Friday, July 21, 2017

Soaring with Sapeornis

Among the diverse avifauna of the Cretaceous, there were none (that we know of) quite like Sapeornis from the Early Cretaceous of China. Though a few other avialans have been described as its close relatives, including "Omnivoropteryx", "Didactylornis", and "Shenshiornis", all of them almost certainly represent additional specimens of Sapeornis itself. Most phylogenetic analyses recover Sapeornis as having been more distantly related to modern birds than Confuciusornis (which is known from the same geologic formations) was, but Sapeornis possessed a strange mosaic of features, some of which were convergently similar to those of modern birds.

Like modern birds, Sapeornis had a reduced third finger that lacked a claw, unlike Confuciusornis, which retained claws on all three of its fingers. Sapeornis was also similar to modern birds in having a backwards-facing first toe that would have allowed it to easily grip a perch, whereas Confuciusornis and other avialans more distantly related to modern birds had a first toe that pointed inward at most. However, unlike Confuciusornis and modern birds, but similar to earlier paravians such as Anchiornis and Archaeopteryx, Sapeornis lacked a bony sternum (breastbone), and may not even have had a cartilaginous one. The sternum anchors many of the chest muscles used for flapping in modern birds, so this suggests that Sapeornis was not a very strong flapper.

A well-preserved specimen of Sapeornis showing the long wing feathers, from Serrano and Chiappe (2017).

Sapeornis was very large for a Mesozoic avialan, likely weighing around 1 kg, comparable in size to a common raven or a red-tailed hawk. Its forelimbs were very long and had long wing feathers attached to them, giving it an estimated wingspan of more than a meter wide, as well as a wing planform very unlike the relatively short, broad wings of most other Mesozoic avialans. So Sapeornis could likely perch skillfully with its reversed first toe, and it likely flew from perch to perch, as indicated by the reduced grasping ability in its hands and its extremely long wings. Yet its anatomy also suggests that couldn't flap its wings very well.

The size of Sapeornis compared to a human, by Matt Martyniuk, licensed.

Perhaps this apparent paradox could be explained if Sapeornis only did quick bursts of flapping flight to get from tree to tree, but such a lifestyle is more common among birds with short, broad wings, more similar to those of other Mesozoic avialans. Another possibility is that Sapeornis was specialized for soaring, given that soaring flight doesn't require a whole lot of flapping. Extant soaring birds tend to have very long wings, similar to Sapeornis, and some previous studies have certainly concluded on anatomical grounds that Sapeornis could likely soar. Until recently, however, detailed investigation into the aerodynamics of Sapeornis had not been carried out to further test these conclusions.

In a newly-published study, Francisco J. Serrano and Luis M. Chiappe used multiple lines of evidence to infer and model the flight style of Sapeornis. First, they found that the size of the deltopectoral crest (a ridge on the humerus, or upper arm bone, that helps anchor the chest muscles) is negatively correlated with flapping frequency in modern birds. In other words, birds that flap less frequently (such as soaring birds) have a larger deltopectoral crest. Sapeornis had a sizeable deltopectoral crest relative to its body size. Score one for soaring.

The forelimb of Sapeornis showing its large deltopectoral crest (DPC), from Serrano and Chiappe (2017).

Next, they used computational modeling to calculate the amount of power necessary for Sapeornis to support itself using sustained flapping as well as the amount of power it could actually generate. These results showed that it would be extremely inefficient for Sapeornis to use sustained flapping flight. This is again similar to extant soaring birds such as hawks and ravens. These birds are capable of sustained flapping, but it is energetically costly for them to do so, and as such they prefer to soar when they can. Score another for soaring.

In addition, Serrano and Chiappe estimated the wing aspect ratio (the length of the wing relative to the width of its chord) of Sapeornis using both multivariate equations as well as direct measurements of reconstructed wing area. In both cases, the wing aspect ratio of Sapeornis plotted among soaring birds, and specifically among thermal soarers such as vultures that primarily soar on columns of warm air rising off the land. This makes sense. There are other soaring birds that are dynamic soarers, specialized for soaring at high speeds over the ocean by exploiting differences in wind velocity at different altitudes. Most of these species are seabirds with narrow, pointed wings. Given that fossils of Sapeornis were preserved in inland lakes near forested environment, it would have been a big surprise if it was found to have been a dynamic soarer!

The reconstructed wing planform of Sapeornis, from Serrano and Chiappe (2017).

