Pregnant hummingbird-like pterosaurs

Earlier
here and here we looked at pregnant pterosaurs. As you may recall, as lepidosaurs pterosaurs could retain their young in utero much longer than archosaurs do. Archosaur embryos are microscopic when laid and they develop in the egg outside of the uterus. Some extant lepidosaurs retain their young in utero to the stage of viviparity. Others lay eggs at an advanced stage. Today, two more tiny pterosaurs are shown to be adult females, based on the embryo inside of each of them.

As long-time readers know, 
phylogenetic analysis of the Pterosauria that includes the tiniest hummingbird-sized individuals from the Solnhofen formation nest them all as adults. They have been phylogenetically miniaturized and generally they nest at the bases of major clades. Generally the smallest pterosaurs are transitional from larger taxa and to larger taxa, but they are also often surrounded by other tiny transitional pterosaurs. That’s how we arrive at pterodactyloid-grade pterosaurs at least 4x. By convergence anurognathids and wukongopterids also added some, but not all, pterodactyloid traits.

Other workers,
who refuse to test the tiny ones, mistakenly assert that the tiny ones are babies. If that were true then, as the other workers suggest, pterosaurs would have to develop isometrically, changing shape with maturity. Several examples of embryo and juvenile pterosaurs demonstrate irrevocably that that is not true. Juveniles and embryos are carbon copies of the adults.

The smallest adult pterosaur is
Pterodactylus? kochi? B St 1967 I 276 (No. 6 of Wellnhofer 1970, (Figs. 1,2).

Figure 1. Pterodactylus? kochi? B St 1967 I 276 (No. 6 of Wellnhofer 1970) is the smallest known adult pterosaur. It is also pregnant. Note the relatively enormous sternal complex, analogous to that of a hummingbird of similar size.

Figure 1. Pterodactylus? kochi? B St 1967 I 276 (No. 6 of Wellnhofer 1970) is the smallest known adult pterosaur. It is also pregnant. Note the relatively enormous sternal complex, analogous to that of a hummingbird of similar size.

I did not realize
how large the sternal complex was on this pterosaur, Such a large pectoral anchor suggests the wings were flapped strongly or rapidly or both, possibly as in similarly-sized hummingbirds. The coracoids are also larger than earlier reconstructed.

Figure 2. The torso of B St 1967 I 276 (No. 6 of Wellnhofer 1970) showing the pectoral girdle and embryo.

Figure 2. The torso of B St 1967 I 276 (No. 6 of Wellnhofer 1970) showing elements of the pectoral girdle, pelvic girdle and embryo. The coracoids are also quite large. 

Nesting with the smallest known pterosaur
in the large pterosaur cladogram, is another tiny Solnhofen specimen, BMNH 42736, which also has a large sternal complex and is, by coincidence, pregnant.

Figure 4. Two of the smallest pterosaurs that both have a large sternal complex. BMNH42736 and B St 1967 I 276.

Figure 4. Two of the smallest pterosaurs that both have a large sternal complex. BMNH42736 and B St 1967 I 276. If your screen resolution is 72 dpi, these are shown > 1.5x larger than they were in life.

All I really wanted to do
was gather the data on this pterosaur to see where mistakes had been made. Finding tiny extra bones in the base of the abdomen was a surprise. These two, despite their differences, nest together in the large pterosaur tree.

Figure 6. Torso region of BMNH 42736 showing various bones, soft tissues and embryo.

Figure 6. BMNH 42736 showing various bones, soft tissues and embryo.

References
Bennett SC 2006. Juvenile specimens of the pterosaur Germanodactylus cristatus, with a review of the genus. Journal of Vertebrate Paleontology 26:872–878.SMNS
Hedges SB and Thomas R 2001. At the Lower Size Limit in Amniote Vertebrates: A New Diminutive Lizard from the West Indies. Caribbean Journal of Science 37:168–173.
Hone and Benton 2006. Cope’s Rule in the Pterosauria, and differing perceptions of Cope’s Rule at different taxonomic levels. Journal of Evolutionary Biology 20(3): 1164–1170. doi: 10.1111/j.1420-9101.2006.01284.x
Unwin D M 2006. The Pterosaurs From Deep Time. 347 pp. New York, Pi Press.
Wang X, Kellner AWA, Zhou Z and Campos DA 2008. Discovery of a rare arboreal forest-dwelling flying reptile (Pterosauria, Pterodactyloidea) from China. Proceedings of the National Academy of Sciences, 106(6): 1983–1987. doi:10.1073/pnas.0707728105
Wellnhofer P 1970. Die Pterodactyloidea (Pterosauria) der Oberjura-Plattenkalke Süddeutschlands. Abhandlungen der Bayerischen Akademie der Wissenschaften, N.F., Munich 141: 1-133.

wiki/Pterodactylus

 

 

A peek beneath the ribs of Pterodactylus scolopaciceps

A not so recent PLOSOne paper (Vidovic and Martill 2014, Late Jurassic) on Pterodactylus scolopaciceps (Meyer 1860, BSP 1937 I 18 (Broili 1938, P. kochi No. 21 of Wellnhofer 1970, 1991) provided the images seen here (Figs. 3, 4). It is one of the best preserved pterosaurs of all. Earlier we critically examined Vidovic and Martill 2014 here. A few short notes and images (Figs. 1,2) below will summarize those criticisms. Otherwise, the photos bring today’s news: tracings of the overlooked coracoids, sternal complex and an early embryo (Figs. 4, 5).

Vidovic and Martill reported,
“The majority of pterosaur species from the Solnhofen Limestone, including P. scolopaciceps are represented by juveniles.” This is utter rubbish.  Several hummingbird- to sparrow-sized adults, yes! …and some with long rostra! …but no verifiable juveniles, EXCEPT the juvenile of the giant Rhamphorhynchus recovered and described here. Remember, pterosaur embryos and juveniles are close matches to their parents as they develop isometrically, able to fly upon hatching, not allometrically. The large pterosaur tree demonstrates the phylogenetic miniaturization is what saved certain pterosaur lineages from extinction following a great radiation in the Late Jurassic. This is evidence Vidovic and Martill refuse to accept.

Vidovic and Martill continue:
“Consequently, specimens can appear remarkably similar due to juvenile characteristics detracting from taxonomic differences that are exaggerated in later ontogeny.” More rubbish based on adherence to Bennett (1996, 1996. 3006) who synonimized dozens of Solnhofen specimens without so much as an attempt at phylogenetic analysis, which lumps and separates the lot into individual taxa here. The Vidovic and Martill cladogram includes only 33 taxa (10 from Solnhofen) and lumps several pterosaurs successfully together (tapejarids, ctenochasmatids, pteranodontids), but fails to put these clades correctly into large clades, nesting sharp beak toothless taxa with broad beak toothy taxa, etc. etc.

Vidovic and Martill dig themselves deeper
“A hypodigm for P. scolopaciceps, comprising of the holotype (BSP AS V 29 a/b) and material Broili referred to the taxon is described. P. scolopaciceps is found to be a valid taxon, but placement within Pterodactylus is inappropriate. Consequently, the new genus Aerodactylus is erected to accommodate it.” As you can see (Figs. 1, 2) and as has been tested, placement within Pterodactylus (Fig. 2)  is MORE appropriate than nesting with purported sisters promoted by Vidovic and Martill (Fig. 1).

Figure 4. Sister taxa of "Aerodactylus" according to Vidovic and Martill 2014 include Gladocephaloides and Cycnnorhamphus. More rubbish.

Figure 1. Sister taxa of “Aerodactylus” according to Vidovic and Martill 2014 include Gladocephaloideus and Cycnnorhamphus. More rubbish. Neither are even related to one another as the former is a ctenochasmatid and the latter, of course, is a cycnohrmphid. Click to enlarge.

Evolution works in minute steps
and the more traits shared between specimens, both overall and in minute detail, the more closely they are related. Vidovic and Martill may also be working under the false assumption that pterosaurs are archosaurs and follow archosaur fusion patterns. No. Pterosaurs are lepidosaurs and follow lepidosaur fusion patterns, which are largely phylogenetic, not ontogenetic, as reported earlier.

Figure 3. Click to enlarge. The large pterosaur tree nests these three taxa together. So this Pterodactylus really is a Pterodactylus.

Figure 2. Click to enlarge. The large pterosaur tree nests these three taxa together. So this Pterodactylus (BSP AS V 28a/b) really is a Pterodactylus (contra Vidovic and Martill)

 

Enough about that paper.
I was drawn to this specimen (Fig. 3) because I did not have data for the coracoids and took another look for them with this excellent photo.

Figure 1. Pterodactylus scolpaciceps from Vidovic and Martill 2014 with elements below the ribs traced in color.

