Tetrapodophis – new four legged very basal, very tiny snake – part 2

Earlier we looked at the skull of Tetrapodophis (Martill et al. 2015), a four-legged very tiny snake.

Figure 1. Tetrapodophis nests at the base of the clade of snakes in the large reptile tree.

Figure 1. Tetrapodophis nests at the base of the clade of snakes in the large reptile tree. Note, the burrowing snakes are not basal in this tree. Rather these very specialized snakes are quite derived. There are more proto-snakes and basal snakes known, so this tree should be considered in that light.

A phylogenetic analysis nested Tetrapodophis at the base of all snakes (Fig. 1).

Figure 2. The skulls of pre-snakes, Tetrapodophis and snakes compared. The orbits move foreword. The jaw muscles enlarge. The upper temporal arch disappears.

Figure 2. The skulls of pre-snakes, Tetrapodophis and snakes compared. The orbits move foreword. The jaw muscles enlarge. The upper temporal arch disappears.

National Geographic featured several Dave Martill quotes. Here are a few:

“And then, if my jaw hadn’t already dropped enough, it dropped right to the floor,” says Martill. The little creature had a pair of hind legs. “I thought: bloody hell! And I looked closer and the little label said: Unknown fossil. Understatement!”

“I looked even closer—and my jaw was already on the floor by now—and I saw that it had tiny little front legs!” he says.

“But no snake has ever been found with four legs. This is a once-in-a-lifetime discovery.”

“This little animal is the Archaeopteryx of the squamate world,” he says.

Martill thinks that Tetrapodophis killed its prey by constriction, like many modern snakes do. “Why else have a really long body?” he says.

Martill thinks that the snake may have used these “strange, spoon-shaped feet” to restrain struggling prey—or maybe mates.

There was a bit of controversy raised about this specimen. Read about it here at Forbes.com. It also includes an illustration of Tetrapodophis wrapped around a mouse-like mammal.

Likely a Crato Formation fossil
Martill et al. thought the Tetrapodophis substrate was from the Crato Formation. Wikipeidea reports, “The Crato Formation is a geologic formation of Early Cretaceous age in northeastern Brazil‘s Araripe Basin. It is an important Lagerstätte (undisturbed fossil accumulation) for palaeontologists. The strata were laid down mostly during the early Albian age, about 108 million years ago, in a shallow inland sea. At that time, the South Atlantic was opening up in a long narrow shallow sea.”

Nevertheless,
Martill et al. consider Tetrapodophis closer to burrowing snakes, not aquatic ones. Distinct from its sea-going predecessors, Tetrapodphis had a longer torso than tail, like living snakes do. It also had a single row of belly scales, like snakes, preserved as soft tissue impressions.

Perhaps,
owing to its small size, Tetrapodophis had returned to the land, or shallows grading to swamps.

Video link 1. Dave Martill describing from large photos Tetrapodophis. Just a few minutes long.

Video link 1. Dave Martill describing from large photos Tetrapodophis. Just a few minutes long.

Finally,
here’s Dave Martill in a video describing Tetrapodophis. Click to play.

References
Martill DM, Tischlinger H and Longrich NR 2015. A four-legged snake from the Early Cretaceous of Gondwana. Science 349 (6246): 416-419. DOI: 10.1126/science.aaa9208

Novel insights – part 1

Not much news lately,
so a bit of a review in the current storms of controversy and disparagement.

In the last four years
adding species- and specimen-based taxa to the large reptile tree and large pterosaur tree and creating reconstructions, at times using DGS, have provided a rich trove of novel insights into reptile evolution heretofore (and too often currently) unnoticed, overlooked and ignored. Of course, these all need to be tested in independent studies using similar taxon lists along with any novel list of character traits exceeding 150-200 in number.

Amniota

  1. Initial split of the Amniota into Lepidosauromorpha and Archosauromorpha clades. That means Amniota = Reptilia.
  2. Gephyrostegus bohemicus is a sister to the last common Viséan (or earlier) ancestor of all Amniotes. It lacks traditional amniote skeletal traits, but lacks posterior dorsal ribs, creating a larger volume for gravid females to hold larger eggs, a deeper pelvic opening and unfused pelvic elements.
  3. Proximal outgroup taxa to the Amniota include sisters to Silvanerpeton, Utegenia and members of the Seymouriamorpha in order of increasing distance.
  4. As in many prior studies, phylogenetic miniaturization is key to the origin of several clades.

