Hamipterus egg accumulation: Wang et al. 2017

Earlier
here and here we looked at the 3-D eggs of Hamipterus, a basal ornithocheirid from Early Cretaceous China. The eggs are scattered in and amongst a wide size/age range of disarticulated, but 3-D fossils. So, according to the authors, the eggs were buried, then bones and eggs were transported by storms, as if bulldozed. No embryos were reported from those eggs. No explanation why the pterosaurs did not fly away in the face of the storm, nor why more sediment wasn’t packed on the buried eggs during the storm.

Today
comes news from this expanding treasure trove site with embryo bones at several stages of development in 16 eggs out of hundreds! That’s good news because full-term embryos (= hatchlings) are identical to parents and eggs keep all the bones from an individual in a neat little package so we can finally put together what Hamipterus looked like.

But that’s not the picture the authors paint.
They said, “some bones lack extensive ossification in potentially late-term embryos, suggesting that hatchlings might have been flightless and less precocious than previously assumed.” Point-by-point:

  1. No nests were found.
  2. 215+ eggs were found
  3. Eggs appeared in moderate size variation
  4. The large number of accumulated eggs (Fig. 1) indicates they were laid by different females
  5. Some were subjected to differential water uptake during transport
  6. Internal content(?) observed in 42 eggs, 16 had embryos
  7. Bones not concentrated on the bottom half of the egg, as in dinosaurs
  8. No embryo is complete. One to several bones only in each of the 16 eggs.
  9. No teeth found in embryos.
  10. The most complete embryo had a lower jaw of 17mm. That’s 4% the size of the largest adult when other full-term pterosaur embryos are 12.5% (1/8) at hatching. So these were not full-term embryos ready to hatch.
  11. In a 2.2m section, eight layers with pterosaur bones have been identified, four of which show egg concentrations in a vertical distance of 1.4 m.

The authors note and conclude:
“This suggests that the hind limbs have developed more rapidly compared to the forelimbs and might have been functional right after the animal hatched. Thus, newborns were likely to move around but were not able to fly, leading to the hypothesis that Hamipterus might have been less precocious than advocated for flying reptiles in general (6) and probably needed some parental care.”

No. Think again.
Pterosaur mothers carried their eggs inside their bodies until just before hatching. That gives their babies warmth and protection until they are ready to hatch. They could do this because they are lepidosaurs, as phylogenetic analysis AND egg shell thickness and pliability tells us.

Figure 1. From Wang et al. 2017, a pterosaur egg and bone accumulation. Eggs float. So do hollow pterosaur bones.

Figure 1. From Wang et al. 2017, a pterosaur egg and bone accumulation. Eggs float. So do hollow pterosaur bones.

Sedimentology report:
“This sedimentological data, associated with the exceptional quantity of eggs and bones, indicate that events of high energy such as storms have passed over a nesting site, causing the eggs to be moved inside the lake where they floated for a short period of time, becoming concentrated and eventually buried along with disarticulated skeletons.”

Bottom line and biggest problem:
The authors assume the eggs were laid. That’s because they think pterosaurs are archosaurs. Birds and crocs are archosaurs and they lay their eggs at an early stage of fertilization. Lepidosaurs wait to lay their eggs, sometimes until the moment before hatching.

Alternative hypothesis:

  1. Mass death of several year-classes of pterosaurs on beach due to lake burping deadly carbon dioxide. That stops the parents from flying away.
  2. Desiccation and insect decomposition reveals eggs inside of female skeletons. This takes just a few days to a week and allows skeletons to easily separate into individual bones (Fig. 1)
  3. Later rising waters (storms optional, melting snow pack will do), overwhelms beached skeletons and exposed eggs. Even a few extra inches of water would be enough for this.
  4. Eggs float. So do pterosaur bones
  5. Wind/ripples push eggs and bones together back against beach bank corner where they accumulate. (This happened several times over dozens to hundreds of years, but not annually.)
  6. Water recedes leaving eggs en masse along with settling disarticulated individual bones of parents and kin
  7. Burial process is later completed with airborne or waterborne sediments overwhelming the bones and eggs in situ.
  8. To point #3 above: moderate egg size variation, we also see this in the chicken eggs we get at our local grocery, but pterosaurs kept growing throughout their lives and larger ones would tend to lay bigger eggs, though this has not been conclusively demonstrated, it seems broadly logical.

Evidently
the lake burping did not always coincide with the pterosaurs flocking together. But it happened four times to a portion of the flock, perhaps over hundreds of years, and evidently at ‘the back of the room where bad things happen’.

References
Wang X and 16 co-authors 2017. Egg accumulation with 3D embryos provides insight into the life history of a pterosaur. Science 358:1197–1201.