All of these features are consistent with Sapeornis having been a thermal soarer, but could it, in fact, soar? Thermal soarers need to be able to make tight turns so that they remain near the center of a thermal where lift is the strongest. Using their computational models, Serrano and Chiappe were also able to calculate the turning radius and minimum sinking speed of Sapeornis. They found that the smaller specimens of Sapeornis could have had a turning radii comparable to those of turkey vultures, some of the most masterful extant thermal soarers. Larger specimens had turning radii closer to those of white storks, which are also capable soarers.

Meanwhile, the large wings of Sapeornis would have been more than enough to slow its sinking speed so that it could be offset by the typical speed of rising air generated by thermals, and Sapeornis is also known to have had several large feathers on its tail, which would have given it even more lift. (In contrast, Confuciusornis had a single pair of long tail feathers at most.) Some specimens of Sapeornis preserve large feathers on the feet, which weren't incorporated into this analysis. I wonder what aerodynamic function, if any, they might have had.

Diagrams showing how Sapeornis likely soared and a graph comparing its estimated turning radii and sinking speeds to those of extant soaring birds, from Serrano and Chiappe (2017).

Thermal soaring also meshes well with the likely ecology of Sapeornis. The shape of its teeth indicate that it was most likely primarily herbivorous, and seeds have been found as its gut contents, suggesting perhaps that it ate fruit. Herbivory being a relatively low-energy diet, Sapeornis would benefited from the low-energy flight style of thermal soaring. Serrano and Chiappe suggest that good ecological analogues for Sapeornis are the screamers, a group of bizarre South American waterfowl that grow daggers on their wings, given that screamers are among the few herbivorous extant birds that both soar and perch in trees. (Though I personally suspect that Sapeornis was less terrestrial than screamers, considering the relative lengths of its forelimbs and hindlimbs as well as its large, curved foot claws.)

On the whole, this study does a convincing job of showing that both the anatomy and biomechanics of Sapeornis support thermal soaring as its main flight style, providing a plausible explanation for its unusual combination of aerially-adapted characteristics with the lack of flapping adaptations. It's easy to envision Mesozoic avialans as mere intermediates, gaining progressively more impressive flying abilities as they approached modern birds. However, the specialized soaring adaptations of Sapeornis, so far unique among Early Cretaceous avialans, set a good example in showing that many of them forged their own innovations that were not directly inherited by the line ancestral to extant birds.

I look forward to seeing this kind of multi-pronged approach applied to studying the locomotion of other extinct flying dinosaurs. Perhaps we can get an enantiornithine next...? ;)

Reference: Serrano, F.J. and L.M. Chiappe. 2017. Aerodynamic modelling of a Cretaceous bird reveals thermal soaring capabilities during early avian evolution. Journal of the Royal Society Interface 14: 20170182. doi: 10.1098/rsif.2017.0182

Saturday, July 8, 2017

Bristol Zoo, This Time with Animatronic Ornithodirans

Bristol Zoo is currently host to one of those temporary animatronic dinosaur exhibits that show up at zoos every so often. Though I didn't visit with the express goal of seeing it, I go to the Bristol Zoo often enough that running into it was almost inevitable.

My general impression is that many zoo aficionados view these animatronic exhibits with disdain, considering them little more than a cheap way to increase zoo attendance, especially when they take up space that could be used to exhibit more real live animals. I certainly sympathize with the view that the live animals should be the main draw of a zoo, but I also appreciate (and encourage) the integration of paleontology and evolutionary biology into a zoo setting. That being said, what little I know about these animatronic dinosaur exhibits suggests that their educational value is frequently lacking.

The first animatronic I came across was not a dinosaur, but this Quetzalcoatlus and its young. They're not flawless (perhaps most notably, pterosaurs are thought to have buried their eggs rather than constructing bird-like nests), but they are surprisingly not terrible, even having a covering of filamentous integument.

The accompanying sign, however, was a complete mess.
  • It uses a... very, very loose translation of "Quetzalcoatl".
  • It incorrectly identifies Quetzalcoatlus as a dinosaur.
  • It wrongly attributes the locale of Quetzalcoatlus to Mexico. (All definite specimens have been found in the U.S.)
  • It uses an outdated mass estimate. (More recent estimates put Quetzalcoatlus over 200 kg).
  • It implies that Quetzalcoatlus fed primarily on fish. (Azhdarchid pterosaurs almost certainly hunted on land. I could envision them foraging in shallow water occasionally, which isn't too different from stalking over the ground, but they were unlikely to have been specialized piscivores.)
  • That silhouette is awful.
It would be quicker to list what this sign doesn't get wrong.