Lepidosauromorpha

  1. Basal lepidosauromorphs include the clade of Urumqia, Brukterepeton and Thuringothyris. Some of these were formerly considered anamniotes.
  2. Captorhinomorph sister taxa include Cephalerpeton, Reiszhorhinus, Concordia and Romeria primus. Romeria texana is a basal captorhinomorph.
  3. A sister to Saurorictus is basal to all remaining lepidosauromorphs, Diadectormorpha + Millerettidae.
  4. Diadectomorphs are lepidosauromorph reptiles.
  5. Procolophon and kin are sisters to diadectomorphs like Oradectes, Silvadectes and Diadectes. A sister to Orobates is their last common ancestor.
  6. Colobomycter is a basal procolophonid.
  7. Tetraceratops is a sister to Tseajaia and Limnoscelis and these three are sisters to the Diadectes + Procolophon clade.
  8. Caseasauria are millerettids, not synapsids and caseasauria is a sister clade to Feeserpeton + Australothyris + Eunotosaurus + Acleistorhinus + Delorhynchus.
  9. Bolosaurids are also millerettids  and are basal to the Stephanospondylus clade.
  10. Stephanospondylus is basal to the pariasaur + turtle clade.
  11. Sclerosaurus is basal to the turtle clade.
  12. ElginiaMeolania nest as basalmost turtles along with Proganochelys.
  13. Odontochelys nests with Trionyx, a soft-shell turtle. Skull emargination and tooth loss was convergent in soft shell  and hard shell turtles.

More later.

Tetrapodophis – new four legged very basal, very tiny snake

A new paper
by Martill, Tischlinger and Longrich (2015) brings us a really tiny, new Early Cretaceous snake, Tetrapodophis amplectus (Fig. 1, BMMS BK 2-2, ), with four limbs and all of its fingers and toes. The authors suggest this basal snake and thus all snakes evolved from burrowing rather than marine ancestors in accord with the  Longrich, Bullar and Gauthier (2012) assessment of another tiny snake, Coniophis, which is known from only a few skull parts. (Also see below.)

Unfortunately
Tetradopodophis (so far based on skull traits only) nests in the large reptile tree between Adriosaurus + Pontosaurus and DinilysiaPachyrhachis + Boa, so an aquatic origin is recovered from the cladogram despite the extremely tiny size of Tetrapodophis (skull length about 1 cm, total length about 16 cm). Martill et al. used mosasaurs and several incomplete taxa (Eophis, Diablophis, Portugalophis and Parviraptor, none included in the large reptile tree) for outgroups and nested Tetrapodophis as a sister to Coniophis and basal to Najash, Dinilysia and all other snakes.

The authors note, “As the only known four-legged snake, Tetrapodophis sheds light on the evolution of snakes from lizards. Tetrapodophis lacks aquatic adaptations (such as pachyostosis or a long, laterally compressed tail) and instead exhibits features of fossorial snakes and lizards: a short rostrum and elongation of the postorbital skull, a long trunk and short tail, short neural spines, and highly reduced limbs.”

I wonder if Tetrapodophis is a hatchling?
Or does it represent yet another example of phylogenetic miniaturization at the origin of a major clade? It is similar in size to Jucaraseps, a more primitive lizard with snake affinities. Tetrapodophis may be a late surviving (Early Cretaceous) very basal snake with likely origins in the Middle Jurassic.

DGS
(digital graphic segregation) was helpful in pulling out details (Fig. 1) overlooked or ignored by the original authors.

Figure1. The skull of Tetrapodophis in situ and colorized (middle) as originally interpreted (below) and reconstructed using DGS (above).

Figure1. The skull of Tetrapodophis in situ and colorized (middle) as originally interpreted (below) and reconstructed using DGS (above). I have not seen the fossil, but examination of the photograph using DGS permits more details to be identified. This image will be tested for validity Monday. Only the major bones were identified here. The skull is about 1 cm in length.

The preparator did an excellent job
on such a tiny (16 cm) specimen, unless it split naturally into part and counterpart. The specimen was in a private collection for decades before getting its museum number.

Like non-snakes,
Tetrapodophis retained a postorbital, squamosal and lacrimal. A broken jugal was also found. Palatal fangs were present along with a deep coronoid process. There is a mass at the back of the throat that makes it difficult to identify the posterior palatal bones. The authors report, BMMS BK 2-2 is distinguished from all other snakes by the following combination of characters: 160 precaudal and 112 caudal vertebrae, short neural spines, four limbs, metapodials short, penultimate phalanges hyper elongate and curved, phalangeal formula 2?-3-3-3-3? (manus) 2-3-3-3-3 (pes).”