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Pterodaustro embryo goes under DGS

Codorniú et al. 2017.
bring us detailed images of a wonderful Pterodaustro embryo (MIC-V246, Early Cretaceous; Chiappe et al. 2004) we’ve seen before. Here (Figs. 1–3) we’ll take a closer look at the fossil itself using DGS to untangle it. This is the same embryo first described by Chiappe et al. 13 years ago. Why did it take so long to bring out these details? There’s nothing really new here, no revision to their original description.

Figure 1. Pterodaustro embryo from Codorniú et al. with DGS color overlays to graphically segregate bones from one another.

Figure 1. Pterodaustro embryo from Codorniú et al. with DGS color overlays to graphically segregate bones from one another. Compare original reconstruction here to that in Figure 3.  Note the broken tibiae. Look how much better color over a photo works than pen and ink.

Turn out
the embryo is far less disarticulated than originally considered. Several bones are newly identified here (Fig. 2).

Figure 2. Original interpretations (2 frames black/white) vs. new interpretations (color).

Figure 2. Original interpretations (2 frames black/white) vs. new interpretations (color).

What Codorniú et al identified as

  1. ascending process of the premaxilla (pmx) is here a right metacarpal 4 (mc4)
  2. right wing phalanx 4 (wp4) is here a maxilla (mx) and premaxilla (pmx)
  3. left metacarpal 4 (mc4) is here a proximal radius (ra)
  4. left coracoid (co) is here a left metacarpal 4 (mc4)
  5. dorsal vertebrae (dv) and ribs at top is here identified as cervical vertebrae and left metacarpals 1–3).
  6. jugal (ju) and maxilla (mx) are here identified as mandible and palatal elements, but there’s also a short jugal in there below the sclerotic ring
  7. right metacarpal 4 (mc4) is here identified as a left scapula
  8. right wing phalanx 1 (wp1) is here identified as a left coracoid
  9. right wing phalanx 2 (wp2) is here identified as a dentary portion + left wp3 (4.3).
  10. gastralia (ga) are here identified as left dorsal ribs with tiny gastralia below
  11. caudal vertebrae (ca) here identified as possible sacral ribs

Parts missing from the Codorniú et al. tracing, but recovered using DGS include

  1. dozens of teeth
  2. prepubes
  3. pteroids
  4. carpals
  5. sternal complex
  6. finger phalanges including wing tip ungual (4.5)
  7. sclerotic rings
  8. cranial bones (parietals + occipitals)
  9. caudal vertebrae
  10. proximal femora
Figure 2. Same as in figure 1 with elements segregated.

Figure 3. Same as in figure 1 with elements segregated. Just a little bit of eggshell is preserved and it is very thin.

Digital Graphic Segregation (DGS)
is the perfect tool to interpret a fossil such as this. Embryos are encased in eggshell. That means every bone is retained in a small area, undisturbed by invaders or currents. Most embryos are going to be articulated, barring violent shaking, which could happen if the egg was tumbled and/or dropped. Some bones will just barely appear below others.

Try DGS yourself.
First color the bones that are easy to identify. Segregate the wings from other skeletal elements. Then color in all the remaining bones, including those that barely appear below other elements. Finally, lift your colors to another file and push them around to create a reconstruction. Check your work for errors and repeat.

Figure 3. Rough reconstruction using color tracings. Note the elongate jaws and small eye, documenting isometric growth in this pterosaur, as in all others where this can be seen.

Figure 4. Rough reconstruction using color tracings. Note the elongate jaws and small eye, documenting isometric growth in this pterosaur, as in all others where this can be seen. Note the relatively short tibia. This could be longer, but pars are hidden by the feet and other bones.

The thin egg shell
is similar to those in lepidosaurs because pterosaurs are lepidosaurs. As lepidosaurs mothers can carry their eggs until just before hatching, taking advantage of her warmth and her ability to avoid predators. Please ignore hypotheses that suggest pterosaurs buried their eggs. No fragile hatchling wants to dig out of a deep steaming pile of wet, rotting leaves filled with insects and centipedes. They’re not like husky little sea turtles. Baby pterosaurs want to dry out, open up their tiny wings and fly!

Pterodaustro adult with manual digit 3 repaired.

Figure 4. Pterodaustro adult with manual digit 3 repaired.

At one-eigth the size of a typical adult (which also vary in size)
embryo Pterodaustro was similar in proportions, but with shorter tibiae, smaller feet, more gracile jaws and other minor differences. With such proportions, there is nothing to prevent hatchling Pterodaustro from flying.