Figure 3. Pterodactylus scolpaciceps from Vidovic and Martill 2014 with elements below the ribs traced in color. Soft tissue is persevered in this specimen, and so is an embryo.

Lo and behold
the coracoids and sternal complex were visible as impressions (Figs. 3, 4) and there was something else further back… an embryo. Not full term. Not fully ossified. The wings are invisible or lost among the ribs and gastralia. Unlike 3D eggs, crushed fossils lay out all the elements into a single bedding plane.

Figure 2. Closeup of the torso of Pterodactylus scolopaciceps showing the coracoids, sternal complex and a passenger.

Figure 4 Closeup of the torso of Pterodactylus scolopaciceps showing the coracoids, sternal complex and a passenger. I was drawn to revisit this specimen because I lacked data for the length of the coracoids. This excellent image provided that data and possibly more. The bones of the embryo are not fully ossified yet. The shell is not formed. Those happen closer to the time just before egg-laying.

The embryo 
is the right size, shape and morphology to someday pass through the pelvis. The bones are soft and underdeveloped. No trace of an eggshell is apparent, but that’s not supposed to happen until the last stages of gestation.

Figure 4. Pterodactylus scolopaciceps reconstructed with the passenger shown here expelled. It is the right size, shape and morphology to be an embryo within an egg.

Figure 5. Pterodactylus scolopaciceps reconstructed with the passenger shown here expelled. It is the right size, shape and morphology to be an embryo within an egg.

Unlike archosaurs
lepidosaurs carry their young for longer terms, sometimes to the point of live birth (viviparity). Earlier I proposed that pterosaurs, like some of their sister lepidosaurs, carried their embryos until just prior to hatching. Other workers, all of whom consider pterosaurs archosaurs, thought egg burial was their method of reproduction. Not sure how they imagine a fragile pterosaur with tearable wing membranes would manage to dig through whatever dirt, sand or debris they were buried in. The aborted egg of Darwinopterus similarly contains an immature and unossified embryo. We also have an aborted fetus in Anurognathus and an aborted egg in the tiny pterosaur, Ornithocephalus added to the Pterodaustro embryo, the ornithocheirid embryo (revised recently) and the (relatively) giant, proto-anurognathid embryo.

How many pterosaur fossils are pregnant?
If they are doing their job, half of the adults should be pregnant, unless females greatly outnumber males, then that percentage goes up. Very few, however, will be preserved with late stage embryos that preserve even impressions of bones. As everyone knows the thinnest bone walls in the animal kingdom are pterosaur bones, thinner yet in embryos and  softer yet in younger embryos.

It’s time people
It’s time to let go of those old paradigms about pterosaur origins, wing shape and interrelationships. Those old hypotheses are not working. They cannot be verified. They are the stuff of myth. I would hate to think that these workers are refereeing manuscripts.

Carl Sagan said this about letting go of old paradigms,
“The essence of the Scientific method is the willingness to admit your’re wrong, to abandon ideas that don’t work, and the essence of religion is not to change anything, that supposed truths are handed down by some revered figure and no one is to make any progress beyond that because all the truth is thought to be in hand.”

References
Bennett SC 1995. A statistical study of Rhamphorhynchus from the Solnhofen Limestone of Germany: Year-classes of a single large species. Journal of Paleontology 69:569-580.
Bennett SC 1996. Year-classes of pterosaurs from the Solnhofen limestone of Germany: Taxonomic and systematic implications. Journal of Vertebrate Paleontology 16: 432–444.
Bennett SC 2006. Juvenile specimens of the pterosaur Germanodactylus cristatus, with a review of the genus. Journal of Vertebrate Paleontology 26:872–878.
Vidovic SU, Martill DM 2014. Pterodactylus scolopaciceps Meyer, 1860 (Pterosauria, Pterodactyloidea) from the Upper Jurassic of Bavaria, Germany: The Problem of Cryptic Pterosaur Taxa in Early Ontogeny. PLoS ONE 9(10): e110646. doi:10.1371/journal.pone.0110646

Hamipterus – a closer look at gender and ontogeny

Wang et al. 2014 introduced us
two years ago to a new collection of pterosaur parts from a monotypic population that was swept together and disarticulated by a flood event. As you may recall, five well-preserved three-dimensional eggs were recovered from the Early Cretaceous site in northwestern China. Sexual dimorphism was identified for the first time in pterosaurs with two different types of crests appeared on a variety of sizes of skulls (Figs. 1, 2). They named the new specimen, Hamipterus tianshanensis and the holotype was described as, One complete presumed female skull (IVPP V18931.1)”.

Figure 1. The female holotype and male paratype from the Hamipterus population assemblage fossil. The second tracing enlarges the male skull to the same length as the female skull. The color bar overprints indicate parts that differ in length from one skull to the other and a second overlay traces tooth position shifts from one to another.

Figure 1. The female holotype and male paratype from the Hamipterus population assemblage fossil. The second tracing enlarges the male skull to the same length as the female skull. The color bar overprints indicate parts that differ in length from one skull to the other and a second overlay traces tooth position shifts from one to another. The vestigial naris appears between the nasal and jugal beneath the crest. Direct comparisons like this help reveal subtle differences that otherwise might be overlooked.

Such a sweeping together of so many individuals
provides an unprecedented insight into several areas of pterosaur biology, but the data need to be rigorously examined so as not to jump to any conclusions.

Visible differences in the two skulls

  1. Crest shape
  2. Tooth placement
  3. Ventral maxilla shape
  4. Lateral extent of the premaxilla
  5. Depth of the skull anterior to the antorbital fenestra
  6. Concave vs. straight rostral margin (sans crest)
  7. Length of the upper temporal fenestra
  8. Placement of the vestigial naris
  9. Suborbital depth of the jugal

Gender
Wang et al. report, “About 40 male and female individuals in total were recovered, but the actual number associated might be in the hundreds. All of the discovered skulls have crests, which exhibit two different morphologies in size, shape, and robustness. Although morphological variation could be interpreted as individual variation, these marked differences suggest that the skulls belong to different genders. Hamipterus tianshanensis contradicts this hypothesis, because this species indicates that morphology of the crest, rather than its presence.”

Consider what we know about gender differences in birds and lizards,
It may be too soon to generalize over gender differences in pterosaurs. While each gender could have its own signature crest, size, etc., likewise each species likely had its own signature identity/crest/color/call, plumage, etc. At present, no other pterosaurs show verifiable gender differences. That’s why the Wang et al. paper was so important. Gender differences described for both Darwinopterus and Pteranodon were shown to be phylogenetic. Darwinopterus does present a mother with an aborted egg, but the father of the egg has not been identified. Hamipterus offers the best opportunity, so far, to bring some data to the table on this topic. And what Wang et al. indicate may indeed be true.

However, not enough care, IMHO, was administered to the non-crest differences in the skull material was made. Considering just the arrangement of teeth in the jaws (Fig. 1), is it possible that two very closely related species lived near one another? Or did individual variation cover a wider gamut than we now think is reasonable? Remember, among all the Pteranodon specimens now known (to me, at least), no two are identical. The same can be said for the Rhamphorhynchus and Pterodactylus specimens. And when you give Hamipterus a rigorous study, several subtle variations arise. Some of these arise from crushing. Others do not. With given data, one wonders if these could be two Hamipterus variations could be very closely related and.or very closely nesting sister taxa. OR… with present data, gender differences could extend beyond just the crest.

It is also possible
that male pterosaurs were rare rogues and this was a colony of females only with lots of individual variation. Do male lizards help raise their young? Do females? No. But pterosaurs might have been different. Wang et al. report on 40 individuals, but not on the male/female ratio or how many skulls are known. There were three in the holotype block. I’m guessing their specimen count was based on 40 skulls.

Figure 2. Finishing up the large skull with the large crest with two smaller candidates reveals that the slightly better fit is with the female skull.

Figure 2. Finishing up the large skull with the large crest with two smaller candidates reveals that the slightly better fit is with the female skull.

Ontogeny
Wang et al. report, “Ontogenetic variation is reflected mainly in the [lateral] expansion of the [spoon-shaped in dorsal view] rostrum.” Wang et al. reinforce what we know from other pterosaurs that they developed isometrically. Note the similarity between the crests of the smaller and larger ‘male’ specimens (Fig. 2). We’ve seen that before with Tupuxuara juveniles (Fig. 3).

Figure 1. Ontogenetic skull and crest development in Tupuxuara. Note the eyes are small and the rostrum is long in juveniles. Only the crest expands and only posteriorly.