Although DGS was able to pull lots of details out of this specimen,
don’t expect the DGS detractors to applaud this example, although It would be nice to get a tip of the hat for this one. It’s a pretty striking example and only took an hour or two to do.

Figure 2. Tiny Tetrapodophis at full scale if your monitor produces 72 dpi images (standard on many monitors).

Figure 2. Tiny Tetrapodophis at full scale if your monitor produces 72 dpi images (standard on many monitors).

This is a major find
and congratulations are due to the authors. More on this specimen in future blog posts.

References
Longrich NR, Bullar B-A S and Gauthier JA 2012. A transitional snake from the Late Cretaceous period of North America. Nature 488, 205-208.
Martill DM, Tischlinger H and Longrich NR 2015. A four-legged snake from the Early Cretaceous of Gondwana. Science 349 (6246): 416-419. DOI: 10.1126/science.aaa9208

Invagination and erosion of the turtle cranium

Turtles have no temporal fenestra,
but some of them have enlarged their jaw muscles by greatly enlarging the cranium, or by invagination of the cranium from the occiput, or both (Fig. 1). Skull temporal fenestra are important traits to categorize most reptiles, but turtles do not follow other clade morphologies. That has made turtles difficult to categorize and nest in traditional studies.

Figure 1. Macrochelys (Macroclemys) skull colorized. Most workers label the bone above the curled quadrate as a squamosal, but here it is considered a supratemporal, which has horns in basal turtles.

Figure 1. Macrochelys (Macroclemys) skull colorized. Most workers label the bone above the curled quadrate as a squamosal, but here it is considered a supratemporal, which has horns in basal turtles. This skull shows a minimum of occiput invagination, but note the great height of the cranium.

Some paleontologists
think turtles are diapsids related to placodonts, but that is not supported by the large reptile tree.

Other paleontologists
think turtles are anapsids related to Eunotosaurus, but that is not supported by the large reptile tree.

Still other paleontologists
USED to think turtles are anapsids related to pareiasaurs, and that IS supported by the large reptile tree. Basal turtles, like pareiasaurs and all basal tetrapods, have both an external (dermal) skull  surrounding and protecting the smaller internal (braincase) skull.

Basal turtles have a solid cranium – with horns!
Elginia and Meiolania are basalmost turtles they have horns and at least we know that Meiolania had a solid carapace and plastron. The outgroup, Sclerosaurus, has horns, but no shell and no broad ribs. In Meiolania the large, horned supratemporal sends a ventral process to contact the quadratojugal leaving a hole for the quadrate and stapes (ear bone). The supratemporal is a large bone in basal turtles that does not go away in derived turtles. Rather, the squamosal continues to shrink.

Figure 2. Elginia and Meiolania, two basal horned turtles without skull invagination.

Figure 2. Elginia and Meiolania, two basal horned turtles without skull invagination. In Meiolania the supratemporal sends a ventral process to contact  the quadratojugal leaving a hole for the quadrate and stapes (ear bone). The supratemporal is a large bone in basal turtles that does not go away in derived turtles. Rather, the squamosal continues to shrink.

Proganochelys (Fig. 3) has long been recognized as a basal turtle. It has no horns or skull invagination, so, in this context, it is not a basal turtle, but a transitional turtle, between horned and invaginated-skull turtles.

Figure 2. The skull of Proganochelys, a basal turtle without skull invagation and without horns.

Figure 3. The skull of Proganochelys, a basal turtle without skull invagation and without horns. Note the identification of the supratemporal on the right matching that of basal turtles like Elginia and Meiloania in figure 2.

Softshell turtles
have an invaginated cranium, no horns and sometimes reduce their bony shells. A basal turtle with teeth, Odontochelys, nests with a soft-shell turtle, Trionyx, in the large reptile tree. The cranium of Trionyx is invaginated from the occiput, creating space for large jaw muscles. The skull of Odontochelys (Fig. 7) is difficult to study with available data, but it appears to have large round holes in the crushed cranium. At least it does not appear to have the solid cranium that was illustrated originally (Fig. 7). Rather the cranium appears to be so badly crushed, even in the low resolution image available, that it may indeed have had a more fragile, less box-like, Trionyx-like cranium. I requested high rez images, but was informed that another paper focusing on the skull of Odontochelys is in progress. Looking forward to that!

Figure 3. Trionyx, a softshell turtle with bones colorized.

