One more look at Rhamphorhynchus growth

Usually I avoid histological (bone microstructure) studies.
But here’s one that merits one more extended report based on its many incorrect assumptions and overlooked comparisons.

Summary of key facts in this long blog:

  1. both phylogenetically miniaturized adult pteros and mammals had juvenile-like “woven” bone texture
  2. Pterosaur embryos develop in utero and had adult proportions, so they could fly upon hatching
  3. Pterosaurs develop isometrically, thus immature pteros can only be identified in phylogenetic analysis (= when larger identical adults are known).

Prondvai et al. 2012 tested growth strategies in Rhamphorhynchus. As noted earlier, Prondvai et al. confused small adults with juveniles and hatchlings, not following the clear data that pterosaurs grow isometrically, not allometrically. Thus the morphological difference shown here (Fig. 1) are phylogenetic, not ontogenetic. Phylogenetic analysis supports this hypothesis.

Figure 1. Bennett 1975 determined that all these Rhamphorhynchus specimens were conspecific and that all differences could be attributed to ontogeny, That is clearly false as shown here and by phylogenetic analysis. Only the juvenile between the two largest specimens is a non-adult. Click to enlarge.

Figure 1. Bennett 1975 determined that all these Rhamphorhynchus specimens were conspecific and that all differences could be attributed to ontogeny, That is clearly false as shown here and by phylogenetic analysis. Only the juvenile between the two largest specimens is a non-adult. Click to enlarge.

 

Age of first flight
Prondvai et al. 2012 report,
The initial rapid growth phase early in Rhamphorhynchus ontogeny supports the non-volant nature of its hatchlings, and refutes the widely accepted ‘superprecocial hatchling’ hypothesis. We suggest the onset of powered flight, and not of reproduction as the cause of the transition from the fast growth phase to a prolonged slower growth phase. Rapidly growing early juveniles may have been attended by their parents, or could have been independent precocial, but non-volant arboreal creatures until attaining a certain somatic maturity to get airborne.” Prondvai et all did not realize they were examining small adult pterosaur specimens, not juveniles. So rapid growth was part of their growth strategy. More refutations relevant to the above statements follow.

Powered flight is one of the most energy-consuming locomotion types in tetrapods, therefore high growth rates and a superprecocial onset of the flying lifestyle in a highly developed hatchling are mutually exclusive developmental parameters. The validity of this simple trade-off model is supported by the fact that the only extant superprecocial fliers, the megapod birds have very low if not the lowest growth rates among extant birds.” Prondvai et al. ignore the fact that megapodes have their rapid growth phase inside the egg shell. Hatchling megapodes are relatively “very large with a wingspan up to half that of the adult).”  By contrast, pterosaurs hatch at 1/8 the height of the adult and 1/8 the wingspan.

In support of supreprecocial flight…
pterosaur hatchlings had adult proportions. Tiny adults, the size of sparrows and hummingbirds, had larger pterosaur proportions. The smallest pterosaur that Prondvai et al. tested had wing tips that extended way over their heads when folded and quadrupedal (Figs. 1, 2). We’ve seen the short wings of flightless pterosaurs. Hatchlings of volant taxa don’t have short wings. Tiny adult pterosaurs may have ‘rapidly growing” bone microstructure because they matured quickly, reproduced as often as possible then died early, like tiny mammals do. More on this below:

Sexual maturity vs. size:
Prondvai et al. report, “According to the hypothesis presented here, the onset of powered-flight in Pterodaustro occurred after attaining 53% of adult size. Here we prefer the hypothesis that bone growth is slowed down by the initiation of a new, and much more energy consuming locomotory activity, namely powered flight.” Not by coincidence, this is the size that Chinsamy et al. (2008) determined that sexual maturity was attained. After observing the morphology of the embryo Pterodaustro, which matches the morphology of the adult, there is no supporting evidence for the Prondvai et al. hypothesis.

Archosaur vs. lepidosaur
Prondvai et al. do not consider the growth strategies and histology of lepidosaurs, only archosaurs. So they are making comparisons to the wrong clade. Pterosaurs nest within the Lepidosauria. Growth patterns in lepidosaurs are distinct and do not follow archosaur growth patterns (Masisano 2002). But this may not be the key factor in observed differences.

