Battle of the Monofenestrata (Stem Pterodactyloidea)

A new paper on pterosaur biogeography (Upchurch et al. 2014) includes a family tree from Andres et al. (2014) that puts a new spin on things. That’s the tree that introduced us to Kryptodrakon, which turns out to be just another Sericipterus (a large, gracile dorygnathid), found in the same locality. As you’ll recall, when that “long” metacarpal of Kryptodrakon was placed on the much larger bauplan of gracile Sericipterus, it wasn’t so long anymore.

More to the headline
The Andres tree demotes Darwinopterus to just another wukongopterid. And that’s to Andres credit! So the combatants for pterosaur tree domination now line up on three fronts, with shifting allegiances.

From their Upchurch et al. 2014 abstract: Although sampling biases and taxonomic problems might have artificially elevated the occurrence of sympatry, we argue that our results probably reflect a genuine biogeographical signal. We propose a novel model to explain pterosaurian distributions: pterosaurs underwent a series of ‘sweep-stakes’ dispersal events (across oceanic barriers in most cases), resulting in the founding of sympatric clusters of taxa.

I have no problem with that scenario or any other ‘rush and rest’ distribution system, but it needs to be built on a base of good phylogeny.

Triassic Pteros
In the Andres tree, their basal split is between Triassic pterosaurs and all others with Preondactylus + Austriadactlyus basal to the former group and DimorphodonParapsicephalus + Campylognathoides basal to the latter. Good golly, that’s a mixed bag already!

No outgroups were mentioned, but earlier trees by Andres used Euparkeria, an archosaur with no pterosaur affinities. And without a good foundation, how well can the house stand?

There aren’t many similarities between long-snouted Parapsicephalus and tall snouted Dimorphodon. And how does Campylognathoides match to these two? Not well. There are more parsimonious sister taxa out there.

The oddest and most controversial aspect of Andres’ tree is the nesting of Anurognathidae far from Peteinosaurus and Dimorphodon. He derives anurognathids like Dendrorhynchoides from Changchengopterus (but which one?), wukongopterids and Sordes (in order of increasing distance), at the base of the Pterodactyloidea, with Krypodrakon at the base.

Andres nests Anurognathidae within the Monofenestrata because he finds no bone to divide the naris from the antorbital fenestra in anurognathids (following the Bennett model) and they have short tails, like pterodactyloids. This needs to be tested with reconstructions. My tests don’t confirm Bennett’s model. I’d like to see someone else step up to the plate and see what they find. We need third and fourth party input.

There remains some traditional confusion about the sharp tooth and toothless pterodactyloids. In the Andres tree toothy Haopterus is basal to toothless, sharp-snouted Pteranodon + Nyctosaurus on one clade that produces more toothy ornithocheirids. And Haopterus is also basal to another toothless (except for dsungaripterids!), sharp-snouted clade, the tapejaridae + dsungaripteridae + azhdarchidae. Eopteranodon is separated from Eoazhdarcho. Noripterus is a sister to Thalassodromeus and other odd pairings abound.

And when did MoganopterusFeilongus and Huanhepterus start nesting with ctenochasmatids?

Perhaps key to the problems here:
No tiny pterosaurs, other than Eudimorphodon cromptonellus, and Nemicolopterus were included. When that happens, all hellzapoppin!

And this is why I always ask my submission editors to not send my work to these chaps to be refereed. They’re not seeing the red flags.

Andres B, Clark JM, Xu X. 2014. The earliest pterodactyloid and the origin of the group. Curr Biol. 24(9):1011–1016
Upchurch P, Andres B,  Butler RJ & Barrett PM 2014. An analysis of pterosaurian biogeography: implications for the evolutionary history and fossil record quality of the first flying vertebrates, Historical Biology: An International Journal of Paleobiology, DOI: 10.1080/08912963.2014.939077

Sharovipteryx Wiki just updated

Once again, like the Pterosaur Origins Wiki page, some well-meaning, but misinformed author/expert on the Sharovipteryx Wiki page claimed that I did not observe the fossil firsthand (which is false) and that my phylogenetic analysis of Sharovipteryx (still the only one in any academic publication in the last 14 years) had less validity than what paleontologists “generally agree”. Yes, in this age of verifiable nestings, can you believe this return to the vagaries of the 1960s? (Actually I think this only occurs when my name is present).

The author/expert claimed that I am not a scientist (ignoring academic publications in 6 or 7 journals now) and put his faith in Bennett’s claim made in a popular publication that my tracings were fantasies (once again, mining the wastebasket). Yes, I made those mistakes, but the new work puts all that crap in the wastebasket, where it should stay.