This Stegoceras tried to give me the side-eye.

Unexpectedly, this Pachyrhinosaurus correctly had clawless outer fingers.

This Chasmosaurus, however, was not so fortunate, and even has its ear opening in the wrong position.

Considering that Chasmosaurus lived a few million years prior to Tyrannosaurus, the only way the former would be on the latter's menu would be if one of them was a time traveler.

Their obligatory Tyrannosaurus. It's not great, but I've seen worse. Largely unremarkable.

When I heard that they had a Utahraptor, I feared the worst. However, it was surprisingly one of the less-terrible animatronics. It still looks a little "gorilla suited" (especially when you see it up close), but the base model is not bad (even having non-pronated hands) and the extent of the plumage is fairly plausible.

Implying that most Mesozoic dinosaurs were cold-blooded though... Uh, no.

This Dilophosaurus could spit "venom" (water) at visitors. Groan.

At least the sign makes it clear that there is no evidence for such behavior.

All in all, it could have been worse, but the signage especially could use some improvement.

Naturally, the actual dinosaurs at the zoo were far more interesting. Here are some marbled teal.

Some greater flamingos, pied avocets, and a common redshank.

A white-winged duck.

A Visayan hornbill.

A Victoria crowned pigeon with a crested partridge below.

Funnily, one of the signs signalling the presence of the animatronics was right next to the Sumatran laughing thrush exhibit. I'd like to think that this was intentional.

A red-billed leiothrix.

I was fortunate enough to see this Palawan peacock pheasant displaying to a female. Unfortunately, he wasn't facing me. I don't make a very good female peacock pheasant.

The female, however, didn't appear to be particularly impressed and soon wandered off. Here is the male on his lonesome (with a wonga pigeon in the background).

A European turtle dove getting some shut-eye.

An Inca tern.

A wild Eurasian jackdaw getting rid of a molted feather.

A humorous sign at the exit of the walkthrough lorikeet aviary (which I hadn't been to on previous visits).

The non-avian residents of the zoo gave a good showing as well. The red panda was out and about.

A yellow mongoose.

Actually got an acceptable picture of a Turkish spiny mouse. It was sitting completely still under decent lighting (by nocturnal house standards). I probably won't get a better photo of one than this!

A Henkel's leaf-tailed gecko

A Lord Howe Island stick insect.

Some purple jewel beetles.

A blue tree monitor.

A Pearse's mudskipper. I see mudskippers more often out of the water than in it, so I took the chance to get this shot.

To accompany the animatronics, the gift shop had been stocked with some dinosaur-themed products. At least the title of this one is honest.

Pterosaurs mistaken for dinosaurs again.

There was also this book. Many of its illustrations look awfully familiar...

Saturday, June 17, 2017

From One Diver to Another: There's Something Loony about Petralca

First things first: we now have approximately half of a baby Mesozoic maniraptor preserved in amber. What a time to be alive.

Onward to the main subject of this post. The seabird Petralca from the Miocene of Austria was first described in 1987 as an auk. However, this identification was (reportedly*) not justified by any anatomical observations and other researchers have subsequently suggested that Petralca may instead belong to a different group of diving birds, the loons (or divers, if you're British). Saying anything conclusive regarding Petralca has been difficult though, given that the holotype is not particularly well preserved. Some of the bones associated with the specimen are not even preserved directly, only evident as impressions.

*I cannot confirm this for myself, as the original description is in German.

The holotype of Petralca, from Göhlich and Mayr (in press).

To uncover more information about the specimen, paleontologists Ursula B. Göhlich and Gerald Mayr initiated further preparation of the fossil as well as casting of the preserved bone impressions. Armed with the new data they collected from these ventures, they were able to better compare the skeleton of Petralca to those of definite fossil loons as well as extant loons and auks.

There's no use beating around the bush: they found that Petralca is a loon. Every available skeletal element in Petralca that could be compared to those of loons and auks was more similar to those of loons. Of particular note is that the radiale (one of the wrist bones) of Petralca has a deep and prominent notch, which is a very distinctive feature of extant loons, but is absent in auks.

Comparison between the radiale of Petralca (B), a red throated loon (Gavia stellata, C), and a razorbill (an auk, Alca torda, D), from Göhlich and Mayr (in press).

