in a reconstructed form that – bar integument – is essentially that of the living animal (Paul 1987, 1988, 1991). His 1987 ‘rigorous how-to guide’ on the reconstruction of dinosaurs and other Mesozoic archosaurs (Paul 1987) remains a classic that has not really been bettered, even if it does now require substantial update.

The Mongolian deinonychosaurs Velociraptor (l) and Saurornithoides (r) squabble over a Protoceratops carcass. Image (c) Greg Paul.

On the subject of integument, Paul has always been careful to restore his archosaurs with the sort of skin known from certain rare fossils – not with the imaginary wrinkled, elephant-like skin so often given to big dinosaurs by naïve artists – or, in the case of smaller or especially bird-like taxa, with feathery or furry plumes, crests or coats. I’m not alone in recalling a time when certain palaeontologists decried the reconstructing of small dinosaurs as feathery or furry as wholly unscientific and as evidence of the obviously inferior intellect and experience of Paul and his artist colleagues, but look where we are now.

Paulian dinosaurs, but not by Greg Paul. This scene - depicting Therizinosaurus, Tarbosaurus and some remarkably stupid avimimid oviraptorosaurs - is by Luis Rey.

In view of all of this, a distinctive ‘Paulian’ style is present throughout the dinosaur art world today, with many artists producing animals clearly inspired in form, pose and posture by – or even direct copies of – Paul’s dinosaurs. The Jurassic Park dinosaurs are Paulian (though, shame about those unfeathered dromaeosaurs and the burly, tree-trunk-like limbs on the brachiosaur): in fact, we even see a Greg Paul skeletal reconstruction in the movie. In view of this movement, artists who produced non-Paulian dinosaurs as late as the 1980s or 90s seemed anachronistic even at the time.

As Paul explains, it has long been an aim to reconstruct, and publish together, the skeletons of “almost all dinosaur species for which sufficient information is available” (Paul 2010, p. 6). A previous effort to produce a compilation of Paulian dinosaurs – the Japanese volume The Complete Illustrated Guide to Dinosaur Skeletons (Paul 1996) – is far less comprehensive and hard to obtain. This new volume is a guide to (nearly) all valid, non-avialan Mesozoic dinosaur species published at the time of going to press. Some species aren’t illustrated because Paul decided that published or illustrated information was insufficient to allow a reconstruction, but all are provided with a brief description. This consists of data on the taxon’s size, age and distribution, but it also includes an ‘Anatomical characteristics’ section.

Ceratosaurus pages from Paul (2010), from Princeton University Press site.

Dinosaurs: A Field Guide really is the sumptuous visual treat that dinosaur fans, and fans of Paul’s artwork and reconstructions, have long wished for. Numerous dinosaurs never before reconstructed by Paul, nor (in cases) by anyone, are present, including the theropods Juravenator, Sinocalliopteryx, Mei long, Jinfengopteryx, a scansoriopterygid (Paul calls it Scansoriopteryx, whereas the name Epidendrosaurus is probably more appropriate), the sauropodomorphs Jingshanosaurus and Gongxianosaurus, the ankylosaurs Saichania and Pinacosaurus, Pachycephalosaurus, and Olorotitan and numerous other hadrosaurs. Surprises come in the form of the apparently ridiculous Atlasaurus, Beipiaosaurus (but see below), the absurdly shaped Gastonia and the impressively long neural spines of Hypacrosaurus altispinus. Paul describes in the Preface how the recent explosion of new dinosaur discoveries – concerning new feathered theropods, new data on dinosaur pigmentation, skin texture, and bizarre new groups like therizinosaurs – make the modern world of dinosaur research radically different from the one he first got to know.

Three negative points

One of the stegosaur pages from Paul (2010), from NHBS site.

Three main points stand out as areas that could definitely have been improved upon. The first concerns the descriptive text, the second the volume’s format, and the third the taxonomy and classification that Paul uses throughout. At this point it has to be noted that this book will primarily be used by artists; it will not be used by, nor is it intended for, technical scientists, the majority of which seem to have little interest in, or knowledge of, inferred dinosaur life appearance.

Negative point 1: descriptive text

On that note, the ‘Anatomical characteristics’ section included in the descriptive text for each species is frequently all but useless, failing to include material that seems appropriate. I think this is because Paul’s general aim was to describe the overall form of the animal as if it were alive, and not to refer to autapomorphies or other distinctive bony attributes. The tendency to describe species as “standard” for their group, or as “uniform” or “fairly uniform” is frustrating, but refers I suppose to the fact that the respective animal is inferred to be typical for its group in general shape. The reference to contemporaneous species as “enemies” is slightly irksome. Should “friends” have been listed as well?

The pages of field guides.

Negative point 2: format

On the second negative point, the book’s format is also frustrating. ‘Field guides’ do not have illustrations of animals scattered throughout: rather, similar species are illustrated together such that the reader can best appreciate their similarities and differences. I really wish that this format had been used in Dinosaurs: A Field Guide – it would have made it so much earlier to compare related animals. To give one example: apatosaurine sauropod skeletons are spread across three pages, making it hard to appreciate how Apatosaurus ajax compares in limb and body proportions to the three species that Paul considers similar enough to be grouped together as ‘Brontosaurus morph’ apatosaurines.