Chinsamy and Hurum 2006
looked at the basal lepidosaur, Gephyrosaurus. “The [bone] compacta consists of essentially parallel−fibred bone tissue interrupted by several lines of arrested growth (LAGs). The first LAG visible from the medullary cavity appears to be a hatchling line with its more haphazardly oriented, globular-shaped, osteocyte lacunae.”  This was not a phylogenetically miniaturized taxon even though it was a basal lepidosaur.

More to the point
Chinsamy and Hurum 2006 also looked at the basal and phylogenetically miniaturized mammal, Morganucodon. They report on, “distinct woven bone tissue with large, irregularly oriented osteocyte lacunae and several primary osteons. No secondary osteons were visible, though several enlarged erosion cavities are evident in the compacta. In the same section, it appears that substantial endosteal resorption had occurred, and parallel−fibred bone tissue is evident only in a localized area peripherally. This area includes several rest lines, which indicate pauses in the rate of bone formation, and hence, pauses in growth.” Perhaps these pauses indicate a lifespan of “several” years. Note the “woven bone” texture description.

Figure 1. Several tiny Rhmphorhynchus adults, among them is the n7 specimen tested by Prondvai et al. and considered a juvenile by them.

Figure 2. Several tiny Rhmphorhynchus adult sister taxa, among them is the n7 specimen tested by Prondvai et al. and considered a juvenile by them shown here about 7/10 of in vivo size. As you can see, these pterosaurs do not appear to have any impediments to flapping and flying. However their tiny hatchlings would probably not have flown based on their high surface/volume ratio. The adults had juvenile traits due to phylogenetic miniaturization.

The smallest sampled Rhamph bone microstructure
Prondvai et al. report about the tiny Rhamph, BSPG 1960 I 470a, (n7 in the Wellnhofer 1970 catalog, Fig. 2): “A thin layer of lamellar bone of endosteal origin rims the medullar [central] cavity. There seem to be only a few longitudinally oriented vascular canals, but these have rather large diameter in relation to the overall thickness of the cortex. The bone matrix is typically woven with some poorly defined, immature primary osteons, hence the majority of the cortex does not show the mature fibrolamellar pattern yet. The osteocyte lacunae are large and plump throughout the cortex, and possess an extremely well-developed system of dense, radially oriented canaliculi implying extensive communication and nutrient-exchange between the osteocytes. No LAGs or any other growth marks can be observed.”  Maybe LAGs were never present in this taxon if it lived for just a short time. Remember, we’re talking about phylogenetic miniaturization here.  If the small precocial Rhamphorhynchus specimens were maturing quickly and laying eggs early, they likely followed the life patterns of other tiny tetrapods, like Morganucodon (above) and died early, perhaps living only one or two years, not five or more as in mid-sized pterosaurs.

Note: Like Morganucodon (above) the phylogenetically miniaturized mammal, 
the bone structure in the smallest tested Rhamphorhynchus is described as “woven”.

Age vs size:
Prondvai et al. report, “The ontogenetic validity of the smallest size category of Bennett is clearly supported by the overall microstructure found in the bones of the three small specimens.” Unfortunately, without a phylogenetic analysis, Prondvai et al. did not realize that the smallest specimens were small due to phylogenetic miniaturization. Their ancestors were larger. Thus small Rhamphs retained juvenile and embryonic traits into adulthood, including the typical short rostrum and smaller wings. These traits also included juvenile “woven” bone tissue. Essentially these tiny pterosaurs were precocious sexually active adults in the former juvenile phase of development.

Precocial hatchling?
Prondvai et al. report, “Superprecocial embryos require substantial amount of nutrients stored in their eggs to reach an advanced level of somatic maturity state by the time the embryo hatches. If the egg volume of Darwinopterus was relatively as low as that of squamates, then how could it have contained so much yolk as to cover the energy requirements of an extremely well-developed, volant hatchling?” Prondvai are assuming that pterosaur eggs developed outside the uterus. As lepidosaurs, pterosaur embryos developed inside the uterus and the super thin eggshell was deposited last. Thus they could “cover the energy requirements.”