In addition to clearing up its phylogenetic affinities, this reassessment of Petralca also provides clues to how it lived. Extant loons can swim quickly underwater by propelling themselves with their feet, whereas previously known early Miocene loons, such as Colymboides minutus, don't appear to have been so specialized for diving. The humerus of Petralca, however, had very thick bone walls, which is characteristic of diving birds (including both extant loons and auks). It looks like Petralca truly lived up to its claim as a diver.

Reference: Göhlich, U.B. and G. Mayr. In press. The alleged early Miocene auk Petralca austriaca is a loon (Aves, Gaviiformes): restudy of a controversial fossil bird. Historical Biology in press. doi: 10.1080/08912963.2017.1333610

Tuesday, June 6, 2017

A Small Basal Paravian Not Preserved as Roadkill? Enter Liaoningvenator!

Near the base of Paraves where we once had only Archaeopteryx as a model for the last common ancestor of modern birds and dromaeosaurids, we now know of a plethora of protobirds that likely occupy the same general region of theropod phylogeny. The Late Jurassic Tiaojishan Formation in northeastern China has proven particularly productive in this regard, its fine-grained lake deposits playing host to Pedopenna, Anchiornis, Xiaotingia, Eosinopteryx, and Aurornis, with rumors of more to come.

Many of these early paravians are known from essentially complete remains, have their soft tissues preserved in exquisite detail, and, in the case of Anchiornis, are represented by hundreds of specimens, to the point where their integument, musculature, and even coloration can be restored with unprecedented accuracy. On the flip side, however, these specimens tend to be preserved as flattened corpses similar to roadkill, making aspects of their skeletal anatomy challenging to interpret.

Another geological formation exposed in northeastern China, the Early Cretaceous Yixian Formation, is also known for lake deposits containing "roadkill" fossil specimens with finely-preserved soft tissues. However, the Yixian has other fossil beds that were formed by volcanic ash, and though the fossils found in these generally lack obvious soft tissue structures, they can preserve complete skeletons in three dimensions. A newly-named basal paravian has been recovered from these Yixian ash beds, making it a potentially quite significant find.

The holotype of Liaoningvenator, from Shen et al. (2017).

Liaoningvenator curriei is known from a nearly complete specimen, and at a glance it has some striking features. It looks extremely leggy, with very long lower legs and and feet for its size. Based on rough measurements done in ImageJ, the longest metatarsal is around 64% the length of the tibia, comparable to the ratio seen in some cusorial noasaurids such as Elaphrosaurus. Its forelimbs, conversely, are quite short, the humerus being less than 60% the length of the femur. (Compare Jinfengopteryx, another short-armed basal paravian, in which this figure is around 70%.)

The tail also looks unusually short and the life restoration provided in the paper appears to take this at face value, depicting the preserved length of the tail as its total length in life. The tail as preserved looks truncated, at least to my eye, so I, for one, am skeptical of this interpretation. (The authors do not comment on this issue one way or another.) Even so, it may not be farfetched to suggest that even the complete tail of Liaoningvenator was fairly short, considering that the phylogenetic analysis in the description finds it to be the sister taxon of Eosinopteryx, which has a relatively short tail with only 20 tail vertebrae in total. (For comparison, Anchiornis and Aurornis both have around 30 tail vertebrae.)

Speaking of phylogenetic affinities, the description recovers Liaoningvenator and Eosinopteryx as part of a clade of basal troodontids along with Anchiornis and Xiaotingia. If these findings are valid, Liaoningvenator would be the geologically youngest known member of this basal clade. It would also be the largest member of the group by far: using the Christiansen and Fariña (2004) method of estimating theropod body mass by femur length, Liaoningvenator is predicted to have weighed around 2 kg, whereas its Tiaojishan brethren have all been estimated as less than 1 kg. (On top of that, bone histology indicates that the holotype of Liaoningvenator was still growing at the time of death, though approaching skeletal maturity.)

That's all very fascinating if true. However, the phylogenetic relationships of these basal paravians are notoriously difficult to figure out. Anchiornis (the best-studied Tiaojishan paravian) has bounced between being an avialan, a troodontid, and neither ever since its initial description. In addition, it is not clear whether all of these protobirds really do clade together to the exclusion of other paravians. Perhaps we should also expect a tumultuous phylogenetic future for Liaoningvenator.

Alternative phylogenetic topologies for Paraves recovered by recent studies, based on Shen et al. (2017), Cau et al. (2015), Brusatte et al. (2014), and Foth et al. (2014).