It is also unfortunate that specimen numbers and scale bars are not included alongside the reconstructions. This would have increased the volume’s value enormously and would make it look more authoritative. It’s not as if there wasn’t the room for the inclusion of such data: there are enormous white spaces alongside every single reconstruction. Paul does say early in the book that the specimen numbers of all reconstructed individuals are provided in an online appendix, archived at http://press.princeton.edu/titles/9287.html. A link available through that site does include specimen numbers, but they’re listed in connection with size estimates – it isn’t immediately clear that they refer to the same specimens used in the reconstructions, but I suppose they must.

Negative point 3: taxonomy, phylogeny, nomenclature

How many 'genera' do you see, one or three? It doesn't matter, so long as we know what we mean. Most workers would interpret these three skulls as (clockwise from top left) representing Centrosaurus, Styracosaurus and Pachyrhinosaurus. Images from wikipedia: see below.

On the third negative point, one of the first rules about this book is that Paul’s taxonomy should be ignored. We all know that Greg has his own views on which taxa should be congeneric, but the use of names across the book is misleading and totally confusing, especially for non-specialists. Paul has argued before (e.g., Paul 1988) that dinosaur genera are oversplit and that tyrannosaurid and ornithomimid species, various of the centrosaurine horned dinosaurs, some of the crested lambeosaurine duckbills and so on should really be placed in the same genera, in part because (say) Styracosaurus and Pachyrhinosaurus are (in Paul’s view) more similar to one another than are species included in such extant genera as Varanus. [In composite image above, Centrosaurus by Sainterx, Styracosaurus by Claire Houck, Pachyrhinosaurus by Sebastian Bergmann.] Authors are of course free to reclassify species based on their own understanding or interpretation of phylogenetic relationships (this is itself an unfortunate consequence of the fact that Linnaean binomials frame a specific hypothesis for a taxon’s affinities; they are not just labels). But authors should only do this when they can justify or explain their reclassifications. Paul doesn’t do this, and in fact his reclassifications seem mostly based on whimsy.

But a little of the morphological variation present within Varanus. Top to bottom: Komodo dragon (V. komodoensis), Savannah monitor (V. exanthematicus), Green tree monitor (V. prasinus).

In any case, it is both naïve and futile to imagine that the entities we term genera should be somehow consistent across different animal clades. It doesn’t make any difference as goes our understanding of diversity or phylogeny whether the name Centrosaurus is understood to be unique to the species apertus, or to apply to all species within the clade typically called Centrosaurinae, or whether Centrosaurus encompasses the same amount of morphological disparity as Varanus, Loxodonta or Cricetus; what matters is that researchers within a given subject area use a consistent nomenclature. Many people – and remember that this is a popular volume, not written just for dinosaur nerds and technical specialists – just won’t get it should they see, for example, the Argentinean Giganotosaurus referred to as a species of the otherwise African Carcharodontosaurus. All in all, the taxonomy that Paul opts to use in his volume represents a real failure of communication. Readers will be perplexed, not enlightened. [In adjacent composite, Komodo dragon photographed at ZSL London Zoo, Savannah monitor image by Shizhao, tree monitors by El Cattivo86.]

On a related note, the way in which Paul classifies the species is frustrating, not necessarily because he is idiosyncratic, but rather because the placements he favours are not explained or justified. To take just one example… Sapeornis chaoyangensis – a long-armed, short-skulled, short-tailed theropod with a pygostyle and reversed hallux – is generally regarded as part of Avialae, close in phylogenetic position to the root of Pygostylia (Zhou & Zhang 2002). Yet Paul regards it as an oviraptorosaur – a placement that many of us have considered informally but one which has yet to be supported by any character analysis. The idea that the long-winged, presumably volant Sapeornis might be an oviraptorosaur is, needless to say, an exciting speculation in view of Paul’s idea about the repeated evolution of flightlessness in Mesozoic maniraptorans (Paul 1988, 2002), but that’s all it is – a speculation. The classification of Epidexipteryx hui within Oviraptorosauria also requires explanation, and why is Epidexipteryx regarded as an oviraptorosaur while the extremely similar Epidendrosaurus is not?

In a few cases, Paul uses new names for certain clusters of species. I can understand that one might need to lump animals together into paraphyletic grades for the purposes of a book like this, but I think it’s misleading to refer to (say) non-lithostrotian titanosaurs as ‘baso-titanosaurs’ (as Paul does), since this immediately creates the impression that such a name is in widespread use (it isn’t) and will be understood by others (it won’t). Minmi and Liaoningosaurus are grouped together as ‘minmids’ and what exactly are ‘paxceratopsians’? So far as I can tell, the latter name is wholly novel (Paul uses it for psittacosaurs and neoceratopsians).

One of my favourite Greg Paul pieces: pair-bonded, nesting azhdarchids with altricial chicks fend off a tyrannosaurid. Image (c) Greg Paul. Art like this is controversial specifically because it is bold and daring. And, no, that doesn't mean that I support the behavioural hypotheses depicted in this painting.

In a work of this size, particularly one containing both novel skeletal reconstructions and a huge amount of controversial re-naming and re-classifying, it’s inevitable that numerous small matters of contention will arise. Some researchers say that the very raison d’etre of this work – Paul’s body of high-fidelity skeletal reconstructions – is problematic, with the underlying reconstructive process being subjective, prone to bias and misinterpretation, and far more artistic than Paul makes clear (Mallison 2010, Norman 2011). Producing skeletal reconstructions of this sort involves extrapolation, interpretation and a degree of guesswork, so perhaps it would be helpful – and this is not a specific criticism of Paul, but one that could be directed at all technical skeletal reconstructions – if the reconstructions were framed as hypotheses where some (SOME, not all) of the details are open to alternative interpretations. Indeed, some of the things that look intuitively reasonable in reconstructions – Paul’s quadrupedally bounding plateosaurian sauropodomorphs are a good example – are unlikely to be correct, in this particular case because, among other things (e.g., Mallison 2010), plateosaurs could not pronate the manus (Bonnan & Senter 2007) and were thus likely incapable of quadrupedal walking or running.