Wiki is generally for information, not for casting aspersions on others. So, when an alternate and testable hypothesis is presented, it is not necessary that the author of that hypothesis be trashed. Simply present the facts. Not the bias, please.

I made changes to the Sharovipteryx Wiki page that stick to the readily observable and testable facts. Let’s see if those changes stick.

If that author/expert wants to put his faith in Chris Bennett, Lord help him. Bennett has made dozens of mistakes, including purposefully creating a fantasy (by his own admission) pterosaur precursor (Bennett 2008), rather than to test any of the hundreds of currently known reptile candidates in phylogenetic analysis, as I have here. And 2008 Bennett had a short list provided by Peters (2000), which he ignored. I tested his 1996 paper by adding a few taxa. Turnabout would have been very welcome.

We’re all guilty. Let’s move forward people. Please, use the latest information and keep the focus on the taxa, not the person. I put all my data in viewable, testable photos and am more than happy to make corrections when made available.


Bottom Line
Sharovipteryx is a complete fossil with many uncontroversial traits shared with Cosesaurus and pterosaurs. Those traits are going unpromoted in Wiki and I think it’s because some people think I’ve poisoned the well in publishing on it without having a PhD. They have to tip-toe around my peer-reviewed publications and they have to trash me because if they started listed characters, they’d soon find out what anyone can find out. Perhaps that’s why no one had published another analysis of Sharovipteryx in the last 14 years. And it’s ripe for a revision because I made several mistakes with it, even firsthand.

Remember, Hone and Benton (2007, 2008) tossed Sharovipteryx out in their search for a pterosaur precursor. Same thinking. Same result.

If you’re thinking of Senter’s (2003) dissertation (which the Wiki author/expert cited), in which he nested Sharovipteryx with Cosesaurus, but pterosaurs with Scleromochlus, take a good look at his scorings. He gave Scleromochlus a sternal complex and a long lateral pedal digit, both of which are absent on it and any sister taxa — among dozens of other rookie mistakes.

Bennett S.C. 1996. The phylogenetic position of the Pterosauria within the Archosauromorpha. Zoo. J. Linn. Soc. 118: 261-309.
Bennett S.C. 2008. Morphological evolution of the forelimb of pterosaurs: myology and function. In: Buffetaut E. and Hone D.W.E. (Eds) – Flugsaurier: pterosaur papers in honour of Peter Wellnhofer. Zit., B28: 127-141.
Peters D. 2000. A Redescription of Four Prolacertiform Genera and Implications for Pterosaur Phylogenesis. Riv. It. Paleo. Strat. 106(3): 293-336.



Temnospondyl eyeballs


Figure 1. Antarctosuchus polygon life restoration with original eyeballs, above, and suggested revision, below. Why way did the eyeballs point? It might make more sense to have 360 degree vision than binocular vision. Just like a frog.

Figure 1. Antarctosuchus polygon life restoration with original eyeballs, above, and suggested revision, below. Why way did the eyeballs point? It might make more sense to have 360 degree vision than binocular vision. Just like a frog.

Sometimes these skull restorations just freak me out.
Sure temnospondyls had orbits on the top of their skulls, but did their eyes stare directly up from the skull with binocular vision? Or did they stare laterally with 360 degree vision, like a frog? Above both variations are shown. The original sort of freaks me out.

How about you?

And what happens at the surface when this croc-like anamniote breathes at the surface? Did it stare at the sun? Was it able to withdraw its eyeballs like frogs do?

Please, someone, fix those eyes or calm my nerves by telling me that’s the way they are supposed to be, so they don’t freak me out.

Sidor CA, Steyer JS and Hammer WR 2014. A new capitosaurid temnospondyl from the Middle Triassic Upper Fremouw Formation of Antarctica. Journal of Vertebrate Paleontology 34(3):539-548. DOI:10.1080/02724634.2013.808205
– See more at: chinleana update

It’s been awhile since I last updated
Just sent off the third of three papers. So, I’ve been busy and it has been interesting discovering things along the way. The family trees for the Pterosauria and the Reptilia will not be updated until the publication of these works. Sorry about that.

Only a few poorly preserved taxa moved.
The tree topology remains the same. The navigation bars have been updated. Should make it easier to get around, especially if you know what you’re looking for.

While we’re on the subject,
It’s been two years since Darren Naish at Tetrapod Zoology took a swipe at I don’t think he’ll notice or appreciate the changes, but I have been checking, double checking and getting new data. Not all of it is up yet (see above for the reason). As you know, I have been criticized, but I’m not married to any of this. I do put out what I’ve learned because Science, in all of its forms, needs to be tested. Are my tests correct? Maybe not. I’ve been wrong thousands of times. Getting it right in the end is my only goal here. At present things seem better than they were. More parsimonious. Sister taxa show gradual evolutionary changes, etc.