Figure 3. Presumed ontogenetic skull and crest development in Tupuxuara. Note the eyes are small and the rostrum is long in juveniles. Only the crest expands and only posteriorly. Are are these two different sized but otherwise related species? With that longer rostrum, the smaller specimen may be distinct phylogenetically. No small crest Tupuxuara specimens are known.

Sedimentology
Wang et al. report, “Tempestite interlayers where nearly all of the pterosaur fossils are found suggest that large storms caused the mass mortality, event deposits, and lagersta¨ tte of the pterosaur population.”

Phylogenetically
Wang et al. discussed what Hamipterus is not. Their analysis nested it at the base of the Ornithocheiridae with complete lack of resolution. The large pterosaur tree nests Hamipterus with complete resolution between Boreopterus and Zhenyuanopterus.

Eggs
Wang et al. report, “A total of five eggs were recovered from the same site. The outer surface is smooth and exhibits no ‘papilla-like ornamentation,’ as was reported of the first pterosaur egg found in China.” Well that was a giant anurognathid egg, for which finding the parent will be big news. I’d be more interested to see comparisons to the second pterosaur egg found in China, the JZMP egg/embryo, which belonged to a rather closely related [to Hamipterus] ornithocheirid.

Wang et al. report, “Due to the close proximity to Hamipterus tianshanensis, the sole taxon found at the site, all of the eggs are referred to this species. Compared with other reptiles, the Hamipterus eggs show more similarities with some squamates,” I love it when every bit of data supports the theory that pterosaurs are lepidosaurs.

Wang et al. report, a 60µm calcareous eggshell followed by a thin 11µm inner membrane. They compared that to a snake egg of similar dimensions with a 60µm calcareous membrane followed by a much thicker 200µm inner membrane. Then they speculate wildly with this imaginative statement, “It is possible that Hamipterus also had a much thicker membrane, which was not completely preserved. We propose that such an eggshell structure, similar to that of some snakes, may well explain the preservation of the outer surface observed in pterosaur eggs.” IMHO, paleontologists go too far when they try to explain away data, rather than dealing with it directly. Elgin, Hone and Frey (2011) did this with their infamous wing membranes which they speculated suffered from imagined “shrinkage” in order to protect their verifiably false deep chord wing membrane hypothesis.

Wang et al report, “The [egg] size differences might also reflect different stages of development, since mass and dimensions differ between recently laid eggs and more developed ones.” There’s another possibility. Since we know that half-sized female pterosaurs were of breeding age (Chinsamy et al. 2008) they could have laid smaller eggs, producing smaller young, one source of rapid phylogenetic miniaturization.

Wang et al. report, “The combination of many pterosaurs and eggs indicates the presence of a nesting site nearby and suggests that this species developed gregarious behavior. Hamipterus likely made its nesting grounds on the shores of freshwater lakes or rivers and buried its eggs in sand along the shore, preventing them from being desiccated.” There’s another possibility. Since pterosaurs are lepidosaurs, they could have retained the eggs in utero until the young were ready to hatch. That also prevents them from desiccation. Since the flood tore the bones apart, any in utero eggs would have been torn away from the mother as well.

Notable by its absence
is any report of embryo bones inside the eggshells. I presume none were found or they would have been reported. That’s a shame, too, because eggs are nice little containers for complete skeletons, something lacking at the Hamipterus site. Some of the eggs appear to be evacuated, as if they were empty when buried. Or maybe all the juices were squeezed out during the rush and tumble of flood waters. If there was an embryo inside one of the Hamipterus eggs, and that is likely as the egg shell is applied just before egg laying, the embryo might have looked something like this (Fig. 3) based on the other pterosaur embryos inside their own two-dimensional eggs and the appearance of more complete sister taxa. During taphonomy the embryo inside would have been shaken AND stirred (but note some skulls are preserved complete without destruction!). The three dimensional egg contents would not accumulate on the randomly chosen longitudinal saw cut.

Figure 3. Wang et al. sliced one of the eggs lengthwise (yellow). if there is an embryo inside, it might have looked something like this. Since the egg has not been crushed to two dimensions, all the bones would not be now located in the plane of the slice, which was a random cut, not recognizing any embryo inside.

Figure 3. Wang et al. sliced one of the eggs lengthwise (yellow). if there is an embryo inside, it might have looked something like this. Since the egg has not been crushed to two dimensions, all the bones would not be now located in the plane of the slice, which was a random cut, not recognizing any embryo inside. Other embryos are typically in this pose.

Pterosaur hatchlings
of this size were precocial, able to fly shortly after hatching and large enough not to suffer from desiccation caused by so much surface area compared to volume.

References
Chinsamy A, Codorniú L and Chiappe LM 2008. Developmental growth patterns of the filter-feeder pterosaur, Pterodaustro guinazui. Biology Letters, 4: 282-285.
Elgin RA, Hone DWE, and Frey E. 2011.
The extent of the pterosaur flight membrane. Acta Palaeontologica Polonica 56 (1), 2011: 99-111 doi:10.4202/app.2009.0145 online pdf
Wang X et al.*, 2014.
 Sexually Dimorphic Tridimensionally Preserved Pterosaurs and Their Eggs from China, Current Biology. http://dx.doi.org/10.1016/j.cub.2014.04.054

Pterosaur reproduction and gender identification – SVPCA talks

Two upcoming SVPCA talks worth discussing:
Kellner et al and Unwin + Deeming both discuss pterosaur reproduction, growth and gender.

Key notes from the Kellner et al. (2015) abstract: All eggs show depressions, clearly indicating their overall pliable nature. SEM analysis shows that the eggshell structure is similar to some squamates. SEM analysis of [another] eggshell did not reveal an external calcareous layer suggesting that it was either removed due to taphonomy or not present at all. Histological section of the femur lacks medullary layer, a bone tissue reported in avian dinosaurs during ovulation and egg-laying phase. Those specimens, associated with experimental taphonomic studies, show that pterosaurs had two functional oviducts and laid eggs even smaller than previously thought, indicating that they have developed a reproductive strategy more similar to basal reptiles than to birds.”

Like I’ve been saying since 2007 and before.
Pterosaurs are non-squamate lepidosaurs. Egg shell morphology is just one more clue to this.

Unwin and Deeming abstract:
“Sexual dimorphism is common in extant vertebrates and almost certainly occurred in extinct species as well, but identifying this phenomenon in fossils is difficult. Meeting two key criteria: a large sample size in which all ontogenetic stages are present; and independent evidence of gender, is rarely possible, but has now been achieved for the early Upper Jurassic pterosaur Darwinopterus modularis. This pterosaur is represented by over 20 individuals ranging from hatchlings through juveniles to mature adults (ontogenetic status determined from osteological, histological and morphometric data). One example, ‘Mrs T’, is preserved with two eggs and thus clearly a female. Approximately half the mature individuals of Darwinopterus exhibit a cranial crest and several of these individuals have a relatively narrow pelvis. The remainder lack a cranial crest and in two cases, including Mrs T, have a relatively broad pelvis. All immature individuals lack a crest, an observation that applies to other species of pterosaur in which immature individuals are known. This pattern of morphological variation shows that the cranial crest and pelvis of Darwinopterus modularis are sexually dimorphic. Datasets for other pterosaurs are less complete and/or lack independent evidence of gender, but many species including Ctenochasma gracile, Germanodactylus cristatus and Pteranodon longiceps, exhibit directly, or closely, comparable patterns of anatomical variation to Darwinopterus and are likely to have been sexually dimorphic. We conclude that the spectacular variability in the shape and size of pterosaur cranial crests was likely generated by sexual selection rather than processes such as species recognition.”

Unfortunately
other than with the presence of eggs in association, sexual dimorphism has not been determined in other pterosaurs in which a large sample size is present (Rhamphorhynchus, Pteranodon, Germanodactylus, Pterodactuylus), even without eggs in association. This is widely recognized, hence the excitement level in the abstract for Darwinopterus. Rather speciation of these taxa has been determined through phylogenetic analysis. Speciation has also been determined for the several Darwinopterus specimens. Currently published specimens don’t divide neatly in two. If that changes with the addition of 15 more, I’ll be happy to note that. Unwin and Deeming do not mention phylogenetic analysis in their abstract. If this is a clue to their methods such laziness in skipping phylogenetic analysis is becoming more and more common, especially when it suits a false paradigm. You can’t just eyeball these things. You have to put your data through analysis. Otherwise the work will always be doubted and you’ll be ‘pulling a Bennett’ (assertion of association without cladogram evidence). The purported hatchlings noted by Unwin and Deeming also need to be run through analysis. Are they examples of phylogenetic miniaturization or actual juveniles? Adding hatchlings and embryos along with tiny adults to analysis has been online for more than four years, so there is no excuse for avoiding it.