Apparently Prondvai et al. are not looking
at verified pterosaur hatchlings (in eggs), which are identical in morphology to adults. In some cases large embryos can be larger than small adult sister taxa! The Prondvai team know that the tiny Rhamps don’t have the same morphology as the medium or big rhamphs. Unfortunately, and this is a continuing problem… they don’t realize those changes are phylogenetic, not ontogneric.

With similar proportions of bone and muscle,
but at 1/8 as tall and therefore (8 cube rooted or) 1/512 as massive, juvenile pterosaur bone tissue would have been strong enough for sustained flight in such lightweight specimens. But that overlooks reality, where the specimens Prondvai are looking at are in fact tiny adults with juvenile bone structure, as in Morganucodon. We don’t know where small, medium and large Rhamphorhynchus laid its eggs, which were likely ready to hatch shortly after deposition. We don’t have any hatchling Rhamphorhynchus fossils. Hatchlings of the small and tiny adults would have been in danger of desiccation (high surface area/volume ratio), so we can presume they grew up in moist environs. Unfortunately Prondvai et al. did not test the one verified juvenile among in the Rhamphorhynchus clade, NHMW 1998z0077/0001 (Fig. 3), the Vienna specimen. No one thinks this juvenile could not fly based on its age/relative size.

Figure 1. Two specimens attributed to Rhamphorhynchus longiceps along with a third specimen that nested with the larger of the two with identical scores, thus identifying it as a juvenile R. longiceps.

Figure 3. Two specimens attributed to Rhamphorhynchus longiceps along with a third specimen, NHMW 1998z0077/0001, that nested with the larger of the two with identical scores, thus identifying it as a juvenile R. longiceps. No one thinks this Rhamph could not fly, despite its young age.

To their credit, Pronvai et al. suggest (following a hypothesis first presented here): “Alternatively, Rhamphorhynchus hatchlings could have been precocial to the effect that they could have left their nests immediately after hatching, but they must have been exclusively terrestrial or rather arboreal. They could have clambered around quadrupedally on the branches of trees feeding themselves with smaller invertebrates or vertebrates without any parental contribution.”

No universal growth strategy in pterosaurs
Prondvai et al. report, “In the light of the histological results it becomes evident that there is no universal pattern in the growth strategy of pterosaurs.” I am concerned that this conclusion was made without the the benefit of a phylogenetic analysis and without knowledge of phylogenetic miniaturization in the clade.

To their credit
Prondvai et al. report, “In contrast to Bennett’s  suggestion, the second size category of Rhamphorhynchus does not only include subadult but also adult specimens, hence it cannot be used as an indicator of real ontogenetic stage.”

References
Chinsamy A, Codorniu ́ L, Chiappe L 2008. Developmental growth patterns of the filter-feeder pterosaur, Pterodaustro guinazui. Biol Lett 4: 282–285.
Chinsamy A and Hurum JH 2006. Bone microstructure and growth patterns of early mammals. Acta Palaeontologica Polonica 51 (2): 325–338.
Maisano JA 2002. Terminal fusions of skeletal elements as indicators of maturity in squamates. Journal of Vertebrae Paleontology 22: 268–275.
Prondvai E, Stein K, O0 si A, Sander MP 2012. Life History of Rhamphorhynchus Inferred from Bone Histology and the Diversity of Pterosaurian Growth Strategies. PLoS ONE 7(2): e31392. doi:10.1371/journal.pone.0031392
Sekercioglu C 1999. Megapodes: A fascinating incubation strategy. Online article. 

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

Another look at the smallest adult pterosaur – AND its hatchling

Earlier we looked at the smallest adult pterosaur, B St 1967 I 276 or No. 6 in the Wellnhofer (1970) catalog. Here (Fig. 1) it is compared to an adult leaf chameleon, Brookesia micro, one of the smallest living lizards and to the Bee hummingbird, one of the smallest living birds. Also shown are their hatchlings and eggs.