My posts have not been produced daily lately, like they were for the first 1000 or so. Whether I’m running out of steam, finding fewer things to talk about in this realm, or there has been a reduced flow of fun new paleo-discoveries, I don’t know.

For those who are interested…
Publication doesn’t guarantee acceptance or even discussion of novel discoveries. It’s been 14 years since Cosesaurus and kin were introduced as more parsimonious ancestors than any known archosaur/archosauromorph. Yet, pterosaur workers are still putting Euparkeria into their matrices when Parasuchus comes out closer on their tree. Very odd. Almost funny if it weren’t so tragic.

Along the same lines, if you google “Pterosaur pteroid” you’ll find a listing for an Ask the Biologist answer on this subject that is surprising given that the origins of the pterosaur pteroid were published in 2002 and 2009. Experts Mark Witton and Dave Unwin in their books on pterosaurs likewise ignore published literature and report that the origin of the pteroid remains an unsolved mystery.

While I’ve gotcha…
I would like to turn you all on to a pretty sharp dinosaur YouTube video I saw recently that I’m probably the last one on the block to hear about. Surprisingly the theropods all have what seems to be an appropriate amount of feathers. Even the big ones!

It’s about an hour and eleven minutes. Not too hokey, either. Enjoy!

Figure 1. March of the Dinosaurs on YouTube. Click to Watch. 1:11:00

Figure 1. March of the Dinosaurs on YouTube. Click to Watch. 1:11:00

The most primitive Choristoderes?

are a difficult clade to figure out both inside and out.

Wikipedia reports, “Champsosaur skulls are actually very similar to lizard skulls, though heavily modified. This has led some researchers to consider champsosaurids as lepidosauromorphs. However, champsosaurs lack the complex quadrate of lepidosaurians. With features of both diapsid groups, the phylogenetic position of Choristodera is highly confused.”

Matsumoto et al. (2007) reports, “This tree confirms the monophyly of Neochoristodera (Evans and Hecht1993) including Champsosaurus, Ikechosaurus, Simoedosaurus, and Tchoiria. The relationships of the non−neochoristoderan taxa have been more controversial. In our analysis, the Jurassic Cteniogenys retains a basal position, with the Chinese Philydrosaurus (Gao and Fox 2005) one node above it.”

Note the loss of resolution at the base of their tree (Fig. 1).

Figure 1. The family tree of the Choristodera according to Matsumoto et al. 2007. Early eras added in color. Note that ancestral taxa were large and long-snouted with four temporal openings, not two.

Figure 1. Click to enlarge. The family tree of the Choristodera according to Matsumoto et al. 2007. Early eras added in color. Note that some ancestral taxa, like Diandongosuchus, were large and long-snouted. Others were small, like the BPI specimen of Youngina, but equally long snouted. So the long-snouted forms may be more primitive. Note the loss of resolution at the clade base.

The question in the Matsumoto et al. 2007 tree is what happened in pre-Jurassic times? In 2007 Youngina was indeed the best outgroup taxon, but several varieties are known and some of these have a longer rostrum. Unfortunately these were overlooked as Matsumoto et al. focused on known choristoderes. In 2012 the large younginidDiandongosuchusbecame known. While it nests at the base of the parasuchia, it also nests at the bases of the Choristodera and Chanaresuchia. Yes, it has an antorbital fenestra, and so do some Youngina specimens. It is possible that the Choristodera secondarily lost their aof. Or they never had one.

In any case, the large reptile tree recovered a similar basal split between the large and small choristoderes, but with complete resolution among the five small ones and three large ones. More precision in the character scoring of the various Youngina specimens should add clues to this mystery. Don’t discount those subtle variations!

Matsumoto R, Evans SE and Manabe M 2007. The choristoderan Monjurosuchus form the Early Cretaceous of Japan. Acta Palaeontologica Polonica 52(2):329-350.


Another lateral temporal fenestra!

Figure 1. Emeroleter skull showing lateral temporal fenestra, as also shown in its sisters Macroleter, Romeriscus and Lanthanosuchus.

Figure 1. Emeroleter skull showing lateral temporal fenestra, as also shown in its sisters Macroleter, Romeriscus and Lanthanosuchus. Images from Tsuji, Müller and Reisz 2012.

This looks like a regular break (Fig. 1), but Emeroleter nests between taxa that also have a lateral temporal fenestra (Fig. 2). So this could be a real lateral temporal fenestra made ragged over the last tens of millions of years.