Life reconstruction of Guanlong wucaii, by Renato Santos (from wikipedia).

On more trivial matters, I especially disliked seeing the tyrannosauroid Guanlong included among the allosauroids as a second species of Monolophosaurus (note that Monolophosaurus itself is no longer regarded as an allosauroid by other workers: Benson 2010, Brusatte et al. 2010). This idea only has its origin in a conference abstract and its associated presentation (Carr 2006) and the full argument has yet to be presented in print. Whether it withstands scrutiny or not (I agree with Brusatte et al. (2010) that it seems unlikely to be correct), Paul shouldn’t have reclassified Guanlong on the basis of a conference presentation. Neovenator, an uncontroversial allosauroid allied with carcharodontosaurids, is said by Paul to be the centre of a disagreement over whether it’s a tyrannosauroid or allosauroid. Unless I’ve missed something, this is incorrect.

Therizinosaur Beipiaosaurus, with 'dorsal fin' visible above its dorsal vertebrae.

What’s with the crazy ‘dorsal fin made of fuzz’ depicted on the therizinosaur Beipiaosaurus? Paul illustrates another, later therizinosaur – Nothronychus – with a subtriangular ridge in the middle of its back (that is, with the peak of the ridge being mid-way along the length of the torso), but here the ridge is formed by tall neural spines. My hypothesis is that Paul is hinting at the possibility that therizinosaurs found mid-dorsal fins highly fashionable, and swapped soft-tissue ones for bony ones at some point during their evolution (elsewhere in the book, Beipiaosaurus gets referred to as “a refugee from a Warner Brothers’s cartoon” (p. 12)). But is there any evidence for a ‘dorsal fin’ in Beipiaosaurus? An excellent referred specimen preserves the subtriangular patch of integument that Paul has restored as a ‘dorsal fin’ (Xu et al. 2009), but, like other integument-bearing patches preserved adjacent to the specimen, I don’t think you can confidently infer that it represents a soft-tissue structure in life position. The Liaoning Province fossils are mostly flattened. Like most fossils with soft tissues attached, their outlines are not sharp and life-like, but messy and the result of decomposition and compression. Taphonomic experiments have shown that, when the bodies of modern birds are crushed flat, large feathery crests not present in life typically result (Foth 2011).

Reconstructed skull of Nemegtosaurus (though - controversially - given 'cheeks'), by palaeozoologist (NOT by Paul).

Paul puts the Nemegtosaurus skull on the Opisthocoelicaudia body [adjacent skull illustration by palaeozoologist]. This is objectionable if you follow phylogenies where the two are recovered as but distantly related (e.g., Curry Rogers 2005), and it isn’t supported by any overlapping material (see discussion in Wilson 2005). For now, it’s an interesting idea, but not a ridiculous one, especially in view of the fact that – the more we learn about sauropod skulls – the more samey they appear. Also on sauropods, Paul’s idea that the raised tails present in some mamenchisaurid specimens might represent the actual life condition has so far proved unpopular (I know: I brought it to attention at a recent meeting devoted to sauropods). I’ll leave this idea for those with more experience of sauropods and their tails.

The panel-mounted chasmosaurine NMC 8538 as we're used to seeing it. Skull comes from a different specimen and the ribs are scrunched up on right side. Photo courtesy of ReBecca Hunt-Foster.

Styracosaurus albertensis (Centrosaurus albertensis of Paul’s usage) is illustrated with an enormously long, tapering nasal horn, despite the fact that Ryan et al. (2007) showed the actual horn to be about half as long as the one reconstructed on mounted specimens. Also on ceratopsids, we now know that the wonderfully complete ‘Anchiceratops ornatus’ skeleton NMC 8538 (shown here), reconstructed by Paul, is a composite (the skull is a cast from another specimen). Furthermore, it turns out that it was only ever identified as Anchiceratops by default, not because it can be convincingly referred to this taxon (Mallon & Holmes 2010). On the subject of ceratopsians, Cerasinops is inadvertently featured twice in the book, and Paul accepts the proposal that Torosaurus is a growth stage of Triceratops, not a separate taxon. On ornithopods, we now know that the cranial crest of Tsintaosaurus spinorhinus is not just a sub-vertical spike as classically depicted, but a more complex structure. I think that some of the dryosaurids in the book are incorrectly labelled.

Another of my favourite Greg Paul paintings. Giraffatitan and Ceratosaurus ingens contemplate one another; pterosaurs and a mammal are in the foreground. The full version includes several diplodocids at far right. Image (c) Greg Paul.

It’s a matter of taste whether you like or dislike Paul’s artwork. The flat horizons, low viewer angle, and tendency of animals to be depicted in tidy profile are features disliked by some. And a minor artistic and scientific movement away from ‘shrink-wrapped’, Paul-style dinosaurs (that is, those where the edges of many of the skull bones, and the distinct margins of the bony skull openings, are far more distinct than they are in any living animals) is currently underway. Anyway, those who enjoy Paul’s colour pieces will be happy to see several new ones included here. Paul also includes modified, ‘colourised’ versions of pieces that were either originally produced in black and white, or originally showed the animals standing alone, without any obvious background. These haven’t worked at all in my opinion (e.g., Lesothosaurus diagnosticus on p. 216, Euoplocephalus tutus on p. 235). The many small life reconstructions of animal’s heads suffer from their fuzzy outlines – a real contrast to the typically sharp lines of Paul’s artwork.