Figure 3. Trionyx, a softshell turtle with bones colorized.

Other tested turtles have a an increasingly invaginated cranium
Chelonia,
the sea turtle, is a basal turtle that has a rather solid skull with a little posterior invagination.

Macrochelys, the alligator snapping turtle (Fig. 1), has a deeper invagination and a much taller cranium.

Pelomedusa and Foxemys are similar to the snapping turtle, but without the grand enlargement of the cranium.

Terrapene (Fig. 5), the box turtle, has lost most of its original cranium, revealing a braincase, like a mammal, snake, amphisbaenid or bird.

Figure 4. Terrapene, the box turtle, with skull bones colorized. Note the lack of a dermal skull and the appearance of the cranial skull, the braincase.

Figure 5. Terrapene, the box turtle, with skull bones colorized. Note the lack of a dermal skull and the appearance of the cranial skull, the braincase, as in birds, mammals, snakes and amphisbaenids.

The large reptile tree
nests Kayentachelys between the soft-shell turtles, Trionyx and the hard-shelled turtles, like Chelonia (Fig. 6). Kayentachelys has a complete cranium without invagination. Separate nestings of soft-shell and hard-shell turtles with skull invagination indicate this trait was convergent, not homologous. Such a tree topology has not been recovered before, but then no prior study (that I can recall) has included Sclerosaurus, Elginia and Stephanospondylus.

Figure 1. Turtle phylogeny showing extent of horns and cranial invagination. Here the skull invagination of soft-shell turtles is convergent with that of most other turtles.

Figure 6. Turtle phylogeny showing extent of horns and cranial invagination. Here the skull invagination of soft-shell turtles is convergent with that of most other turtles.

Parts of the skull of Odontochelys cannot be accurately reconstructed with available data (Fig. 7). There are apparent temporal fenestrae in the in situ specimen exposed in dorsal view. These would ordinarily have the appearance of diapsid openings and would lend credence to the diapsid hypothesis of turtle origins. Instead, let’s wonder if these holes represent either: 1) geological erosion; or 2) erosion of the posterior cranium in spots transitional to the morphology seen in Trionyx. There’s nothing else I can say at present until better data comes along.

There is a large circular plate
(Fig. 7) in the palatal view of the smaller Odontochelys that was labeled a possible squamosal. I don’t think there is room on the skull for that bone at present. So that elliptical bone may be from elsewhere, perhaps on the carapace or plastron.

Most turtles have anterior nares.
The anterolateral placement of the large naris in Odontchelys is different from all other turtles and similar only to Elginia (Fig. 2).

Figure 7. Data and tentative interpretations of skull elements for Odontochelys. I can't make more sense than this of the bones. Sorry. Gray areas appear to represent holes in the cranium.

Figure 7. Data and tentative interpretations of skull elements for Odontochelys. I can’t make more sense than this of the bones. Sorry. Gray areas appear to represent holes in the cranium. Note the difference between the original drawing and the photo with color overlay. The skull bones of Odontochelys appear to be more fragile than boxy turtle skulls are.

The skull of Stephanospondylus (Fig. 8) is a good starting point for both pareiasaur and turtle skulls. It nests (Fig. 7) at the base of both clades.

Figure 2. Stephanospondylus skull in two views. Note the rotation of the post parietals to the dorsal skull along with the transformation of the supratemporals into small horns.

Figure 8. Stephanospondylus skull in two views. Note the rotation of the post parietals to the dorsal skull along with the transformation of the supratemporals into small horns.

Perhaps more taxa
will someday unite soft-shell turtles with hard-shell turtles, but at present, the convergence is remarkable among all turtles with an invaginated skull. With regard to Odontochelys, I think we’ll see a strong revision of the original drawing.

References
Li C, Wu X-C, Rieppel O, Wang L-T and Zhao L-J 2008. An ancestral turtle from the Late Triassic of southwestern China. Nature 456: 497-501.

Urumqia – a very basal lepidosauromorph

Urumqia liudaowanensis (Zhang et al. 1984) ~20 cm snout-vent length, Lower Permian.

Figure 1. Urumqia liudaowanensis (Zhang et al. 1984) ~20 cm snout-vent length, Late Permian.

Here’s a gephyrostegid/basal amniote/basal lepidosauromorph
you may not have heard of. (Remember lepidosauromorphs in the large reptile tree constitute about half of all amniotes). It is considered China’s oldest known tetrapod.