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 embryo compared to post-hatchling. Note the right specimen is a little larger with more robust wing finger phalanges and a larger sternal complex.

We looked at other pterosaur embryos
here, here, here, here, and here.

Bone fusion patterns in pterosaurs
Codorniú et al. discuss the lack of scapulacoracoid (s/c) bone fusion in the embryo and considered that an ontogenetic trait. A long time ago, bone fusion was phylogenetically mapped on the pterosaur cladogram. No current embryos have s/c bone fusion, but then neither do any of their adults or sisters. What we need is to find an embryo of a pterosaur genus with a fused s/c as an adult. We have not found one yet. As lepidosaurs, pterosaurs follow lepidosaur bone fusion patterns (Maisano 2002), which are phylogenetic, not ontogenetic. Fused bones keep growing. So do unfused bones into adulthood.

It would be great
if this embryo finally puts to rest the invalid hypothesis of allometric growth in pterosaurs promoted by Bennett and many Bennett followers who considered small, short rostrum, large-eyed Solnhofen pterosaurs the juveniles of larger, longer-snouted specimens. Zhejiangopterus and Pterodaustro growth series provide further strong evidence against the invalid allometric growth hypothesis. Pterosaur workers: GET OVER IT! IT’S WRONG!

References
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, Chiapped L and Rivarola D 2017. Neonate morphology and development in pterosaurs: evidence from a Ctenochasmatid embryo from the Early Cretaceous of Argentina. From: Hone DWE, Witton MP and Martill DM (eds) New Perspectives on Pterosaur Palaeobiology. Geological Society, London, Special Publications, 455, https://doi.org/10.1144/SP455.17
Maisano JA 2002. Terminal fusions of skeletal elements as indicators of maturity in squamates. Journal of Vertebrae Paleontology 22: 268–275.

 

 

Dr. David Unwin on pterosaur reproduction – YouTube

Dr. David Unwin’ talk on pterosaur reproduction 
was recorded at the XIV Annual Meeting of the European Association of Vertebrate Palaeontologists, Teylers Museum, Haarlem, Netherlands and are online as a YouTube video.
Dr. Unwin is an excellent and engaging speaker.
However, some of the issues Dr. Unwin raises have been solved at www.ReptileEvolution.com
The virtual lack of calcite in pterosaur eggs were compared to lepidosaurs by Dr. Unwin, because pterosaurs ARE lepidosaurs.  See: www.ReptileEvolution.com/reptile-tree.htm
Lepidosaurs carry their eggs internally much longer than archosaurs, some to the point of live birth or hatching within hours of egg laying. Given this, pterosaurs did not have to bury their eggs where hatchlings would risk damaging their fragile membranes while digging out. Rather mothers carried them until hatching. The Mrs. T external egg was prematurely expelled at death, thus the embryo was poorly ossified and small.
Dr. Unwin ignores the fact that hatchlings and juveniles had adult proportions as demonstrated by growth series in Zhejiangopterus, Pterodaustro and all others, like the JZMP embryo (with adult ornithocheirid proportions) and the IVPP embryo (with adult anurognathid proportions).
Dr. Unwin also holds to the disproved assumption that all Solnhofen sparrow- to hummingbird-sized pterosaurs were juveniles or hatchlings distinct from any adult in the strata. So they can’t be juveniles (see above). Rather these have been demonstrated to be phylogenetically miniaturized adults and transitional taxa linking larger long-tailed dorygnathid and scaphognathid ancestors to larger short-tailed pterodactyloid-grade descendants, as shown at: www.ReptileEvolution.com/MPUM6009-3.htm
Thus the BMNH 42736 specimen and Ningchengopterus are adults, not hatchlings. And the small Rhamphorhynchus specimens are also small adults.

Rhamphorhynchus n28: unidentified food mass? or overlooked egg in the abdomen?

Figure 1. Rhamphorhynchus intermedius (n28) reconstructed.

Figure 1. Rhamphorhynchus intermedius (n28) reconstructed.

Rhamphorhynchus intermedius (Koh 1937, n28 in the Wellnhofer 1975 catalog) is a well preserved basal specimen (derived from the C3 specimen of Campylognathoides) with a mass inside of its torso, only part of which has been identified as a Solhnhofen fish (Figs. 2,3).

Figure 2. Wellnhofer 1991 illustrates the abdominal mass as part of a fish and other unidentified elements.

Figure 2. Wellnhofer 1991 illustrates the abdominal mass as part of a fish and other unidentified elements.