Some of the spelling is sloppy. ‘Ornithsichians’ are mentioned on p. 273 and Paul’s own taxon Dollodon bampingi is consistently referred to as Dollodon bambingi (pp. 289-290). Then there’s Crylophosaurus (p. 11) (= Cryolophosaurus), Pacycephalosaurus (p. 22) (= Pachycephalosaurus), Archiornis (p. 52) (= Anchiornis), Pycnoneosaurus (p. 79) (= Pycnonemosaurus), Aerosteons (p. 99) (= Aerosteon), Ansermimus (p. 113) (= Anserimimus) and so on.

What is the purpose of this book? People will enjoy looking at it, for sure, but doesn’t Paul produce those high-fidelity skeletal reconstructions so that other scientists and artists have access to excellent, useable data on the animals that Paul reconstructs? Well, no, apparently not, for the great irony is that a major ruckus erupted in the dinosaur art world at about the same time as this book appeared in print.

The 'no GSP' logo you can find online. I won't say who produced it.

Apparently inspired by the fact that he was losing paying work to competitors, Greg Paul spent considerable time and effort during 2011 demanding that other artists stop using his work as reference points for their own art, that people compensate him adequately should they use or pass his reconstructions on to others, and that those who produce skeletal reconstructions of their own should ensure that the poses they choose are visually distinct from the Greg Paul standard. Paul’s proposals understandably caused great animosity from many quarters: an enormous amount of discussion is archived online on this issue at blogs and the dinosaur mailing list and I don’t wish to cover it here. Suffice to say that, while Paul should get appropriate credit for his work and art, he cannot and will not stop people from using his published work as an available resource.

Princeton University Press cover.

Any lengthy piece of work by Greg Paul is likely to attract a degree of criticism, either because of his taxonomic and phylogenetic proposals, because of certain controversial or uncertain details of his reconstructions or art, or because of his outspoken views on dinosaur physiology or behaviour. But, then, these areas are part of the appeal. People like reading what Paul writes because he is well known for being controversial; for promoting new and sometimes daring and weird ideas about dinosaur evolution, biology, locomotion and behaviour. This is the person who was arguing for feathered theropods and fuzzy ornithischians before such ideas went mainstream, argued for the evolution of widespread secondary flightlessness across maniraptoran theropods, and worked to emphasise the idea that watching live dinosaurs would be like watching modern animals on the Serengeti – there would be dust in the air, interspecies conflicts, occasional herbivory in predators and occasional carnivory in herbivores. If Dinosaurs: A Field Guide is anything, it is indeed a visual treat, with reconstructions of tens of species representing most of Mesozoic dinosaur diversity as we currently understand it. What’s more, the book is extremely affordable for its size and quality. Bottom-line: most people interested in dinosaurs will value it, despite its problems.

The book comes in two editions. The US version, published by Princeton University Press, features two running tyrannosaurs on the cover and is properly titled The Princeton Field Guide to Dinosaurs. The UK version, published by A & C Black, is black with six life restorations on the cover. The cover image shown at the very top of this article has three ostrich dinosaurs at the top and I’m not sure if this version exists in the real world: the one I have with me here now features Leptoceratops, Euoplocephalus and Shantungosaurus at the top.

Paul, G. S. 2010. Dinosaurs: A Field Guide. A & C Black (London). Hardback, index, refs, pp. 320. ISBN 978-1-4081-3074-2.

For previous Tet Zoo articles relevant to some of the stuff discussed here, see…

• Evidence that too much Greg Paul is bad for you
• My old, sad t-shirt
• Terrestrial stalking azhdarchids, the paper
• A quick history of tree-climbing dinosaurs
• No-one talks about Anchiceratops, boo hoo
• Goodbye super-inclusive Iguanodon, hello Mantellisaurus, Owenodon, Dakotadon, Dollodon, Barilium, Kukufeldia, Hypselospinus, Sellacoxa, Proplanicoxa etc. etc.
• Luis Chiappe’s Glorified Dinosaurs: The Origin and Early Evolution of Birds
• Long and Schouten’s Feathered Dinosaurs, a review
• The ‘Birds Come First’ hypothesis of dinosaur evolution
• What they’re saying about The Great Dinosaur Discoveries

Refs – -

Benson, R. B. J. 2010. A description of Megalosaurus bucklandii (Dinosauria: Theropoda) from the Bathonian of the UK and the relationships of Middle Jurassic theropods. Zoological Journal of the Linnean Society 158, 882-935

Bonnan, M. F. & Senter, P. 2007. Were the basal sauropodomorph dinosaurs Plateosaurus and Massospondylus habitual quadrupeds? Special Papers in Palaeontology 77, 139-155.

Brusatte, S. L., Benson, R. B. J., Currie, P.  J. & Zhao, X. 2010. The skull of Monolophosaurus jiangi (Dinosauria: Theropoda) and its implications for early theropod phylogeny and evolution. Zoological Journal of the Linnean Society 158, 573-607.

Carr T. 2006. Is Guanlong a tyrannosauroid or a subadult Monolophosaurus? Journal of Vertebrate Paleontology 26, 48A.