Figure 1. The smallest of all adult pterosaurs, B St 1967 I 276 or No. 6 in the Wellnhofer (1970) catalog compared to scale with the living leaf chameleon (Brookesia micro) sitting on someone's thumb. Also shown are hypothetical eggs and hatchlings for both. These lepidosaurs had tiny eggs and hatchlings.

Figure 1. The smallest of all adult pterosaurs, B St 1967 I 276 or No. 6 in the Wellnhofer (1970) catalog compared to scale with the living leaf chameleon (Brookesia micro) sitting on someone’s thumb. Also shown are hypothetical eggs and hatchlings for both. These lepidosaurs had tiny eggs and hatchlings, relatively larger in the chameleon, based on pelvis size and average 1/8 size for other pterosaur hatchlings.

 

Traditional paleontologists
don’t buy the argument that No. 6 was an adult, even though it is much larger than the smallest lizard and about the size of the smallest bird. Worse yet, they refused to test it in phylogenetic analysis. So, the  impasse remains.

Figure 2. Smallest known bird, Bee hummingbird, compared to smallest known adult pterosaur, No. 6 (Wellnhofer 1970). Traditional workers consider this a hatchling or juvenile, but in phylogenetic analysis it does not nest with any 8x larger adults.

Figure 2. Smallest known bird, Bee hummingbird, compared to smallest known adult pterosaur, No. 6 (Wellnhofer 1970). Traditional workers consider this a hatchling or juvenile, but in phylogenetic analysis it does not nest with any 8x larger adults. This image is slightly larger than life size at 72dpi. Note the much smaller eggs produced by the tiny pterosaur. 

 

Pictures tell the tale.
You can see for yourself. No. 6 is substantially smaller than other tiny pterosaurs just as the bee hummingbird is substantially smaller than other hummingbirds.The hatchling was substantially smaller than both the leaf chameleon and bee hummingbird hatchlings based on their larger egg size/pelvis opening.

Earlier we looked at isometric growth in several pterosaurs, with hatchlings matching adults in morphology. Earlier we also took note of the danger of desiccation to hatchling pterosaurs until they reached a certain size/volume, so they probably roamed the leaf litter, which is probably when pterosaurs became quadrupeds and developed elongate metacarpals 4x.

References
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.
Wellnhofer P 1970. 
Die Pterodactyloidea (Pterosauria) der Oberjura-Plattenkalke Süddeutschlands. Abhandlungen der Bayerischen Akademie der Wissenschaften, N.F., Munich 141: 1-133.

wiki/Pterodactylus

Pterodaustro isometric growth series

Tradtional paleontologists think pterosaur babies had a cute short rostrum that became longer with maturity and a large orbit that became smaller with maturity (Fig. 1). This is a growth pattern seen in the more familiar birds, crocs and mammals.

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.

Unfortunately
these paleontologists ignore the fossil evidence (Figs 2, 3). These are the data deniers. They see things their own way, no matter what the evidence is. The data from several pterosaur growth series indicates that hatchlings had adult proportions in the skull and post-crania. We’ve seen that earlier with Zhejiangopterus (Fig. 2), Tapejara, Pteranodon, Rhamphorhynchus and others. Still traditional paleontologists ignore this evidence as they continue to insist that small short rostrum pterosaurs are babies of larger long rostrum 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.

As readers know,
several pterosaur clades went through a phase of phylogenetic miniaturization, then these small pterosaurs became ancestors for larger clades. Pterosaurs are lepidosaurs and they grow like lepidosaurs do, not like archosaurs do.

Today we’ll look at
the growth series of Pterodaustro (Fig. 1), previously known to yours truly only from adults and embryos. Today we can fill the gaps with some juveniles.

This blog post is meant to help traditional paleontologists get out of their funk.

A recent paper
on the braincase of odd South American Early Cretaceous pterosaur Pterodaustro (Codorniú et al. 2015) pictured three relatively complete skulls from a nesting site (Fig. 1). I scaled the images according to the scale bars then added other available specimens.

Figure 1. Pterodaustro skulls demonstrating an isometric growth series. One juvenile is scaled to the adult length. One adult is scaled to the embryo skull length. There is no short rostrum and large orbit in the younger specimens.