Tiny wukongopterids are welcome news, by the way. This clade is one of a few that currently lacks any tiny representatives and that lack is the current best reason why wukongopterids left no descendants in the Cretaceous.

The bone originally identified as an ischium on Mrs. T was a misidentified displaced prepubis. The actual ischia were preserved in the counter plate and they were relatively narrow.

Unwin does not support isometric growth during ontogeny, which is otherwise a well established fact in pterosaurs. So he may be accepting dissimilar morphologies as juvenile examples (pulling another Bennett). Very dangerous. As in all other pterosaurs, like Pteranodon, you have to evolve crested derived forms from non-crested basal forms. Unwin and Deeming, if you’re reading this: before you publish your paper, send me your data, if you don’t want to do the analysis yourself. I’ll send back the recovered cladogram. Don’t make the same mistakes again. However, if the juveniles are isometric copies of the adults, then congratulations and remember to give credit where credit is due.

References
Kellner  AWA et al. 2015. Comments on pterosaur reproduction based on recently found specimens from the Jurassic and Cretaceous of China. Among the most spectacular pterosaur finds done in recent years is the bone-bed from the Tugulu Group (Lower Cretaceous) discovered in the Hami area, Xinjiang Uyghur Autonomous Region of China. SVPCA 2015 abstracts.
Peters D 2007. The origin and radiation of the Pterosauria.
Flugsaurier. The Wellnhofer Pterosaur Meeting, Munich 27
Unwin DM and Deeming CD 2015. New evidence for sexual dimorphism in the basal monofenestratan pterosaur Darwinopterus. SVPCA 2015 abstracts.

More tiny birds and tiny pterosaurs

Earlier we took a peek at a few tiny birds and pterosaurs. Here (Fig. 1) are several more.

Traditional paleontologists
insist that these tiny pterosaurs were babies of larger forms that looked different, (Bennett 1991, 1992, 1994, 1995, 1996, 2001, 2006, 2007, 2012, 2014) ignoring or not aware of the fact that we know pterosaur embryos and juveniles were virtually identical to their adult counterparts (Fig. 2). Bennett (2006) matched two tiny short-snouted pterosaurs (JME SoS 4593 and SoS 4006 (formerly  PTHE No. 1957 52) to Germanodactylus, but they don’t nest together in the large pterosaur tree.

Figure 1. Tiny pterosaurs and tiny birds to scale showing that tiny pterosaurs were generally about the size of the tiny Early Cretaceous bird.

Figure 1. Tiny pterosaurs and tiny birds to scale showing that tiny pterosaurs were generally about the size of the tiny Early Cretaceous bird. I have, for over a decade, promoted the fact that these tiny pterosaurs were adults, the size of modern hummingbirds and wrens.

One of the most disappointing aspects of modern paleontology
is the refusal of modern pterosaur workers to include in their analyses the small and tiny pterosaurs. They were all the size of living hummingbirds and wrens. Many were similar in size to extinct Early Cretaceous birds (Fig. 1). Those workers don’t want to add these taxa to their lists on the false supposition that the tiny pterosaurs are babies of, so far unknown adults. Note Bennett’s long body of work (see below) indicated otherwise, but never with phylogenetic analysis.

Phylogenetic analysis (Peters 2007) reveals these tiny pterosaurs are adults or can be scored as adults. They are surrounded by adults and they often form transitional taxa in the evolutionary process of phylogenetic miniaturization between larger long-tailed pterosaurs and larger short-tailed pterosaurs.

Figure 1. Click to enlarge. There are several specimens of Zhejiangopterus. The two pictured in figure 2 are the two smallest above at left. Also shown is a hypothetical hatchling, 1/8 the size of the largest specimen.

Figure 2. Click to enlarge. There are several specimens of Zhejiangopterus. The two pictured in figure 2 are the two smallest above at left. Also shown is a hypothetical hatchling, 1/8 the size of the largest specimen. This is evidence that juveniles were virtually identical to adults, except in size.

More importantly,
earlier we discussed several examples of juvenile pterosaurs morphologically matching adults here, here and here. So young pterosaurs have been shown to match their adult counterparts. They don’t transform like young mammals and dinosaurs do. They were ready to fly upon hatching IF they were the minimum size to avoid desiccation, as discussed earlier here.

The most interesting aspect
to the whole tiny pterosaur story is how small their smallest hatchlings would be. We looked at that earlier here.

References
Bennett SC 1991. Morphology of the Late Cretaceous Pterosaur Pteranodon and Systematics of the Pterodactyloidea. [Volumes I & II]. Ph.D. thesis, University of Kansas, University Microfilms International/ProQuest.
Bennett SC 1992. 
Sexual dimorphism of Pteranodon and other pterosaurs, with comments on cranial crests. Journal of Vertebrate Paleontology 12: 422–434.
Bennett SC 1994. 
Taxonomy and systematics of the Late Cretaceous pterosaur Pteranodon (Pterosauria, Pterodactyloidea). Occassional Papers of the Natural History Museum University of Kansas 169: 1–70.
Bennett SC 1995. A statistical study of Rhamphorhynchus from the Solnhofen limestone of Germany: year classes of a single large species. Journal of Paleontology 69, 569–580.
Bennett SC 1996. 
Year-classes of pterosaurs from the Solnhofen limestones of Germany: taxonomic and systematic implications. Journal of Vertebrate Paleontology 16:432–444.
Bennett SC 2001.
 
The osteology and functional morphology of the Late Cretaceous pterosaur Pteranodon. Part I. General description of osteology. Palaeontographica, Abteilung A, 260: 1–112. Part II. Functional morphology. Palaeontographica, Abteilung A, 260: 113–153
Bennett SC 2006. Juvenile specimens of the pterosaur Germanodactylus cristatus, with a revision of the genus. Journal of Vertebrate Paleontology 26(4): 872–878.
Bennett SC 2007. A second specimen of the pterosaur Anurognathus ammoni. Paläontologische Zeitschrift 81(4):376-398.
Bennett  SC (2012) [2013] 
New information on body size and cranial display structures of Pterodactylus antiquus, with a revision of the genus. Paläontologische Zeitschrift (advance online publication) doi: 10.1007/s12542-012-0159-8
http://link.springer.com/article/10.1007/s12542-012-0159-8
Bennett SC 2014. A new specimen of the pterosaur Scaphognathus crassirostris, with comments on constraint of cervical vertebrae number in pterosaurs. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen, 271(3): 327-348.
Peters D 2007. The origin and radiation of the Pterosauria. Flugsaurier. The Wellnhofer Pterosaur Meeting, Munich 27

 

Another Pterodaustro embryo reconstructed

I missed this one last June.

Grellet-Tinner et al. (2014)
report on a new Pterodaustro egg, concentrating on the eggshell morphology and its chemistry while strangely ignoring the contents. Here’s an egg they identify by its proximity to a loose Pterodaustro skeletal element and the egg’s similarity to the earlier pterosaur egg (Chiappe et al. 2004), which includes a Pterodaustro embryo.

And yet, there IS a nicely preserved embryo inside the 2014 egg! 
Here it is (Figs. 1, 2) traced using DGS and after reassembling the color graphic elements. Nearly every bone in the pterosaur is visible, down to the prepubes and wing tip unguals. The cranium, unfortunately, is below the two sets of free fingers, so it is difficult to restore without additional resolution (which I have requested).

Figure 1. Egg image from Grellet-Tinner et al. 2014, DGS tracing of elements reveals a complete Pterodaustro embryo with a small cranium and long rostrum.

Figure 1. Egg image from Grellet-Tinner et al. 2014, DGS tracing of elements reveals a complete Pterodaustro embryo with a small cranium and long rostrum. Click to enlarge. Grellet-Tinner et al. identified this egg not by its contents but by its proximity to Pterodaustro skeletal element and its similarity to the 2004 egg, which includes an embryo.

Earlier we looked at the first Pterodaustro embryo (Fig. 2) and discussed attempts by others at reconstructing the embryo without tracing the bones(!) here. Several years ago Laura Codorniu kindly sent me a high resolution image of the 2004 egg and embryo from which I produced that reconstruction. On agreement with her I have not published online that loaned egg image nor the tracing of it. (You might recall the 2004 egg was illustrated with a line drawing (Fig. 3), not a photo). Here (Fig.1) the egg photo is published (Grellet-Tinner et al. 2014. You’ll note the two reconstructions are essentially identical (Fig. 2) except for the slightly larger size of the 2014 specimen. Evidently it was in the egg a little longer and closer to hatching.