Curry Rogers, C. 2005. Titanosauria: a phylogenetic overview. In Curry Rogers, C. & Wilson, J. A. (eds) The Sauropods: Evolution and Paleobiology. University of California Press (Berkeley), pp. 50-103.

Foth, C. (2011). On the identification of feather structures in stem-line representatives of birds: evidence from fossils and actuopalaeontology Paläontologische Zeitschrift DOI: 10.1007/s12542-011-0111-3

Mallison, H. 2010. The digital Plateosaurus II: An assessment of the range of motion of the limbs and vertebral column and of previous reconstructions using a digital skeletal mount. Acta Palaeontologica Polonica 55, 433-458.

Mallon, J. C. & Holmes. R. 2010. Description of a complete and fully articulated chasmosaurine postcranium previously assigned to Anchiceratops (Dinosauria: Ceratopsia). In Ryan, M. J., Chinnery-Allgeier, B. J. & Eberth, D. A. (eds) New Perspectives on Horned Dinosaurs: the Royal Tyrrell Museum Ceratopsian Symposium. Indiana University Press (Bloomington and Indianapolis), pp. 189-202.

Naish, D. 2009. The Great Dinosaur Discoveries. University of California Press (Berkeley and Los Angeles).

Norman, D. B. 2011. Ornithopod dinosaurs. In Batten, D. J. (ed.) English Wealden Fossils. The Palaeontological Association (London), pp. 407-475.

Paul, G. S. 1987. The science and art of restoring the life appearance of dinosaurs and their relatives – a rigorous how-to guide. In Czerkas, S. J. & Olson, E. C. (eds) Dinosaurs Past and Present Vol. II. Natural History Museum of Los Angeles County/University of Washington Press (Seattle and London), pp. 4-49.

- . 1988. Predatory Dinosaurs of the World. Simon & Schuster (New York).

- . 1991. The many myths, some old, some new, of dinosaurology. Modern Geology 16, 69-99.

- . 1996. The Complete Illustrated Guide to Dinosaur Skeletons. Gakken.

- . 2002. Dinosaurs of the Air: the Evolution and Loss of Flight in Dinosaurs and Birds. Baltimore: Johns Hopkins University Press (Baltimore), pp. 536.

- . 2010. Dinosaurs: A Field Guide. A & C Black (London).

Ryan, M. J., Holmes, R. & Russell, A. P. 2007. A revision of the late Campanian centrosaurine ceratopsid genus Styracosaurus from the Western Interior of North America. Journal of Vertebrate Paleontology 27, 944-962.

Wilson, J. A. 2005. Redescription of the Mongolian sauropod Nemegtosaurus mongoliensis Nowinski (Dinosauria: Saurischia) and comments on Late Cretaceous sauropod diversity. Journal of Systematic Palaeontology 3, 283-318.

Xu, X., Zheng, X. & You, H. 2009. A new feather type in a nonavian theropod and the early evolution of feathers. Proceedings of the National Academy of Sciences 106, 832-834.

Zhou, Z. & Zhang, F. 2002. Largest bird from the Early Cretaceous and its implications for the earliest avian ecological diversification. Naturwissenschaften 89, 34-38.

We all love identifying – or, trying to identify – weird carcasses. Back in December 2011, marine biologist and world chiton expert Douglas Eernisse of the University of California (Santa Cruz) sent me the series of photos you see here and below. They show a smallish cetacean (total length about 3 m), found beached and in partly decomposed condition at Loreto, Baja California. A large damaged region on the animal’s tailstock seemed to demonstrate death by collision with a boat.

The people who discovered the carcass identified it as a Pygmy sperm whale Kogia breviceps and noted that it possessed the asymmetrical blowhole typical for sperm whales. Hmm.. oh really?

Douglas and I have been discussing the carcass in an effort to identify it, but in the interests of sharing the fun, I have Doug’s permission to post the images here. Check out the many anatomical details you can see in the photos and see if you can pin down the carcass’s precise identity. To the winner – the spoils!

Thanks again to Douglas Eernisse for the photos. For previous Tet Zoo articles on weird marine mammal carcasses, see…

• Santa Cruz’s duck-billed elephant monster
• It had wool, and armour plates, a massive beak, horns, and it smelled veeeeery bad: whatever happened to the Tecolutla monster?
• Where are all the dead sea monsters?
• Skull of the Moore’s Beach monster revealed!
• England ‘does a Montauk’
• A Russian sea monster carcass is claimed to be that of an ancient ‘archaeocete’ whale
• In which the Conakry Monster carcass leads to a digression on ‘tubercle technology’

A few good books have already reviewed Australian big cat sightings. Karl Shuker’s 1989 Mystery Cats of the World is a classic, and Tony Healy and Paul Cropper’s 1994 Out of the Shadows includes a good section on Australian big cats too. One of the most influential books on the subject is David O’Reilly’s 1981 Savage Shadow: the Search for the Australian Cougar [recently republished by Strange Nation Publishing; adjacent cover image of 1981 edition from Mike Williams’s Australian Big Cats blog]. O’Reilly’s book mostly centres around the experiences of those who clamed to have seen (or experienced the depredations of) the ‘Cordering Cougar’ in West Australia during the 1970s. One of the main contentions about the ‘puma’ phenomenon wasn’t just that people were seeing big, puma-like cats in the West Australian bush, but also that government officials were unwilling to investigate or make announcements about it. This apparent lack of government action has been a consistent theme throughout the Australian ‘mystery big cat’ experience.