Figure 1. Pterodaustro skulls demonstrating an isometric growth series. One juvenile is scaled to the adult length. One adult is scaled to the embryo skull length. There is no short rostrum and large orbit in the younger specimens. If you can see differences in juvenile skulls vs. adult skulls, please let me know. All these specimens come from the same bone bed.

You can’t tell which skulls are adults or juveniles
without scale bars and/or comparable specimens. As we established earlier, embryos are generally one-eighth (12.5%) the size of the adult. Pterodaustro follows this pattern precisely.  We have adults and 1/8 size embryos and several juveniles of intermediate size.

No DGS was employed in this study.

If you know any traditional paleontologists, 
remind them that the data indicates that pterosaurs matured isometrically, like other  lepidosaurs. Those small, short rostrum specimens, principally from the Late Jurassic Solnhofen Formation, are small adults, transitional from larger ancestors to larger descendants. Tiny pterosaurs experiencing phylogenetic miniaturization(as in birds, mammals, crocs, turtles, basal reptiles, and many other clades) that helped their lineage survive while larger forms perished, Sadly, no tiny pterosaurs are known from the Late Cretaceous when they all became extinct.

References
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, Paulina-Carabajal A and Gianechini FA 2015.
 Braincase anatomy of Pterodaustro guinazui, pterodactyloid pterosaur from the Lower Cretaceous of Argentina. Journal of Vertebrate Paleontology, DOI:10.1080/02724634.2015.1031340

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

 

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)

Pterosaur tails tell tales… Unwin et al. 2014 JVP abstract

Unwin et al. (2014)
describe an increasing number of tail vertebrae in a purported ontogenetic series (hatchling to juvenile to adult in a series of purported Darwinopterus specimens.) Although this is unheard of elsewhere among vertebrates, Unwin et al. link this trait to the origin of pterodactyloid-grade pterosaurs. And it should be mentioned that Unwin et al. are the only workers who nest darwinopterids basal to pterodactyloids. Andres nests anurognathids there. Kellner nests Rhamphorhynchus there. I nest tiny dorygnathids and scaphognathids there by convergence (e.g. Fig. 1) four times.

From the Unwin et al. 2104 abstract:
“The evolution of pterodactyloids from basal pterosaurs in the Early-Middle Jurassic involved a complex series of anatomical transformations that affected the entire skeleton. Until recently, almost nothing was known of this major evolutionary transition that culminated in the Pterodactyloidea, a morphologically diverse and ecologically important clade that dominated the aerial environment throughout the mid-late Mesozoic. The discovery of Darwinopterus, a transitional form from the early Late Jurassic of China, provided the first insights into the sequence of events that gave rise to the pterodactyloid bauplan and hinted at an important role for modularity, but was largely silent regarding the anatomical transformations themselves, or the evolutionary mechanism(s) that underlay them. A series of recent finds allowed us to construct a complete postnatal growth sequence for Darwinopterus. By comparing this sequence with those for Rhamphorhynchus and Pterodactylus, pterosaurs that phylogenetically bracket Darwinopterus, it is possible to map key anatomical transformations such as the evolution of the elongate, complex tail of basal pterosaurs into the short, simple tail of pterodactyloids. In Darwinopterus hatchlings the tail is shorter than the dorsal-sacral series (DSV) and consists of around 18 simple vertebral ossifications. The tail is longer (1-2 x DSV) in juveniles and has a normal complement of about 30 caudals, but only reaches its full length (2-3 x DSV) and complexity in adults. Basal pterosaurs largely conform to this pattern, although some species, including Rhamphorhynchus, have longer tails with up to 40 caudals. Generally, the tail of adult pterodactyloids, including Pterodactylus, resembles that of Darwinopterus hatchlings (≤18 ossified vertebrae; tail ≤0.7 x DSV; vertebrae simple, blocky), but occasionally develops a little further (e.g. in Pterodaustro) corresponding to the condition seen in early juveniles of Darwinopterus and paralleling the developmental pattern observed in long-tailed pterosaurs. The short tail of adult pterodactyloids, and anurognathids, basal pterosaurs that also have relatively short tails, appears to be neotenic, resulting from a sharp decrease in growth rate compared to the rest of the skeleton. This mechanism, heterochrony acting upon a distinct anatomical module to effect a large-scale morphological transformation, can be applied to other modules to generate the derived features (e.g. elongate neck and metacarpus, reduced fifth toe) that typify the pterodactyloid bauplan.”