Figure 2. Macroleter, Emeroleter, Romeriscus and Lanthanosuchus in phylogenetic order and to scale. All have a lateral temporal fenestra.

Figure 2. Macroleter, Emeroleter, Romeriscus and Lanthanosuchus in phylogenetic order and to scale. All have a lateral temporal fenestral, even if small. This is a better evolutionary sequence leading to Lanthanosuchus than Acleistorhinus can provide.

At this stage in lepidosaurormorph evolution, taxa were just beginning to experiment with the lateral temporal fenestra, which was “here to stay” in owenettids, like Sauropareion.

However, the last time we see the lower temporal bar (jugal/ quadratojugal connection) is right here (Fig. 2) until these two bones are reconnected again in basal sphenodontians, drepanosaurs and fenestrasaurs.

DeBraga and Reisz (1996) and Cisneros (2004) reported that Acleistorhinus was the sister to Lanthanosuchus. However, the large reptile tree indicates a better match with Romeriscus, Emeroleter and Macroleter. Acleistorhinus makes a better sister to the turtle-like millerettid, Eunotosaurus.

Tsuji et al. 2012 reported, “A clade consisting of the nycteroleters and pareiasaurs, here termed Pareiasauromorpha, is supported by both methods.” Unfortunately, it is not supported by the large reptile tree, unless all subsequent taxa that have these taxa at their base are also called pareiasauromorphs, and that would include all turtles, lizards and snakes, and Lanthanosuchus, which Tsuji et al. 2102 also nested with Acleistorhinus far from Emeroleter. 

Cisneros et al 2004. A procolophonid reptile with temporal fenestration from the Middle Triassic of Brazil. Proceedings of the Royal Society London B (2004) 271, 1541–1546 
deBraga M and Reisz RR 1996. The Early Permian reptile Acleistorhinus pteroticus and its phylogenetic position. Journal of Vertebrate Paleontology 16(3): 384–395. doi:10.1080/02724634.1996.10011328.
Modesto SP, Damiana RJ and Sues H-D 2002. A reappraisal of Coletta seca, a basal procolophonid reptile from the lower Triassic of South Africa. Palaentology 45(5):883-895.
Tsuji, Müller and Reisz 2012. Anatomy of Emeroleter levis and the Phylogeny of the Nycteroleter Parareptiles. Journal of Vertebrate Paleontology 32 (1): 45-67. doi:10.1080/02724634.2012.626004.



Scandensia is a key taxon at the base of the Squamata

Changes made December 04, 2014 to renest Scandensia basal to the Squamates, not the Lepidosauria.

From the Early Cretaceous of Spain, 
Scandensia has been described as “an unusual lizard” and “enigmatic.”

From Bolet and Evans 2011
“The original description of Scandensia (Evans and Barbadillo 1998) established that it was a squamate (e.g. jaw and tooth morphology, emarginated scapulocoracoid fenestrate clavicle, absence of gastralia) and this is confirmed by the new specimen (e.g. co-ossification of the pelvic bones, pubic morphology). The first phylogenetic analysis (Evans and Barbadillo 1998), using PAUP and a modified version of the Estes et al. (1988) matrix, put Scandensia in a basal position on the squamate stem.”

Figure 1. Scandensia a basal lepidosaur

Figure 1. Scandensia a basalmost squamate.

The present analysis confirms that nesting
Here Scandensia nests at the base of the Squamata, which means it was relic from the Early Permian living in the Early Cretaceous. Tendril-like fingers and toes mark it as arboreal.

The misnamed “Langobardisaurus” rossii, MFSN 19235, is the current larger outgroup taxon.

Figure 2. Langobardisaurus(?) rossii (MFSN 19235) reconstructed. Here it nests between basal sphenodontids and basal tritosaurs + squamates.

Figure 2. Langobardisaurus(?) rossii (MFSN 19235) reconstructed. Here it nests between basal sphenodontids and basal tritosaurs + squamates. Probably not a climber, but a digger.

There’s a whole other world of lepidosaurs being discovered that don’t fit into the established clades like the Sphenodontia and the Squamata. The Tritosauria are among them, but other clades, like the one that includes Scandensia and MFSN 19235, are out there and more should be expected.

Evans SE and Barbadillo LJ 1998. An unusual lizard (Reptilia: Squamata) from the Early Cretaceous of Las Hoyas, Spain. Zoological Journal of the Linnean Society 124:235-265.
Bolet A and Evans SE 2011. New material on the enigmatic Scandensia, an Early Cretaceous lizard from the Iberian Peninsula. Special Papers in Palaeontology 86:99-108.