A long term interest and involvement in Australian and world mysteries led Michael Williams and Rebecca Lang to research and produce what is now the definitive volume on Australian mystery big cats; it’s titled Australian Big Cats: An Unnatural History of Panthers (Williams & Lang 2010). At 434 pages, it’s substantial. It’s also highly readable, nicely formatted and very well illustrated. The authors have collated a vast amount of information gleaned not only from published sources but also from interviews with both eyewitnesses and people who have examined evidence firsthand. Williams and Lang clearly travelled widely across the country, photographing locations, people, documents, taxiderm specimens and so on at what must have been great personal expense. They obtained freedom of information acts and other previously undisclosed documents. A lengthy appendix (c. 120 pages) includes copies of numerous letters and documents produced by government officials, veterinarians, ecologists, geneticists and others. The volume is fully referenced (though with the citations given at the bottom of the respective pages, rather than at the end of the text) and with an index.

So, to anyone seriously interested in mystery animals, mystery big cats or Australian mammals in general, this book is a must-have. Never before has so much data been gathered together on the subject: well done to the authors on this substantial achievement. [Graphic below borrowed from Centre for Fortean Zoology Australia].

Map of Victoria showing locations of reported sightings. From a May 2010 newspaper article.

There are wild big cats in Australia

As I (generally) always say when talking about mystery animals, remember that these phenomena aren’t just interesting because there might be real, flesh-and-blood animals at the bottom of the reports: even if there are not, mystery animal sightings, accounts and stories are still fascinating and research-worthy subjects, combining as they do psychology, sociology, folklore and human observational skills and biases. Remember that some people who would ordinarily be labelled cryptozoologists are quite happy to be regarded as folklorists.

However, while the mystery big cat phenomenon does indeed involve sociology and folklore, there’s no doubt that at least some sightings involve real animals. Large (sometimes black) feral dogs and dingoes, foxes and even wallabies explain some ‘big cats’ sightings, but not all of them. Australian big cats aren’t just represented by eyewitness accounts and hazy photos, but by some pretty good photos, and also by a number of dead bodies. Let’s look at some of these cases.

The St Arnaud puma, shot in 1924.

Among the more impressive photos is that taken by Barry Morris in 1978 near Carnarvon in West Australia. It shows a big, black cat walking along the top of a hillside, its long, cylindrical tail held in a curve up over its rump. A puma that escaped from a travelling circus, and lived wild for a time, was shot at St Arnaud in Victoria in 1924 while another puma was shot at Woodend, Victoria, during the 1960s. The Woodend animal was stuffed and then pretty much forgotten about until 2005. And in 1985, a lioness – the ‘Broken Hill lioness’ – was shot in New South Wales. This case has become notorious due to its apparent lack of investigation by the Department of Agriculture (Williams & Lang 2010). Remarkably, vocalisations apparently made by large cats living wild in the Australian bush have been caught on tape at least twice.

Footprint cast at Hawkesbury, NSW, in 2011, and thought to have been made by an animal that killed a captive alpaca. Sure looks like a large felid print to me.

Various footprints, scat and large animal kills attributed to big cats have also been recorded [adjacent footprint photo from here]. Williams & Lang (2010) publish many of these. Many of the tracks do look unmistakeably cat-like, and vets, ecologists, professional mammalogists and government officials are on record as saying that large cats are indeed the most likely, or only likely, culprits (Williams & Lang 2010, pp. 253-272). As Williams & Lang (2010) explain, some of the more positive assessments (including those penned by Charles Sturt University ecologist Johannes Bauer, Deakin University’s John Henry, and veterinarians Keith Hart and Ron Hynes) have been essentially buried or kept quiet by some of the governmental bodies that have been asked by farmers and stock-owners to investigate. So, there are non-native big cats running around in the Australian outback, at least sometimes.

The so-called 'Territory Tiger': a sandy-coloured cat, photographed by Jan Donovan in 2007. It looks large but it's just about impossible to accurately judge scale. It might be a puma or a large, skinny feral.

Pumas, moggies, marsupials: competing or overlapping hypotheses of origin

Black jaguar photographed in captivity. There's something weird about this animal's tail.

In my view, the Australian ‘mystery big cat’ phenomenon is made especially interesting by the fact that three very different hypotheses have been invoked to explain the identity of the creatures involved. Some researchers contend that two or even all three of these hypotheses have merit.

Hypothesis 1 is that some or all of the cats are the descendants of military mascots or escapees from circuses, private collections, zoos and so on [adjacent Black jaguar photo by Cburnett]. They thus represent pumas from the America, lions from Africa, leopards from Africa or Asia, and so on. Hypothesis 2 is that feral cats have grown to extraordinary size in the Australian bush, and that these monster moggies account for some or all ‘big cat’ sightings. Hypothesis 3 – the most extreme and interesting idea, I think – is that some of the animals are not cats at all, but big, cat-like marsupials that either represent new species, or late-surviving members of one of the thylacoleonid (= marsupial lion) taxa.

As should be clear from what I’ve already said, I think we can be pretty confident that some sightings of Australian big cats really do represent encounters with lions, pumas and members of other species (perhaps including Golden cat, Jaguar and even Tiger). What about hypothesis 2? Long-time readers will recall my ‘Australia’s new feral megacats’ article of 2007. Various photos and bits of film seem to show feral cats – that is, members of the same species as the domestic cat (Felis catus or whatever you choose to call it) – that are extraordinarily big, with shoulder heights of about 60 cm and total lengths exceeding 1.5 m.