2004 specimen: MIC-V 246 – 6.0 x 2.2 cm, preserved in a concretion in two-dimensions with eggshell remains delimiting the egg outline around the embryo

2014 specimen: MIC-V 778 – 6.0 x 2.2 cm, the first partial three-dimensional egg, same deposit, but the authors note the matrix records offshore lacustrine environments characterized by very fine, laminated siltstones

Figure 2. Pterodaustro embryos compared. Note the 2004 specimen is a little larger with more robust wing finger phalanges and a larger sternal complex.

Figure 2. Pterodaustro embryos compared. Note the 2014 (778) specimen is a little larger with more robust wing finger phalanges and a larger sternal complex. Click to enlarge.

Let’s talk about the embryos first.
The two Pterodaustro embryos (Fig. 2) are essentially identical, but the 2014 specimen is slightly larger with more robust and longer wing phalanges, a more robust proximal tail, more robust free fingers, and a larger sternal complex. These elements suggest it was getting ready to hatch and to fly shortly after hatching.

Both embryos had the proportions of the adult Pterodaustro.
That embryo data supports the hypothesis of isometry during ontogeny found in other pterosaurs, like Zhejiangopterus. The embryos had the basic proportions of the adult.

Unfortunately
this hypothesis has been rejected by all other pterosaur experts that I am aware of, even in the face of incontrovertible evidence such as this. Bennett (1993-2007) is the traditional  proponent of allometry (or great change) during ontogeny, having written several papers on it.

The fact remains
There is no short rostrum in the two Pterodaustro embryos, much as some would like to imagine. Furthermore Phylogenetic analysis nests tiny pterosaurs as miniaturized taxa transitional between large taxa of one morphology and large taxa of another morphology. This happens often in the clade Reptilia and all of its subclades, like the origin of Dinosauria, Aves and Mammalia. It’s not a big deal. It’s common.

Pterodaustro adult with manual digit 3 repaired.

Figure 3. Pterodaustro adult to scale with the 2004 3mbryo and egg.

Now on to eggshells…
Pterosaurs have very thin eggshells. The Asian eggs (Wang 2004, Ji et al. 2004, Lü et al. 2009) were considered ‘leathery.’ By contrast the South American Pterodaustro eggs have a calcium coating, but still extremely thin. Did that calcium coating maintain an elliptical shape? Apparentely so.

Grellet-Tinner et al. 2014 report, “the only pterosaur eggs with embryos obtained from the Yixian Formation in China are soft-shelled, whereas those from Argentina display a well-mineralized calcitic shell. This conundrum may reflect a similar species-specific reproductive strategy observed in modern Geckonids, which lay both hard- and soft-shelled eggs.”

That’s a great explanation for the differences seen in pterosaur eggs.

Grellet-Tinner et al. 2014 report, “The calcareous eggshells of amniotes must allow the exchange of respiratory gases to support embryonic metabolism, while simultaneously regulating flux of water vapor. The thickness of the eggshell and the size and number of the pores, which both determine the conductance of the eggshell to respiratory gases and water vapor, are key features that facilitate oxygen supply to the embryo, allow it to excrete carbon dioxide, and prevent excessive water loss.”

Studies demonstrate
an inverse correlation between eggshell thickness and length of uterine egg retention. So, with such thin eggshells, pterosaur females likely held their eggs in utero, as many lizards do, depositing them shortly before hatching. That hypothesis is also in direct contrast to that of most pterosaur workers (like Grellet-Tinner et al) who continue to consider pterosaurs archosaurs, but are stymied by the lack of a thick calcium carbonate eggshell, like those of birds and crocs.

Eggshell pores
are important features of amniote eggs. They allow the exchange of gases and water between the developing embryo and the environment. Since pores are open to the atmosphere for gas exchange, they are visible on the eggshell surface sometimes without a microscope. However, distinct from archosaur eggshells, Pterodaustro eggshells have no pores, another clue that they were held in utero by their lizard-derived mothers.

Grellet-Tinner et al. 2014 report, “Although the shell of MIC-V 778 is superbly preserved, the scanning electron micrographs do not reveal any pores, for which there could be a number of explanations: (i) pores may not have been evident in a scanning electron micrograph of a radial fracture of the eggshell, which is possible but not likely; (ii) pores were not present or noticeable at the ontogenetic stage at which the egg died; or (iii) all pores were located in the larger pole like in extinct troodontids.”

Unfortunately the G-T assumed that Pterodaustro was an archosaur. But that’s a mistake. Pterosaurs were lepidosaurs. And most, if not all of them retain the egg in utero longer than archosaurs do.

Eggshell thickness
Grellet-Tinner et al. 2014 report, “The thickness of the shell of MIC-V 778 is approximately 50 micrometers, which is less than one third of the thickness expected for the eggshell of an avian egg of similar size (179 micrometers). As eggshell conductance is inversely proportional to eggshell thickness the MIC-V 778 calcareous eggshell would have a conductance more than three times higher than for an avian egg of equivalent size.”

“if considering reptilian incubation (hard-shelled eggs of crocodilians, geckos, and turtles generally do not lose water during incubation), and given the relative egg: female mass ratio of pterosaurs, it is very unlikely that pterosaur eggs would lose water during incubation. Thus, nest humidities would need to be even higher than our calculations suggest.”

Like inside a mother?

Apparently not.
Grellet-Tinner et al. 2014 report, “Such a nesting environment is likely to result in the egg being incubated in a nest with a very high moisture content during some of the incubation time, conditions matching those of grebes or primitive flamingos living in similar  eological/environmental settings. The eggs of P. guinazui and grebes, which incubate their eggs in moist nests, are of similar size and have similar estimated gas conductances that are about three times higher than in other avian eggs of similar size. The difference between these two species is the way in which the higher conductance is achieved. In P. guinazui, high conductance is achieved by having a thin eggshell (short pore length), whereas in grebes high eggshell  conductance is achieved through having a larger number of pores through an eggshell that is as thick as that in other birds.”

Another outer layer on the pterosaur egg.
Grellet-Tinner et al. 2014 report, “Another curious feature of MIC-V 778 is the ubiquitous 2 micrometer outer layer of calcium carbonate, an eggshell feature unknown in reptiles but observed in a few avian taxa, notably Mirandornithes (grebes and flamingos). Termed the “accessory layer or surface crystal layer”, it may function in avian eggs that are incubated in relatively wet or muddy nests to prevent water from occluding the pores through the eggshell and thus prevent a reduction in the eggshell conductance to oxygen.”

Since pterosaurs are not birds, this is a convergent appearance of the surface crystal layer. So at least some reptiles (pterosaurs) had this new layer.

Grellet-Tinner et al. 2014 report, “the sum of all the taphonomic and biological evidence gathered for MIC-V 246 and MIC-V 778 allow us now to reconsider this original hypothesis in favor of incubation in nests built in palustrine ecosystems, where the water could have intermittently permeated in the nests under a semi arid climatic regime.”

Or the egg could have been retained by the mother…

Taphonomic transportation of the egg
Grellet-Tinner et al. 2014 report: “A grey-brown matrix encrusts the egg, but no sediment was present inside the egg when discovered, implying that the egg was complete and not broken at the time of burial. This preservation is most unusual considering the fragility of the specimen and the thinness of its eggshell, so even a short distance of transportation by underwater debris flow seems unlikely.” 

And yet some of the legs bones were broken and other bones displaced (Fig. 1). The tail extends beyond the shell boundaries.

It is also rare
in that locality to find complete Pterodaustro specimens. Most are fragments and loose bones. I know of only one complete specimen. As in the Hamipterus find that includes a nesting colony disassociated bones and associated pterosaur eggs, the possibility must be considered that the eggs found had not been laid yet, but were inside the mother when she was killed, then as the carcass rotted, was transported and buried, the egg became disassociated with the mother and her disassociated bones. I’m no ob-gyn, but something tells me the pterosaur uterus likely also produced a mesohaline environment.

Grellet-Tinner et al. 2014 report, “our results demonstrate that the nesting paleoenvironment of this pterosaur species was closely linked to a mesohaline [salt concentrations between 5 and 18 ppt,] lacustrine ecosystem in a basin governed by regional tectonic subsidence, a setting characteristic for the feeding and reproduction of modern flamingos.”

Or the eggs could have been held in utero.
See what happens when the first domino tells you ‘pterosaurs are archosaurs’?

One final thought
Darren Naish and other detractors are fond of saying that I see things with DGS that aren’t there. This Pterodaustro embryo is evidence to the contrary. All the bones looked like the other Pterodaustro embryo and both looked like those of the adult.