Problem #1
Among professional pterosaur workers, only Unwin et al. nest Darwinopterus as the stem pterodactyloid. No one else does. Andres nests anurognathids with pterodactyloids. Kellner nests Rhamphorhynchus with pterodactyloids. Readers of this blog and reptile evolution.com know that when you add the sparrow- to hummingbird-sized Solnhofen pterosaurs, you get four clades of pterodactyloid-grade pterosaurs.

Figure 1. Scaphognathians to scale. Click to enlarge.

Figure 1. Scaphognathians to scale. Click to enlarge.

Problem #2
Are the specimens truly juvenile Darwinopterus? Or do they represent smaller genera or species, perhaps closely related, or not? Currently no two Darwinopterus specimens are conspecific. No two are identical. See them here. By comparing purported Rhamphorhynchus and Pterodactylus juveniles to putative adults I’m afraid Unwin et al. are playing with a pack of Jokers. Those smaller specimens are distinct species and genera, as recovered in the large pterosaur tree. Everyone should know by now that pterosaur juvenile pterosaurs are isometric matches to their adult counterparts, from several well-known examples. Any differences in Darwinopterus likewise mark phylogenetic, not ontogenetic differences.

Problem #3
Rhamphorhynchus and Pterodactylus only phylogenetically bracket Darwinopterus if the inclusion set is reduced to these three taxa. Otherwise they nest several nodes away from each other with lots of intermediate taxa as you can see here.

Problem #4
Unwin et al. claim the caudal count increases with maturity in Darwinopterus (18 in hatchlings, 30 in juveniles and adults). Put these into a cladogram and they probably become disparate taxa. Where else does the vertebral count nearly double during ontogeny? Nowhere. Those caudal counts for the larger specimens have to be estimates. Not every tail is complete. It appears as if the caudal count could vary among the larger specimens as well.

Problem #5
I see no mention of a phylogenetic analysis with regard to the various Darwinopterus specimens. This is a problem as Unwin et al. do not want to test their observations with the only method known to lump and split taxa. In the large pterosaur tree IVPP V 16049 nests with YH2000. 41H111-0309A nests with ZMNH M 8782. All four Darwinopterus taxa nest as a sister clade to Kunpengopterus + Archaeoistiodatylus and this combined clade is a sister to Wukongopterus, then the PMOL specimen of Changchengopterus, then Pterorhynchus. This major clade nests between Dorygnathus and Scaphognathus, both of which ultimately give rise to the two pairs of basalmost pterodactyloids.

Possible Solution 
I noted earlier that the Darwinopterus clade left no descendants. They also did not produce any small taxa like Dorygnathus and Scaphognathus did. Other workers thought the smaller Scaphognathus specimens were juveniles, despite the morphological differences. I can only wonder if the same situation is happening in the Darwinopterus clade? Perhaps what the Unwin team found are the smaller specimens previously missing from their clade branch. Even so, and sadly, this clade was not able to survive into the Cretaceous, small or not, because no known Cretaceous pterosaurs share darwinopterid traits. They are all accounted for with presently known tiny ancestors.

References
Unwin D, Lü J-C, Pu H-Y, Jim X-S  2014. Pterosaur tails tell tails of modularity and heterochrony in the evolution of the pterodactyloid bauplan. JVP 2014 abstracts

Caiuajara dobruskii – new tapejarid pterosaur bone bed

We’ll call this:
“When discovery confirms heretical hypotheses.”

Figure 1. Caiuajara adult skull. Color bones added.

Figure 1. Caiuajara adult skull. Color bones added. Their premaxillary crest also includes the nasal. Blue = jugal. Yellow = missing teeth. Fo = foramina. Wonder if those represent ancient tooth sockets? For now they are blood vessel holes. Exp = ventral expansion of premaxilla, but it’s really the nasal. That’s where the descending process drops on certain other pterosaurs.