Kurt Engel, a dead megacat, and a lot of forced perspective.

Two keys bits of evidence in particular seem to support the ‘feral mega-cat’ hypothesis. One is the cat shot dead by hunter Kurt Engel in Gippsland, Victoria, in 2005. This animal was claimed to be somewhere round about 1.6 m long (in Williams & Lang (2010), Engel says that it was over 2 m long in total), with its tail alone being 60 cm in length. The fact that Engel discarded the body and deliberately used forced perspective in his photos of the carcass haven’t exactly helped add credibility to the case, but a photo published by Williams & Lang (2010) does seem to support these rough measurements. Williams & Lang (2010) cover this case at some length and explain how they helped arrange for DNA testing on tissue from the animal’s tail. These tests (the reports are included in the volume’s appendices) identified the animal as Felis catus.

Still from the 'Lithgow panther' footage.

The second bit of evidence is the so-called ‘Lithgow panther’ footage, filmed in 2001 by Gail and Wayne Pound at Lithgow, New South Wales. After catching sight of a surprisingly large black cat in the scrub near their house, the Pounds decided to film it and managed to get 15 minutes of footage. The cat was in close association with a normal-sized feral cat, yet (as demonstrated by people who visited the site and measured the height of adjacent vegetation) had a shoulder height of about 50 cm and hence was more like a puma in size (shoulder height 60-70 cm) than a domestic cat (shoulder height 25-30 cm).

Opinion differs as to whether the existence of these really big feral cats is remarkable or not. I think it’s at least very interesting, in part because I find appealing the idea that something about the Australian ecosystem is encouraging large size in some Australian feral cat populations. And once a cat of any sort approaches or exceeds a metre in total length, people who see it will refer to it as a ‘big cat’. So, some ‘big cats’ are not big cats in the strict sense at all, but big ‘small cats’.

Modern marsupial lions and other marvels

Cartoon showing (l to r) big black cat, Queensland tiger as reconstructed by Heuvelmans, the Rilla Martin animal, and a thylacine.

Hypothesis 3 rests on the idea that a few eyewitnesses have described animals that, while cat-like, supposedly exhibit weird, sometimes marsupial-like traits. A few decades ago, the idea that a long-tailed, stripy, leopard-sized Australian animal might exist in Queensland (and perhaps elsewhere) was fairly popular (Heuvelmans 1995). Dubbed the ‘Queensland tiger’, it was regarded by some as a possible living species of marsupial lion (a group of extinct marsupials, properly called thylacoleonids, otherwise thought to have become extinct during the Pleistocene: see this article for more) (Shuker 1989, Healy & Cropper 1994). Alas, the idea that the Queensland tiger was real has mostly fallen away now given the total absence of material evidence, photos and recent eyewitness accounts.

But while belief in the Queensland tiger has mostly evaporated, it’s thought by some cryptozoologists that various of the black or tan-coloured Australian ‘big cats’ might be marsupial lions too. I learnt of this idea from both Healy & Cropper (1994) and from Rex Gilroy’s terrible but entertaining book Mysterious Australia. Among the various tales that Gilroy recounts is one where the witness describes seeing a rear-facing pouch and joey in a big black ‘cat’ (Gilroy 1995). The witness surmised – and Gilroy agreed – that at least some Australian ‘big cats’ are not cats at all, but extant marsupial lions. That’s pretty neat stuff; shame that other Australian researchers have failed to record similar accounts, or indeed to verify the existence of the witnesses that Gilroy quoted… though read on.

Rilla Martin's 1964 'Ozenkadnook tiger' photo.

The face of 'Jaws'. Poor puss.

Williams and Lang devote a chapter to the supposed existence of modern day, thylacoleonid-like predators. Some of my favourite cases included in this chapter include Rilla Martin’s creature, photographed in 1964 and previously covered here on Tet Zoo, and the ‘Jaws’ carcass, found on a beach sometime in the 1980s (Shuker 1996, Williams & Lang 2010) and suggested by some writers to be a dead thylacoleonid (as the authors note, and as I hoped to demonstrate, it was actually just a dead domestic cat). Incidentally, thanks to the authors, I’ve actually gotten to see a filmed interview with Rilla Martin where she describes her encounter. It was very interesting to see her recount the tale in her own words. [Thylacoleo image below by Karora, from wikipedia.]

Thylacoleo skeleton, on display at Naracoorte Caves. Photo by Karora.

A few accounts, photos and bits of film are highly intriguing in view of ‘hypothesis 3’. Williams & Lang (2010) might not verify Rex Gilroy’s accounts, but they do provide some equally surprising ones. A farmer, searching for a missing cow in 2005, found that it had been severely wounded by a broad-headed predator, still in attendance, that “seemed to have some marsupial-like attributes” and was long-bodied, short-legged and long and thick in the tail. The cow’s calf had been killed. The photo taken in 1981 by Martin Williams as she “wandered down to the lagoon on her Moyston, Victoria property to take photographs one afternoon” (p. 205) is peculiar. The photo isn’t great (the animal is facing away from the photographer and its outlines are hard to demarcate from the waterhole behind it), but the animal’s apparently muscular hindlimbs and short, very slender tail don’t look right for a cat, dog or just about anything else you might think of.