References
Bennett SC 1993a. The ontogeny of Pteranodon and other pterosaurs. Paleobiology 19, 92–106.
Bennett SC 1993b. Year classes of pterosaurs from the Solnhofen limestone of southern Germany. Journal of Vertebrate Paleontology. 13, 26A.
Bennett SC 1995. A statistical study of Rhamphorhynchus from the Solnhofen limestone of Germany: year classes of a single large species. Journal of Paleontology 69, 569–580.
Bennett SC 1996a. Year-classes of pterosaurs from the Solnhofen limestones of Germany: taxonomic and systematic implications. Journal of Vertebrate Paleontology 16:432–444.
Bennett, SC 1996b. The phylogenetic position of the Pterosauria within the Archosauromorpha. Zoological Journal of the Linnean Society 118:261–309.
Bennett SC 2001a, b. The osteology and functional morphology of the Late Cretaceous pterosaur PteranodonPart I. General description of osteology. Palaeontographica, Abteilung A, 260:1–112. Part II. Functional morphology. Palaeontographica, Abteilung A, 260:113–153.
Bennett SC 2006. Juvenile specimens of the pterosaur Germanodactylus cristatus, with a review of the genus. Journal of Vertebrate Paleontology 26:872–878.
Bennett SC 2007. A review of the pterosaur Ctenochasma: taxonomy and ontogeny. Neues Jahrbuch fur Geologie und Paläontologie, Abhandlungen 245:23–31.
Bonaparte J F 1970. Pterodaustro guinazui gen. et sp. nov. Pterosaurio de la Formacion Lagarcito, Provincia de San Luis, Argentina y su significado en la geologia regional (Pterodactylidae)”, Acta Geologica Lilloana, 10: 209-225.
Chiappe LM, Codorniú L, Grellet-Tinner G and Rivarola D. 2004. Argentinian unhatched pterosaur fossil. Nature, 432: 571.
Chinsamy A, Codorniú L and Chiappe LM 2008. Developmental growth patterns of the filter-feeder pterosaur, Pterodaustro guinazui. Biology Letters, 4: 282-285.
Codorniú L and Chiappe LM 2004. Early juvenile pterosaurs (Pterodactyloidea: Pterodaustro guinazui) from the Lower Cretaceous of central Argentina. Canadian Journal of Earth Science 41, 9–18. (doi:10.1139/e03-080)
Deeming C and Unwin DM 2007. Eggshell structure and its implications for pterosaur reproductive biology and physiology. Flugsaurier, the Wellnhofer pterosaur meeting, Munich, 12.
Grellet-Tinner G, Wroe S, Thompson SB and Ji Q 2007. A note on pterosaur nesting behavior. Historical Biology 19:273–277.
Grellet-Tinner G, Thompson c MB, Fiorelli LE, Argañaraz E, Codorniú L, Hechenleitner EM 2014. The first pterosaur 3-D egg: Implications for Pterodaustro guinazui nesting strategies, an Albian filter feeder pterosaur from central Argentina. Geoscience Frontiers 5 (2014) 759e765
Ji Q, Ji S-A, Cheng Y-N, You HL, Lü J-C, Liu Y-Q and Yuan CX 2004. Pterosaur egg with leathery shell. Nature 432:572.
Lü J, Unwin DM, Jin X, Liu Y and Ji Q 2009. Evidence for modular evolution in a long-tailed pterosaur with a pterodactyloid skull. Proceedings of the Royal Society London B  (DOI 10.1098/rspb.2009.1603.)
Lü J, Unwin DM, Deeming DC, Jin X, Liu Y and Ji Q 2011a. An egg-adult association, gender, and reproduction in pterosaurs. Science, 331(6015): 321-324. doi:10.1126/science.1197323
Lü J, Xu L, Chang H and Zhang X 2011b. A new darwinopterid pterosaur from the Middle Jurassic of Western Liaoning, northeastern China and its ecological implicaitions. Acta Geologica Sinica 85: 507-514.
Wang X-L and Zhou Z 2004. Palaeontology: pterosaur embryo from the Early Cretaceous. Nature 429: 623.

New smallest Pteranodon: Bennett 2014 JVP abstract

Figure 1. Pteranodon ingens. Full size and little Ptweety the baby Pteranodon, not curated.

Figure 1. Pteranodon ingens. Full size and little Ptweety the baby Pteranodon, not curated. Alongside in black is a hypothetical hatchling half the size of Ptweety. With a 1.5m wingspan, Ptweety is still the smallest, compared to Bennett’s 1.76 m wingspan. The Bennett Pteranodon is not shown.

I was hoping a curated specimen would follow Ptweety, the baby Pteranodon (Fig. 1). It was just a matter of time. Here it is in the 2014 JVP abstracts.

From the Bennett abstract:
“An earlier study of all available specimens of the pterosaur Pteranodon from the
Smoky Hill Chalk Member of the Niobrara Formation found a bimodal size distribution. The small size class with estimated wingspans in life of ~3.1-4.8 m was twice as abundant as the large, with wingspans of ~4.8-6.7 m, and immature specimens formed ~15% of each class suggesting that they cannot be age classes. The bimodal distribution was interpreted as evidence of sexual dimorphism and the absence of specimens smaller than ~3 m wingspan was interpreted as evidence of bird-like parental care during rapid growth to adult size before flying and feeding independently. A new immature specimen of Pteranodon with an estimated wingspan of only 1.76 m demonstrates that juveniles were capable of flying and feeding independently, contradicting the interpretation of parental care during rapid growth. Instead Pteranodon apparently was precocial, flying and feeding independently during several years of growth to adult size as previously observed in Rhamphorhynchus, Pterodactylus, and Pterodaustro. Therefore, the absence of Pteranodon juveniles and a similar absence of Nyctosaurus juveniles from the Smoky Hill Chalk indicates those taxa had multi-niche ontogenies, occupying distinct niches in different locations and environments at different stages of their life history. Thus, the Smoky Hill Chalk represents a pelagic feeding environment of Pteranodon and Nyctosaurus adults whereas hatchlings and juveniles presumably fed on smaller prey in lacustrine, riverine, estuarine, or coastal environments. The pterosaur records of most other Lagerstätten are consistent with multi-niche ontogeny being the norm in pterosaurs. For example, the record of Azhdarcho in the Bissekty Formation consists of hatchlings and adults and represents a breeding ground, that of the Solnhofen Limestone consists primarily of hatchlings and juveniles and represents a nursery environment of juveniles in sheltered lagoons near breeding grounds whereas those of the Romualdo and Cambridge Greensand Formations consist of adults and represent coastal feeding environments of adults. One exception seems to be the record of Pterodaustro in the Lagarcito Formation, which consists of eggs, hatchlings, juveniles, and adults in a single location and environment; however, that may reflect a special environment required to effectively utilize the filter-feeding specializations of the taxon.”

Bennett has been the target of many Pterosaur Heresies blogposts.
And for good reason: (no gender classes, this represent several species evolving from small, small-crest forms to several clades of large, large-crest forms, etc. etc. etc.).

Here Bennett is right on the money
when he agrees to different niches for juvenile and adult pterosaurs, which we discussed earlier here, due to the rarity of juvenile pterosaurs in the fossil record, a topic in which Bennett takes the opposite stance.

Not mentioned in the Bennett abstract
is the fusion of the extensor tendon process to manual 4.1, which occurs in all Pteranodon specimens (even Ptweety) and no Nyctosaurus specimens except the crested ones. The same goes for scapula and coracoid fusion (fused in Pteranodon, not in Nyctosaurus). I wonder what the data is on his new juvenile Pteranodon?

Figure 2. Ptweety the juvenile Pteranodon. Note the presence of a fused extensor tendon process, a long rostrum and small orbit in this isometrically identical juvenile pterosaur.

Figure 2. Ptweety the juvenile Pteranodon. Note the presence of a fused extensor tendon process, a long rostrum and small orbit in this isometrically identical juvenile pterosaur.

The presence of hatchling Azhdarcho specimens in a breeding ground comes as something of a surprise. Good news! The literature (Averianov 2010) only refers to a juvenile/immature specimen represented by a notarium (4cm long, compared to a 6.5 notarium for an unrelated adult mid-size Pteranodon adult.)

Bennett does not mention the growth series in Tapejara and Zhejiangopterus. The abstract was probably written before the Caiuajara nesting site. Pterodaustro embryos and hatchlings are well known (Chiappe et al. 2004, Chinsamy et al. 2008, Codorniú and Chiappe 2004). The various growth series described by Bennett (1995, 1996) actually represent individual species if not genera. This he would discover by phylogenetic analysis.

Ptweety will never be published because it was extracted and prepared without documentation and the last I heard it was a standing mount at an online retail store. Good to hear that another juvie Pteranodon is out there and hopefully will soon be published.

On a side note: 
The blog post on the evolution of frogs is getting an unusually large number of hits. Not sure why. Let me know if there is anything else you want to learn more about.