Another pterosaur bone bed,
this time with subadults and juveniles (no eggs or hatchlings) of a new tapejarid, Caiuajara dobruskii (Manzig et al. 2014). Contra traditional paradigms, there is no indication of a large orbit and short rostrum in juveniles (confirming earlier posts here and at reptileevolution.com. Yes, the crest developed in adults, because it wouldn’t have fit inside the eggshell! At least 47 individuals here. Smallest were twice the size of hatchlings, one quarter the size of adults.

Also,
you can’t tell the females from the males. All had crests.

Figure 2. from Manzig et al. 2014. Note the lack of change in the size of the orbit vs rostrum in Caiuajara.

Figure 2. from Manzig et al. 2014. Note the lack of change in the size of the orbit vs rostrum in Caiuajara.

Bone beds are great for individual bone size comparisons, but difficult for creating reconstructions as small individuals are mixed in with large ones. From Manzig et al. (2014) “The presence of three main levels of accumulation in a section of less than one meter suggests that this region was home to pterosaur populations for an extended period of time. The causes of death remain unknown, although similarities with dinosaur drought-related mortality are striking. However, it is also possible that desert storms could have been responsible for the occasional demise of these pterosaurs.”

Figure 2. Typical portion of bone bed of Caiuajara.

Figure 3. Typical portion of bone bed of Caiuajara.

The size of the crests, both below and above the jaws, became larger with age. Most of the individuals were young with only a few adults present.

Figure 4. Caiuajara skulls to scale.

Figure 4. Caiuajara skulls to scale.

The authors found no allometry during ontogeny in post-cranial elements, but adults appear to be more robust and the scapula fused to the coracoid in adults only. This confirms what I’ve found in the fossil record in Zhejiangopterus, Pteranodon, Pterodaustro and generally in phylogenetic analysis. Now, after so much evidence, I hope the naysayers will give the hypothesis of isometry during ontogeny in pterosaurs its day in court.

Figure 5. Caiuajara post crania. Hypothetical hatchlings added at 1/8 adult size.

Figure 5. Caiuajara post crania, a. humerus, b. femur, c. coracoid and scapulocoracoid, d. sternal complex. Hypothetical hatchling elements added at 1/8 adult size. Finally, a fused adult coracoid along with an unfused juvenile and subadult coracoid. Scale bars = 1 cm.

Caiuajara is a small tapejarid, very similar to other tapejarids. This brings up the subject of lumping and splitting with nomenclature, whether a new genus is warranted or not. Is Caiuajara just another species of Tapejara? If not then we need to start splitting up other genera clades containing a wide variety of morphologies as in Rhamphorhynchus, Pteranodon, Germanodactylus, Darwinopterus and other pterosaurs, in which essentially, no two are identical. I’ll leave that to the experts. It’s going to take more than consensus.

Figure 6. Caiuajara size comparisons. There is quite a variety of tapejarids, approaching the variety in Pteranodon, Rhamphorhynchus and other pterosaurs. Note that in the larger Tapejara there is still a suture in the scapulocoracoid.

Figure 6. Caiuajara size comparisons. There is quite a variety of tapejarids, approaching the variety in Pteranodon, Rhamphorhynchus and other pterosaurs. Note that in the larger Tapejara there is still a suture in the scapulocoracoid.

A little speculation
Here we have a large number of juveniles (not hatchlings) and only a few adults in a sandy environment sometimes flooded by rising waters from a nearby lake. What does this mean?

A little backstory:
Pterosaur eggs are large enough that only one could be produced at a time, and held within the mother until just prior to hatching. So the large number of juveniles in this case (no hatchlings here) huddling together, did not belong to a single set of parents. The authors were right, pterosaurs of a certain size (perhaps hatchlings, but up to twice the size of hatchlings in this case) were able to fly. Since they were hatched individually the hatchlings/juveniles sought each other out at an early age, and sought out the company of older, larger tapejarids. Those crests made identification easy. It did not matter that the adults were their parents or not (distinct from the nuclear family illustration at Nat Geo) because the numbers don’t match up. Now IF the adults were found in a distinct layer from the juveniles, the speculation about the adult influence has no basis in evidence.