Urumqia liudaowanensis (Zhang et al. 1984, Fig. 1) ~20 cm snout-vent length, Late Permian Lucaogou Formation), was originally considered a discosaurid seymouriamorph. Here it nests at the base of the lepidosauromorph reptiles. Shifting Urumqia to the discosaurid seymouriamorphs adds 39 steps to the large reptile tree.

Derived from 
Gephyostegus bohemicusUrumqia was basal to Bruketererpeton, Thuringothyris, and all lepdiosaurs, turtles, diadectids, pterosaurs and other various lepidosauromorphs starting with Saurorictus and Cephalerpeton. Phylogenetically Urumqia must have made a first appearance in the Viséan (335 mya, Mississipian, Carboniferous) despite its late appearance in the Late Permian (255 mya).

Figure 1. Basal amniotes to scale. Click to enlarge.

Figure 2. Basal amniotes to scale. Click to enlarge. Urumqia nests on the right hand column with Cephalerpeton and Thuringothyris.

Distinct from G. bohemicus,
Urumqia had shorter limbs, longer (but not long) posterior dorsal ribs and a robust tail with elongate caudals. The palate included a suborbital fenestra. The cheek may have included a small lateral temporal fenestra convergent with others. The carpals and tarsals were poorly ossified.

Figure 4. Extant lizards, A. gravid, B. in the process of laying eggs, C. with egg clutch.

Figure 3. Extant lizards, A. gravid, B. in the process of laying eggs, C. with egg clutch.

Notably
the posterior dorsal ribs were much shorter than the gastralia. So the gastralia create a wide posterior torso, ideal for carrying large amniote eggs (Fig. 3), as we learned earlier.

The new topology of basal reptiles
is based on the inclusion of several more species based taxa not previously considered. This new topology show that synapsids were not the first clade to branch off. Rather all taxa closer to archosaurs (here considered the new Archosauromorpha) split from all taxa closer to lepidosaurs (here considered the new Lepidosauromorpha) at the onset of the Reptilia (=Amniota).

References
Zhang F, Li Y, and Wan X. 1984. A new occurrence of Permian seymouriamorphs in Xinjiang, China. Vertebrate Palasiatica22(4):294-306.

Comments from readers

I don’t get very many comments from readers.
Rarely do any of my blogposts get any feedback. The few rare comments I do get usually arrive whenever I make a mistake among the bird-like theropod dinosaurs, who have their own large fan base. Oddly, many of those readers also become further angered whenever I correct those mistakes, something I thought they were encouraging me to do! In the world of the Internet, and scientific discovery, such feedback is par for the course and must be expected. People in general, and scientists in particular, like their paradigms and don’t want outsiders tampering with them.

As a matter of practice, 
I try to be very specific and show images in my comments on the work of others, keeping anger and other negative emotions out of it.

Even more rarely
do I get replies that include specific instructions and data on how to correct my tracing errors. That has probably happened less than ten times in 1350 blog posts. Nevertheless, all of those rare comments are gratefully appreciated and acted upon. As my readers know, I’m only trying to get everything right, hoping only to provide new ideas to colleagues, whether they like those ideas or not. In Science, testing is supposed to be an okay thing to do. And if the tests are not valid, they can be done again and again until they are valid.

After 4+ years of reptileevolution.com
and pterosaurheresies.wordpress.com, I still haven’t seen any other paleontologist attempt to provide large gamut reptile cladograms based on specimens and species, now hovering around 560 taxa (exclusive of the pterosaur cladogram). The bird, dinosaur, croc and lizard paleontologists have done similar large gamut work, so I’m trying to avoid those well-studied clades, concentrating only on their origins. Let’s face it, a large gamut study of the basal reptiles needs to be published. The problem is, no PhD is interested or capable (time and travel constraints) of doing so, so far. Perhaps one is in progress.

The ‘hate mail’ I get reminds me of the 1961 Yankees
and specifically of the plight of Roger Maris, who, in 1961 approached and ultimately exceeded Babe Ruth’s hallowed 60 home runs in a season record. Teammate Mickey Mantle (Fig. 1) was also in that race that year that also featured an extended season. No one liked the fact that Maris, an outsider, was doing something so important.

Figure 1. Roger Maris and Mickey Mantile in 1961, two Yankees with a chance to break Babe Ruth's home run record.

Figure 1. Roger Maris and Mickey Mantile in 1961, two Yankees with a chance to break Babe Ruth’s home run record. The press and the fans were not kind to Maris during that season or to Mantle several years earlier.