I would be remiss if I neglected to mention that Liaoningvenator is not the only small basal paravian known from a complete, three-dimensionally preserved specimen. Mei is known from two such specimens, also found in ash beds from the Yixian Formation. Infamously, both of these specimens have been preserved in what appears to be a sleeping posture, which has led to jokes that Mei spent all of its time sleeping, despite efforts by paleoartists to depict alternative behaviors. Unlike Liaoningvenator, however, Mei has never been considered a close relative of the Tiaojishan "problem paravians", and is uniformly recovered as a troodontid.

Reference: Shen, C., B. Zhao, C. Gao, J. Lü, and M. Kundrát. 2017. A new troodontid dinosaur (Liaoningvenator curriei gen. et sp. nov.) from the Early Cretaceous Yixian Formation in western Liaoning Province. Acta Geoscientica Sinica 38: 359-371. doi: 10.3975/cagsb.2017.03.06

Sunday, June 4, 2017

ProgPal 2017 and New Walk Museum

Being in the UK opens up access to a whole host of paleontological conferences that I might not have considered attending before. Among these is Progressive Palaeontology (ProgPal), a conference run specifically by grad students for grad students.

This year's ProgPal was held in the city of Leicester. The first afternoon of the conference was host to a couple of workshops, but the first event I attended was the icebreaker later in the evening, which took place at the New Walk Museum.

The paleontology exhibit at this museum is particularly rich in Jurassic marine fossils. Here are the magical jaws of Liopleurodon, poised to clamp down on the head and neck of Muraenosaurus.

As is also visible in the previous photo, many of the fossils were accompanied by small models representing life restorations.

Direct evidence that plesiosaurs ended up on the menu of other sea creatures, in the form of bite marks on some of their bones.

The skull of Rhomaleosaurus.

An ichthyosaur showing off a massive sclerotic ring.

The New Walk Museum is home to some newly-discovered marine reptiles, such as this currently unnamed plesiosaur.

Here is Wahlisaurus, an ichthyosaur that was only described last year.

Naturally, there are also a few dinosaurs on display, including this model of a Neovenator skeleton.

The centerpiece of the hall is the sauropod Cetiosaurus.

Some Ediacaran fossils get in on the act, including several specimens of Charnia.

This is said to be the only solo portrait that David Attenborough has agreed to sit.

The museum has some interesting osteological material of extant species as well, such as this skeleton of a tree kangaroo.

The skull of an Indian (or Ganges) softshell turtle.

The main events of the conference, however, happened on the second day. These were, naturally, the talks and poster presentations. I didn't present anything, given that I had barely begun to do my Master's research by the time abstracts were due. Next year, perhaps.

Even so, the Bristol contingent was well represented (unsurprisingly, considering the size of our research group), accounting for around a quarter of the total number of both attendees and presentations given. (Yes, I counted.) Dare I say, this may have been a double-edged sword in some ways; I noticed that many of us Bristolians (myself included) tended to cluster together during free periods rather than taking the time to meet new people. Nonetheless, I did have quite a few worthwhile interactions with some non-Bristolian delegates, such as meeting with future collaborator Juan Benito Moreno for the first time. I also received some (positive) attention for my continuous livetweeting of the conference talks. (Livetweeting at SVP has given me plenty of opportunities for practice.)

Most of the talks were livesteamed and can be viewed here (though the sound appears to be out of sync towards the end of the recording). Institutional loyalties aside, my favorite talks include (in order of presentation):
  • Virginia Harvey's talk on using ancient collagen to study the fauna of the Cayman Islands (which deservedly went on to win best talk)
  • Alessandro Chiarenza's talk on modelling the ecological niches of dinosaurs in the latest Cretaceous of North America
  • David Marshall's talk on the paleoecology of Acutiramus (a Bristol talk, but the fact that he had to give a hand-drawn presentation due to spending his weekend trying to salvage a vandalized fossil site and still knocked it out of the park means he more than deserves a mention)
  • Andrew Jones's talk on phytosaur phylogeny (I especially liked his slide designs)
  • Alexander Askew's talk on Middle Devonian depositional dynamics in Spain (his declaration that "animal fossils are useless" doubtless being memorable to many)

On the whole, I had a great time at ProgPal and found it to be very effective at what it sets out to do (providing a relatively laid-back environment for early-career paleontologists to share their research). I intend to be back next year.