There’s also a very odd piece of film taken in 1994 (stills are provided in the book) where a stocky, short-tailed animal with a distinctive gait and deep, boxy head runs alongside some overgrown railway tracks. The animal looks weird and I could probably convince myself that it doesn’t represent a big cat, feral dog or a member of any other known species. However, as is typical for footage of this sort, the animal is at a distance, the footage is fuzzy, and I conclude that it’s probably not possible to say for sure just what the animal is.

And what to make of the weird, bushy-tailed animal – apparently a predatory marsupial of some sort – described by Gary Opit after his 1969 night-time encounter? Based on Opit’s (not wholly unique) account, the animal couldn’t have been a surviving thylacine or anything known to be alive today.

Gary Opit's weird, bushy-tailed ?marsupial. From Williams & Lang (2010).

Necessary pedantry: things to dislike

There are a few things that I really don’t like about Australian Big Cats. The book includes a huge number of photos, eyewitness drawings and other illustrations – that’s great. But I often found it difficult to relate the figures to the nearby text, and to find that specific part of the text that describes something shown in one of the figures. So things would have really been improved had the figures been numbered (as in: Fig. 1, etc.). I’m also not too keen on the way the figures are presented. All too often, they’re reproduced at tiny size and the details are hard to make out due to their reproduction in black and white. To make matters worse, virtually all of the images have thick white borders round their edges. These serve only to make the images even smaller. So, bigger, clearer pictures would have been nice. Given that the book is already 437 pages long, you might think that the authors (and/or the designers) were very keen to do things not to make it any larger. But this brings us to another issue – the extraordinary amount of wasted space in this book.

Firstly, there are many sections where the authors write in short, one-sentence paragraphs. That works for a tabloid newspaper, but not for a detailed tome such as this where it can be assumed that the reader has an attention span exceeding a few seconds. This style of writing also ruins the flow of the text and creates a lot of wasted space on the page. Demonstrating to your prospective readership that you have the facts and figures at your fingertips – especially when writing about something as controversial and problematic as Australian big cats – is half the battle, and I feel that this is destroyed by this kind of choppy, disjointed writing. I’m sure I’m not alone in disliking a book once I see that much of it is empty (to anyone with the book at hand: random examples include pages 163, 234, 241 and 284).

This spacey look isn’t just created by the disjointed writing style; it’s exacerbated by the book’s design. I just do not understand why some publishers choose to waste enormous amounts of paper by designing books where the pages have huge, bland borders. Granted, you don’t want text to disappear in the recesses of the book’s spine, and there has to be enough room for the reader’s fingers, but… come on: each and every page in Australian Big Cats includes a 35 mm border at the bottom, a 42 mm border at the top, a 25 mm border on the right, and a 17 mm border on the left! Again, I dislike this sort of thing because it creates an airy, data-free look to a volume.

l to r: Mike Williams, Darren Naish, Rebecca Lang. Hopefully we're still friends.

As I hope is clear from this review, overall I found Australian Big Cats an impressive piece of work. The amount of research involved in its production was clearly vast, and the authors did a great job in presenting the enormous body of data they collected in a readable, enjoyable format. Williams and Lang have also done us all a great service in going ‘straight to source’ behind the scenes by digging out previously unreleased documents using freedom of information requests and so on. As outlined above, I do wish that Australian Big Cats had been presented in a different, in general ‘more technical’, way, but I say again that it is, to date, the ultimate book on the subject and one that should definitely be consulted by anyone seriously interested in Australian cryptozoology, or indeed in cat lore or biology in general. Given that Australian ‘big cats’ – whatever they are – have received all too little ‘official’ attention, will this huge book be the catalyst that helps break down the stigma that still surrounds this fascinating subject?

Williams, M. & Lang, R. 2010. Australian Big Cats: An Unnatural History of Panthers. Strange Nation, Hazelbrook, NSW. ISBN 978-0-646-53007-9. Softback. References. Index. Illustrations. 437 pp.

For more on Australian mystery animals see…

• Australia’s new feral mega-cats
• Lithgow mega-cat footage goes live
• What was the animal in the ‘Jaws’ photo?
• Identifying that ‘Jaws’ carcass
• The 2006 Night parrot: dead, decapitated, evidence for collision with a fence… but otherwise the news is good
• What to make of the Yowie?
• Rilla Martin’s 1964 photo of the ‘Ozenkadnook tiger’

And for previous Tet Zoo articles on cats worldwide, see…

• Belated welcome to a ‘new’ clouded leopard.. named in 1823
• Peter Hocking’s big cats: where are you now?
• So what was that mysterious black gracile felid?
• Pumas of South Africa, cheetahs of France, jaguars of England
• Super-size cougars
• What is the Snodland mystery cat?
• The Pogeyan, a new mystery cat
• Leopard cats: exotic and (sometimes) wild in the UK
• Big spotted pumas… Miracinonyx redux?

Refs – -

Gilroy, R. 1995. Mysterious Australia. Nexus Publications, Mapleton, Queensland.

Healy, T. & Cropper, P. 1994. Out of the Shadows: Mystery Animals of Australia. Ironbark, Chippendale, Australia.

Heuvelmans, B. 1995. On the Track of Unknown Animals. Kegan Paul International, London.

Shuker, K. P. N. 1989. Mystery Cats of the World. Robert Hale, London.

- . 1996. The Unexplained: An Illustrated Guide to the World’s Natural and Paranormal Mysteries. Carlton, London.

Williams, M. & Lang, R. 2010. Australian Big Cats: An Unnatural History of Panthers. Strange Nation, Hazelbrook, NSW.