Wikipedia reports, “In 1956, the New York press had been protective of Ruth when Mantle challenged Ruth’s record for most of the season. When Mantle fell short, finishing with 52, there seemed to be a collective sigh of relief from the New York traditionalists. The New York press had not been kind to Mantle in his early years with the team; he struck out frequently, was injury prone, was a true “hick” from Oklahoma, and was perceived as being distinctly inferior to his predecessor in center field, Joe DiMaggio. Mantle, however, over the course of time (with a little help from his friend and teammate Whitey Ford, a native of New York’s Borough of Queens), had gotten better at “schmoozing” with the New York media, and consequently gained the favor of the press. This was a talent that Maris, a blunt-spoken Upper Midwesterner, never attempted to cultivate. Maris was perceived as surly during his time on the Yankees.

“More and more, the Yankees became “Mickey Mantle’s team” and Maris was ostracized as an “outsider” and “not a true Yankee.” The press at that time seemed to be rooting for Mantle and belittling Maris. Mantle, however, was felled by a hip infection causing hospitalization late in the season, leaving Maris as the single remaining player with the opportunity to break Ruth’s home run record.”

Much of the same sort of human psychology is at play here.
In this case, yours truly, an outsider, not a true paleontologist, and not a PhD, has created a large gamut set of cladograms for reptiles and pterosaurs. The expanded data recovers a different topology than smaller studies, often handicapping themselves by using suprageneric taxa. And not all of those smaller studies match one another. The new topologies featured here and here were due in large part to taxon inclusion that was not attempted in the smaller studies. No one should see this as a threat.

That same outsider (yours truly) also broke a cardinal rule among paleontologists, “You have to see the fossil.”  Due to the large number of specimens involved, I have not seen every fossil, nor will anyone else in my lifetime. Referencing the literature is also common practice. That’s what it is there for!

Instead, after concentrated study,
I have reconstructed every included fossil and compared one with another graphically. That is something most paleontologists don’t do or do only rarely. As you should expect of such a large cladogram, all sister taxa actually look like they could be related, something that is too often lacking at certain nodes in certain other traditional cladograms.

In my attempt at making sure all the data was verifiable,
I have traced photos of in situ specimens and reconstructed them. That, evidently, is a sin, but one that is getting to be increasingly popular. And like most paleontologists, I have made a few mistakes along the way. These seem to happen most often when working with images of low resolution. When alerted to those mistakes, and provided better data, I have made corrections. That’s should be considered, “a good thing,” just as it is with the new data on Pluto (note the earlier fuzzy images that still have/had scientific value). Unfortunately, like Roger Maris’s situation in 1961, the jeers keep coming, but the large gamut reptile studies have not arrived yet.

I encourage more reader feedback,
but please, make replies constructive and include data if you have it. I don’t want anybody to be embarrassed by brash comments as future data and cladograms confirm current findings. And if you find two taxa that should not nest together, please let me know where the errant one should nest. If there are any mistakes in my presentation, I want to fix them.

 

 

Germanodactylus cristatus ventral reconstruction

Every so often
old reconstructions get updated. This time Germanodactylus cristatus gets the treatment with a new ventral view, matching the exposure of the cervicals and anterior dorsals (Fig. 1).

Figure 1. Germanodactylus cristatus in ventral view, wings outspread.

Figure 1. Germanodactylus cristatus in ventral view, wings and legs outspread. in that configuration the legs act as horizontal stabilizers and create their own lift. The torso appears to have been wider than deep. The prepubes were fused medially. The pelvis and three toes are conjectural based on matrix impressions, perhaps buried bones.

Note
the femoral heads are nearly at right angles to the femoral shafts. This gives the hind limbs more of an erect configuration, which may have aided this taxon in terrestrial stalking, something we talked about earlier here.

Figure 2. Germanodactylus cristatus in lateral view, bipedal/quadrupedal configuration.

Figure 2. Germanodactylus cristatus in lateral view, bipedal/quadrupedal configuration. The pelvis and three toes are based on matrix impressions.

References
Plieninger F 1901. Beiträge zur Kenntnis der FlugsaurierPaläontographica 48, 65–90 and pls 4–5.
Wellnhofer P 1970. Die Pterodactyloidea (Pterosauria) der Oberjura-Plattenkalke Süddeutschlands. Abhandlungen der Bayerischen Akademie der Wissenschaften, N.F., Munich 141: 1-133.
Wiman C 1925. Aus dem Leber der Flugsaurier. Bulletin of the Geological Insititute of the University of Uppsala 19: 115-127.

wiki/Germanodactylus