References
Manzig PC et al. 2014. Discovery of a Rare Pterosaur Bone Bed in a Cretaceous Desert with Insights on Ontogeny and Behavior of Flying Reptiles. Plos ONE 9 (8): e100005. doi:10.1371/journal.pone.0100005.

NOT a new Zhenyuanopterus: XHPM1088

Very, very close, but no cigar.

And not a juvenile either.
A new paper by Teng et al. (2014) reports on a small partial Zhenyuanopterus (XHPM1088, Fig. 1) that does quite fit the morphology of the holotype. No worries. They said it was a juvenile with some odd sorts of allometry going on.

I hate to say it, but we can blame Chris Bennett for this bit of wishful thinking as his 1995 and 1996 papers on Solnhofen pterosaurs opened the doors to letting almost any small specimen become the juvenile of any somewhat similar, but much larger specimen based on the false notion of allometry during ontogeny. Several specimens falsify that little fantasy, including all the embryos now known.

Phylogenetic analysis would have put a stop to such nonsense, but no analysis was undertaken, either in 1995, 1996 or 2014.

Figure 1. XHPM1088 in situ. Only the posterior half is preserved here.

Figure 1. XHPM1088 (mistakenly referred to Zhenyuanopterus) in situ. Only the posterior half is preserved here.

Here’s the problem
The new specimen has a relatively long and robust tail (15 caudals) and a more robust forelimb than hindlimb, plus a Yixian Formation (Early Cretaceous) locality. These facts identified this pterosaur as Zhenyuanopterus to its authors. With identical length ratios between the humerus and femur, Teng et al. thought growth was isometric in these bones, but not others. The scapula has an odd sort of shape otherwise found only in Zhenyuanopterus. However the coracoid was not the same shape or size ratio (Fig. 1). They thought the length of the coracoid would slow dramatically during growth compared to other bones, not realizing that taxa just outside of Zhenyuanopterus (i.e. Boreopoterus, Arthurdactylus, Fig. 2) had a similar long, straight coracoid. They also blamed the coracoid length problem on the holotype of Zhenyuanopterus, saying it was not well-preserved and giving it a longer redicted length based on XHPM1008. That’s not good Science, especially when the coracoids are well preserved and articulated in the holotype.

Unfortunately
Teng et al. thought one of the unique characters of Zhenyuanopterus was its small feet, but the reality is ALL ornithocheirids (more derived than the JZMP embryo) had tiny feet.

Figure 2. The partial pterosaur XHPM1088 to scale with Boreopterus and Zhenyuanopterus and also scaled up to a similar humerus length with Zhenyuanopterus.  Note the coracoids don't match. This is one of the few pterosaurs in which the tibia is shorter than the femur. Boreopterus is similar in this regard.

Figure 2. Click to enlarge. The partial pterosaur XHPM1088 to scale with Boreopterus and Zhenyuanopterus and also scaled up to a similar humerus length with Zhenyuanopterus. Note the coracoids don’t match. This is one of the few pterosaurs in which the tibia is shorter than the femur. Boreopterus is similar in this regard.

A beautiful illustration of Zhenyuanopterus is included in the paper (Fig. 3) sadly flawed by bat-like, deep chord wing membranes and an odd sort of hanging posture for a pterosaur, especially one with such small feet. Some traditions are very hard to kill.

Zhenyuanopterus-illustration

Figure 3. Zhenyuanopterus illustration by Zhao Chuang, a very talented artist. Sadly the wing membranes are wrong and the hanging posture is unlikely based on the tiny feet.

I encourage pterosaur workers
to start putting bones together in reconstructions, then adding new taxa to good phylogenetic analyses before assigning a juvenile status to a small pterosaur that doesn’t match a large one. Here’s a new genus that Teng et al. could have named, but didn’t.

Reference
Teng F-F, Lü J-C, Wei X-F, Hsiao Y-F and Pittman, M 2014. New Material of Zhenyuanopterus (Pterosauria) from the Early Cretaceous Yixian Formation of Western Liaoning. Acta Geologica Sinica (English) 88(1):1-5.