References
Averianov AO 2010. The osteology of Azhdarcho lancicollis Nessov 1984 (Pterosauria, Azhdarchidae) from the Late Cretaceous of Uzbekistan. Proceedings of the Zoological Institute RAS. 314(3):264–317.
Averianov AO 2013. 
Reconstruction of the neck of Azhdarcho lancicollis and lifestyle of azhdarchids (Pterosauria, Azhdarchidae). Paleontological Journal 47 (2): 203-209. DOI: 10.1134/S0031030113020020
Bennett SC 1995. A statistical study of Rhamphorhynchus from the Solnhofen limestone of Germany: year classes of a single large species. Journal of Paleontology 69, 569–580.
Bennett SC 1996. Year-classes of pterosaurs from the Solnhofen limestones of Germany: taxonomic and systematic implications. Journal of Vertebrate Paleontology 16:432–444.
Bennett, SC 2014. New smallest specimen of the pterosaur Pteranodon and multi-niche ontogeny in pterosaurs. Journal of Vertebrate Paleontology abstracts, Berlin Conference 2014.
Chinsamy A, Codorniú L and Chiappe LM 2008. Developmental growth patterns of the filter-feeder pterosaur, Pterodaustro guinazui. Biology Letters, 4: 282-285.
Chiappe LM, Codorniú L, Grellet-Tinner G and Rivarola D. 2004. Argentinian unhatched pterosaur fossil. Nature, 432: 571.
Codorniú L and Chiappe LM 2004. Early juvenile pterosaurs (Pterodactyloidea: Pterodaustro guinazui) from the Lower Cretaceous of central Argentina. Canadian Journal of Earth Science 41, 9–18. (doi:10.1139/e03-080)

Two really big anurognathids

Yesterday we looked at the adult sisters to the JZMP embryo. Today we’ll do the same with the IVPP embryo.

Figure 1. Large anurognathids and their typical-sized sisters. Here the IVPP embryo enlarged to adult size is larger than  D. weintraubi and both are much larger than more typical basal anurognathids, Mesadactylus and MCSNB 8950.

Figure 1. Large anurognathids and their typical-sized sisters. Here the IVPP embryo enlarged to adult size is larger than D. weintraubi and both are much larger than more typical basal anurognathids, Mesadactylus and MCSNB 8950.

The IVPP embryo pterosaur (Wang et al. 2004), the first ever described, was wrongly considered a juvenile ornithocheirid based on its small tail and short rostrum. At the time pterosaur juveniles were purported to have a short rostrum, but this has been proven wrong at every turn. First on that list: the JZMP embryo is an ornithocheirid and it has a long rostrum.

Phylogenetic analysis nests the IVPP embryo pterosaur together with Mesadactylus, a poorly known anurognathid. Both are sister to another large anurognathid, the misnamed “Dimorphodon” weintraubi. And all three were derived from a sister to MCSNB 8950, wrongly considered a juvenile “Eudimorphodon.

It’s a wonder to see a giant anurognathoid with an embryo the size of other anurognathoids. Only D. weintraubi approaches the embryo enlarged to adult size. Can’t wait for someone in China to come out with the big news of the discovery of the adult.

References
Wang X-L and Zhou Z 2004. Palaeontology: pterosaur embryo from the Early Cretaceous. Nature 429: 623.

Witton’s baby Pterodaustro fantasy

Pterodaustro embryo as falsely imagined in Witton 2013. The actual embryo had a small cranium, small eyes and a very long rostrum.

Figure 1. Pterodaustro embryo as falsely imagined in Witton 2013. The actual embryo had a small cranium, small eyes and a very long rostrum.

Sad for everyone involved,
Witton 2013 completely imagined his embryo Pterodaustro (Fig. 1), giving it giant eyes and a tiny rostrum, as if it was a baby bird. This complete disregards the data that shows the rostrum was very long while the cranium and orbit were very small (Fig. 2), virtually matching the 8x larger adult.

Earlier we examined the ontogenetic growth of another pterosaur with a long rostrum and tiny eye, Zhejiangopterus. So isometric growth is the pterosaur pattern, not allometric growth. Pterosaur hatchlings were not cute!

Pterodaustro embryo

Figure 2. Pterodaustro embryo. There certainly is no short snout/large eye here!

As everyone knows
tracing does have its limitations, but Science can’t support the complete disregard for data that Witton 2013 promotes when he free hands from his hopes and dreams.

Unfortunately this is not the first time I’ve had to shine a light on Witton’s promotion of false data. Here, here and here are others and they also have links.

I encourage Mark Witton to start tracing skeletal elements for more accuracy.  

References
Witton M. 2013. Pterosaurs. Princeton University Press. 291 pages.

Mark Witton’s “Pterosaurs” – a book review part 1

Dr. Mark Witton is fantastic artist and devotee of pterosaurs. He has a new book called Pterosaurs (with an Amazon.com preview). I’ve ordered the book and will make an in depth report after it arrives. The following is based on the online preview of chapter 1. Witton’s writing style is entertaining and engaging. The book should have popular appeal on that level.

The cover portrays a magnificent crested Nyctosaurus at sunrise or sunset. Gorgeous!

Then things tumble.

Witton’s Table of Contents shows an embryo Pterodaustro with a very short rostrum, unlike any Pterodaustro I’ve ever seen. And I’ve seen the embryo. The rostrum extends nearly the entire length of the egg. An agreement with Laura Codorniú prohibits me from publishing the image until she does, but the reconstruction of the long-beaked embryo Pterodaustro is based on that tracing. As we learned earlier, pterosaurs grew isometrically, resembling their parents on hatching.

Witton’s Rhamphorhynchus image on page 2 portrays the infamous cruro/uropatagium, a membrane spanning the hind limbs and not including the tail. The image also includes the infamous deep chord wing membrane, for which there is no evidence whatsoever as the Sordes situation was falsified. Witton’s two Rhamphs also have much shorter wings than any Rhamphorhynchus I’ve ever seen. One of Witton’s wonders has brought its wrists (carpals) in close to the base of the neck, which is novel, at least, but kills the tension on the extensor tendon that keeps the wing membrane aerodynamic. As in birds, when the elbow flexes, the wing folds. Having the wings fold in flight isn’t bad. Birds do it all the time for a brief low drag rest. At least the feet are properly positioned in Wittons’ illustration.

Page 3 portrays several dozen pterosaurs doing the forelimb leap that is such a travesty and fantasy that I slap my head every time I see it again and again. It has become firmly entrenched. Gadzooks@!# what is the ptero-world coming to?

Page 4 has a fine picture of Pterodactylus antiquus, the first pterosaur known to science, with a big round head crest. Not quite ready to buy into that one quite yet. Some Pterodactylus did have a crest, but not that one.

Page 12 portrays a hypothetic pterosaur ancestor. It looks like Peteinosaurus with a short digit 4 leaping from a branch (using muscular hind limbs). The caption reads, “The fossil record has yet to reveal an “intermediate” between fully formed pterosaurs and possible ancestors, meaning we can only speculate on their anatomy and appearance.” And once again, pterosaur professors are casting a blind eye toward the hard evidence presented in the large reptile tree where dozens of ancestors are lined up. As you’ll recall, ludicrous as it sounds, we can even put turtles up as the closest known sisters to pterosaurs if we delete all the other sisters and candidates from the new Lepidosauromorpha, as demonstrated here. This just proves that pterosaur workers are actively avoiding the issue and the answer. But, I have to say, it’s a beautiful and evocative image that Witton has created, wrong though it may be.

Page 16 portrays three purported pterosaur ancestor/sisters, Sharovipteryx, Euparkeria and Scleromochlus. Witton calls Sharovipteryx an archosauromorph protorosaur, when it is neither. It is a fenestrasaur tritosaur lepidosaur, as we learned earlier. Euparkeria is closest to erythrosuchids, about as far from pterosaurs as one could imagine. Scleromochlus, shown hopping in Witton’s illustration with a dino quadrate leaning the wrong way, is a basal crocodylomorph. Witton strongly leans toward the “pterosaurs are ornithodires” direction despite the tiny hands and lack of pedal digit 5 in Scleromochlus.

Witton takes aim at my placing pterosaurs within the Squamata as the most unlikely hypothesis currently under consideration. See a recent post on this here. Witton writes, “There seems little similarity between the skulls of pterosaurs and the highly modified, mobile skulls of squamates or any similarity between the trunk and limb skeletons of each group.” Well, frequent readers will know that pterosaurs are tritosaur lepidosaurs, an outgroup clade to the two that make up the Squamata, the Iguania and the Scleroglossa. Pterosaurs are neither of these. Tritosaurs do not have the mobile skulls found in some squamates. They also don’t have the fused tarsals of squamates. They are distinct. Witton has whitewashed the tritosaur fenestrasaur hypothesis with this “red herring,” while virtually ignoring the fenestrasaurs, following in the less than noble footsteps of our colleague Dr. David Hone, whose exploits you can read about here. In chapter one, at least, Witton avoids any discussion of the pteroid and prepubis in Cosesaurus and other fenestrasaurs. Why should he ignore these key and readily observable traits? Dr. Pierre Ellenberger saw them first without recognizing their significance.