The rest of the abdominal mass
is unidentified (Fig. 2). Wellnhofer 1991 considered it food. Considering its shape, size and placement, I wonder if the posterior mass is actually a nearly full term egg (Fig. 3). I don’t think it makes much sense to consider such an abdominal mass as “unidentified food” when no other known specimen has a similar mass of undigested food. That would mean the stomach could expand to fill the abdomen. Usually the only thing that crowds out other organs and air sacs is an egg or a number of eggs in other reptile taxa. Check out this kiwi X-ray for an extreme example.

Figure 3. Skeletal elements of Rhamphorhynchus intermedius (n28) along with an ingested fish and what appears to be a possible egg.

Figure 3. Skeletal elements of Rhamphorhynchus intermedius (n28) along with an ingested fish and what appears to be a possible egg. Note the overlapping sets of gastralia. It looks like the jaws of the fish are displaced here. Wellnhofer did not see the large eyeball identified here. Note the left scapula and coracoid are inverted. There sternal complex has a similar unexpectedly bumpy texture as the purported egg.

Rhamphorhynchus intermedius
is a medium-sized Rhamphorhynchus nesting at the very base of the clade between the larger Campylognathoides and the smaller Bellubrunnu. Thus it is a transitional taxon, step one in an extreme example of phylogenetic miniaturization. No one understood this nesting prior to the phylogenetic analysis documented at ReptileEvolution.com because no one included a long list of Rhamphorhynchus specimens in analysis prior to or since. I’d like to encourage other pterosaur workers to do so and test this hypothesis of relationships.

Texture
The “egg” has an odd texture, but then so does the sternal complex. Not sure why.

Previous examples?
Two smaller examples of “something else” in the abdomen of – or just aborted from the abdomen – other specimens of Rhamphorhynhcus can be seen here and here.

References
Koh TP 1937. Unterscuchungen über die Gattung Ramphorhynchus. – Neues Jahrbuch Mineralogie, Geologie und Palaeontologie, Beilage-Band 77: 455-506.
Wellnhofer P 1975a-c. Teil I. Die Rhamphorhynchoidea (Pterosauria) der Oberjura-Plattenkalke Süddeutschlands. Allgemeine Skelettmorphologie. Paleontographica A 148: 1-33. Teil II. Systematische Beschreibung. Paleontographica A 148: 132-186. Teil III. Paläokolgie und Stammesgeschichte. Palaeontographica 149: 1-30.

wiki/Rhamphorhynchus

Another look at the tiny pterosaur, Nemicolopterus

Not content
with a fully resolved cladogram, I wanted higher Bootstrap scores at certain nodes to ascertain nesting pairs. So I reviewed the data for several taxa, among them Nemicolopterus. I found mistakes and oversights, nearly all of which more closely match Nemicolopterus to its much taller sister, Shenzhoupterus (Fig. 1) within the larger encompassing Germanodactylus/Tapejara clade.

Figure 1. Germanodactylus cristatus and members of the Shenzhoupteridae, Nemicolopterus and Shenzhoupterus.

Figure 1. Germanodactylus cristatus and members of the Shenzhoupteridae, Nemicolopterus and Shenzhoupterus.

When first announced
(Wang et al. 2008), Nemicolopterus was hailed as the smallest, or one of the smallest known pterosaurs. And it is. But there is one other that is only half as tall (Fig. 2) which we looked at in more detail yesterdayB St 1967 I 276.

Figure 2. Nemicolopterus has been described as the smallest pterosaur, but No. 6 in the Wellnhofer (1970) catalog was only half as tall.

Figure 2. Nemicolopterus has been described as the smallest pterosaur, but B St 1967 I 276, No. 6 in the Wellnhofer (1970) catalog was only half as tall.

An insitu tracing 
animated in a GIF movie reveals the bones segregated by digital layers (Fig. 3).

Figure 3. Two images of Nemicolopterus superimposed and traced with transparent colors. Note, not all of the shapes seen in photo 1 can be seen in photo 2. There appear to be extra tiny bones in the belly of this specimen.

Figure 3. Two images of Nemicolopterus superimposed and traced with transparent colors. Note, not all of the shapes seen in photo 1 can be seen in photo 2. There appear to be extra tiny bones in the belly of this specimen where all the ribs and gastralia have been accounted for. Click to enlarge. 

A displaced sliver of bone in the cheek
would appear to be the ventral portion of an orbit-dividing lacrimal, as in sister taxa. No one would identify this bone as such without phylogenetic bracketing. Hairlike soft tissue arising from the rostrum evidently precedes the rostral crest found in Shenzhoupterus. The free fingers remain unknown.

Phylogenetic miniaturization 
appears to be at work once again with Nemicolopterus, a tiny adult at the base of a major clade or two of pterosaurs in the large pterosaur tree.

References
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

wiki/Nemicolopterus

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