Page 17 Witton then discusses the possible protorosaur origins of pterosaurs, pointing to the shared trait of an elongated neck and forgetting the not-so-elongated neck of the basalmost  pterosaur, MPUM6009.  Witton points up the “fact” that protorosaurs lack an antorbital fenestra, but recent finds show that two protorosaurs had such a fenestra by virtue of convergence (really a side issue of little consequence). Witton finishes with protorosaurs by noting the body shapes are not at all pterosaurian, which is true.

Witton invites a closer look at Sharovipteryx and notices similarities to pterosaurs in the hind limbs and their membranes, but notes, “It’s hard to find other features that reliably link this animals with pterosaurs.” He may not have looked at the actual specimen as I have. Evidently he did not notice the ilium was anteriorly elongated, prepubes were present, more than five sacrals were present, the tail was attenuated with parallel chevrons, the bones were hollow, the feet have the same morphology as pterosaurs with a short metatarsal 5 and an elongated and robust p5.1 as obvious and compelling similarities. Once again, the blind eye rules. Witton reports that the Sharovipteryx skull lacks an antorbital fenestra and the foot is unlike that of any pterosaur. Where does he get his information? Certainly not from any sort of direct observation or adherence to the literature. Of course he doesn’t back up any of this with evidence. Witton concludes by noting that gliding with hind limbs is unique, failing to find parallels in Microraptor and the uropatagia of fenestrasaurs including pterosaurs. Sharovipteryx had fore limbs. Witton just doesn’t know or doesn’t show what they look like. But you can see them here.

Page 18 Witton prefers the archosauriform ancestry hypothesis due to the shared features of an antorbital fenestra and reduced bone counts in the fifth pedal digit, perforated lower jaws, and “many other anatomical similarities.” Really? Witton equates an evaporating pedal digit 5 in archosauriforms with the robust element in pterosaurs (and, of course he doesn’t count the ungual on the pterosaur digit). A robust pedal digit 5 is also found in Huehuecuetzpalli and all the tritosaur lepidosaurs that followed (except Macrocnemus and the drepanosaurs). Why doesn’t Witton consider these and put some study into them? The antorbital fenstra of archosauriforms is always (except for proterosuchians) surrounded by a fossa, a trait lacking in any pterosaurs.

Witton also prefers archosaurs as pterosaur sisters, and, in particular, Scleromochlus, despite the tiny hands that were, ironically, used to rule out Sharovipteryx. Evidently Witton prefers to have it both ways, so long as he stays within tradition. Witton lists fusion of the two proximal ankle bones to the shin (which does not occur in pterosaurs), reduction of the fibula (also in tritosaurs), the structure of the foot (actually more like that of tritosaur lizards like Cosesaurus, which retain an elongated pedal digit 5, which archosaurs lack), “several limb and hip proportions” (can Witton get even more vague here?) and the lack of bony scales along the back (then why is he ignoring those on Scleromochlus and Scutellosaurus).

Witton notes the shield-like pelves were different than in dinosaurs, but defends that by saying, “This may not be surprising, however, given, that pterosaur hindlinmbs were, uniquely among ornithodirans, used to support the wing in flight.” Utter rubbish!!! on the face of it and not pertinent to any phylogenetic discussion. You take the traits as they are and you let the computer decide where the taxa belong most parsimoniously. The “why” question or reason is never in play. By the way, similar pelves to pterosaurs can be found in fenestrasaurs, but these are ignored by Witton.

Witton writes, “arguments that basal pterosaurs were bipedal and digitigrade may be flawed” because basal ornithodires (aka: Asilisauruswhich bears no resemblance whatsoever to pterosaurs) were quadrupeds. This is far-reaching and totally bogus. I would be ashamed and would expect heavy chastisement having made such a comparison, especially after promoting bipedal Scleromochlus as a potential ancestor. But then Witton tops that bungle of reasoning by saying that Scleromochlus is “suspected of hopping about on plantigrade feet.” More fantasy! Few creatures, other than deer and horses, have feet more obviously digitigrade than Scleromochlus. Witton also ignores the known bipedal pterosaur footprints  (more here, here and more info here).

Page 21 Witton prefers an imagined hypothetical ancestor to a real one, and it glides from trees. Of course, this does nothing to explain the origin of flapping (because no gliders flap, unless they started off as flappers). Witton ascribes the mobility and length of the fifth toe to its use as a stabilizing tool, ignoring the fact that most tritosaurs from Tanystropheus to Sharovipteryx, have such a fifth toe, thus it cannot be developed for flight. Witton reports that the fifth toe, which is lateral, elongates to frame the medial membrane, which should strike you as odd and implausible. In reality the fifth toe is not connected to a membrane, except in Sharovipteryx, and each membrane trails each hind limb. They don’t cross to connect with each other.

Page 22 Witton reports that the hind limbs rotate out sideways to create efficient airfoils, but even that is fraught with error. One: Archosaurs can’t do this with their erect femurs. Two: Basal pterosaurs can’t do this either with their erect femurs. Raising the hind limbs to the horizon happens in later, more derived pterosaurs with a more sprawling femur.

Witton reports that during the evolution of pterosaurs that the fourth finger became so enlarged and unwieldy that it needed to be stowed away when grounded. We can all stow away our fingers by pressing them against our palms, but Witton ignores this. He also ignores the axial rotation of metacarpal 4 so that digit flexion puts digit 4 along the posterior rim of the hand, not the palmar side any longer. Witton reports ungual 4 was missing, since it was no longer necessary. We’ve seen so many several cases of ungual 4 present on pterosaurs that it needs to be considered universal.

Witton adds fibers to wing membranes as they need to be more sophisticated in their unsupported regions, ignoring that Cosesaurus had trailing fibers before it had wing membranes (Ellenberger 1993, Peters 2009).

With regard to flapping, our expert Dr. Witton reports, “At some point, manipulation of these wings in the vertical plane produced flapping, and self-propelled flight was achieved.” Gee, he makes it sound almost as if it was that easy. At ReptileEvolution.com and the PterosaurHeresies blog you learned the exact steps the exact taxa took to achieve flapping prior to the development of wings in pterosaurs, paralleling that same development in birds. So if Witton’s book leaves you unsatisfied and yearning for real answers, come see these websites and blogs.

Witton ascribes the development of flight muscles and bones to the ability of quadrupedal pterosaur ancestors to chiefly employ the forelimbs during leaps. He sort of leaves the larger hips and thighs out of the equation, evidently incapable of creating all the power necessary for a leap and leaving the unused arms in this bipedal model to do something else, like flap as a secondary sexual trait.

Dr. Witton does take the brave leap of including my published works in his reference list, something Dr. Unwin did not do in his less recent pterosaur book.

Let’s face it
If Dr. Witton does not even know what pterosaurs are (which he has acknowledged in his book), he has no business acting as an expert on pterosaurs and writing books about them. Unfortunately this is an acceptable trend continued by Dr. Unwin from Dr. Peter Wellnhofer. In chapter one Witton has already published too many errors. It’s too late in the game to fold ones’ hands and politely tell your readers, “Good question… we really don’t know. It’s one of the mysteries of paleontology.” There’s something called phylogenetic analysis that is guaranteed to give you an answer when you’re looking for an ancestor. However, you’ll have to include at least a few of the right taxa (among the tritosaurs in this case), to get close to the right answer. If you’re looking for the ancestors of pterosaurs, they’re right here in one place.

We’ll look at other Witton chapters in the future. But this one on pterosaur origins really irks me. It’s rather embarrassing that this sort of crap (a complete avoidance of certain data) is still being circulated. But I _do_ love the artwork.

References
Ellenberger P 1993. Cosesaurus aviceps . Vertébré aviforme du Trias Moyen de Catalogne. Étude descriptive et comparative. Mémoire Avec le concours de l’École Pratique des Hautes Etudes. Laboratorie de Paléontologie des Vertébrés. Univ. Sci. Tech. Languedoc, Montpellier (France). Pp. 1-664.
Peters D 2009. A reinterpretation of pteroid articulation in pterosaurs. Journal of Vertebrate Paleontology 29: 1327-1330.
Witton M. 2013. Pterosaurs. Princeton University Press. 291 pages.