The larger specimen of Sinopterus atavismus enters the LPT basal to dsungaripterids

Many pterosaur fossils attributed to Sinopterus
have been described. They vary greatly in size and shape.

Presently four Sinopterus specimens have been added
to the large pterosaur tree (LPT, 253 taxa). They are all sister taxa, but as in Archaeopteryx, no two are alike, one is basal to the others, which are, in turn, basal to large clades within the Tapejaridae.

1. Sinopterus dongi (the holotype) nests basal to the Tupuxuara clade.
2. Sinopterus liui nests in the Tupuxuara clade.
3. Sinopterus jii (aka Huaxiapterus jii) nests basal to the Tapejara clade.
4. Sinopterus atavisms (Figs. 1-4; Zhang et al. 2019; IVPP V 23388) nests basal to the Dsungaripterus (Fig. 4) clade, outside the Tapejaridae.

Figure 1. Sinopterus atavismus in situ. IVPP V 23388

From the Zhang et al. 2019 abstract:
“Here, we report on a new juvenile specimen of Sinopterus atavismus from the Jiufotang Formation of western Liaoning, China, and revise the diagnosis of this species.”

Zhang et al. note that several elements are unfused including a humeral epiphysis. Several pits and grooves in the distal ends of the long bones are also pitted and grooved. Normally these would be good indicators in archosaurs and mammals, but pterosaurs are lepidosaurs and lepidosaurs follow distinctly different ‘rules’ for growth (Maisano 2002). As an example, some pterosaur embryos have fused elements. Some giant pterosaurs have unfused elements. Here the new specimen (IVPP 23388) is considered an ontogenetic adult as its size is similar to other phylogenetic relatives.

“Sinopterus atavismus does not present a square-like crest. Moreover the feature that groove in the ventral part of the second or third phalanx of manual digit IV is not diagnostic of the species.”

Zhang et al. are comparing the new larger IVPP specimen to the smaller, previously described (Lü et al. 2016) XHPM 1009 specimen (then named Huaxiapterus atavismus), which they considered conspecific. The XHPM specimen has wing phalanx grooves while the IVPP specimen does not. The shapes of the skulls do not match (Fig. 3) and we know that pterosaurs grew isometrically. Thus these two specimens are not conspecific.

“In the new material, the skull preserves a pointed process in the middle part of the dorsal marginof the premaxillary crest, which is different from other Chinese tapejarids. Considering the new specimen is known from a large skeleton that differed from the holotype, this difference may be related to ontogeny, as the premaxillary crest of the holotype is short and does not extend as long as that of the new specimen.”

These two specimens are not conspecific, so ontogenetic comparisons should not be made.

Figure 2. Sinopterus atavismus reconstruction.

From the Zhang et al. 2019 discussion:
“Except for D 2525 which represents an adult individual of Sinopterus (Lü et al. 2006b), all Chinese tapejarid pterosaurs known so far were immature individuals at the time of death. The new specimen (IVPP 23388) shares some features with the holotype of Sinopterus atavismus. The wingspan of the new material is about twice as long as that of the holotype of S. atavismus.”

As mentioned above, the IVPP V 23388 specimen is here considered an adult with unfused bone elements. It needs both a new generic and specific name. The XHPM 1009 specimen (Fig. 3) requires further study.

Figure 3. Sinopterus atavismus size and shape comparison.

The present confusion about the ontogenetic status of pterosaurs 
could have been largely resolved with the publication of “The first juvenile Rhamphorhynchus recovered by phylogenetic analysis” and other papers suppressed by pterosaur referees. Sorry, readers, we’ll have to forge ahead with the venues we have.

Figure 4. Sinopterus atavismus skull restored (gray areas).

Figure 5. Sinopterus atavisms compared to Dsungaripterus to scale.

Sinopterus atavismus (Zhang et al. 2019; Early Cretaceous; IVPP V 23388) was originally considered a juvenile member of the Tapejaridae, but here nests as a small adult basal to Dsungaripteridae. The antorbital fenestra is not taller than the orbit. The carpals are not fused. No notarium is present. The antebrachium is robust. The distant pedal phalanges are longer than the proximal pedal phalanges. An internal egg appears to be present (but half-final-size adults were sexually mature according to Chinsamy et al. 2008,)

Sinopterus dongi IVPP V13363 (Wang and Zhou 2003) wingspan 1.2 m, 17 cm skull length, was linked to Tapejara upon its discovery, but is closer to Tupuxuara.

Sinopterus? liui (Meng 2015; IVPP 14188) is represented by a virtually complete and articulated specimen attributed to Sinopterus, but nests here at the base of Tupuxuara longicristatus.

Sinopterus jii (originally Huaxiapterus jii, Lü and Yuan 2005; GMN-03-11-001; Early Cretaceous) is basal to the Tapejara in the LPT, distinct from the other sinopterids basal to Tupuxuara.

Figure 5. Click to enlarge. The Tapejaridae arise from dsungaripterids and germanodactylids.

The present LPT hypothesis of interrelationships
appears to be a novel due to taxon inclusion, reconstruction and phylogenetic analysis. If not novel, please let me know so I can promote the prior citation.

Traditional phylogenies falsely link azhdarchids with tapejarids
in an invalid clade ‘Azhdarchoidea‘. The LPT has never supported this clade (also see Peters 2007), which is based on one character: an antorbital fenestra taller than the orbit (that a few sinopterids lack). Pterosaur workers have been “Pulling a Larry Martin” by counting on this one character and by excluding pertinent taxa that would have shown them this is a convergent trait ever since the first cladograms appeared in Kellner 2003 and Unwin 2003.

Figure x. Gene studies link swifts to hummingbirds. Trait studies link swifts to owlets. Trait studies link hummingbirds to stilts.

Unrelated update:
The stilt, Himantopus (Fig. x) has moved one node over and now nests closer to the hummingbird, Archilochus. Both arise from the Eocene bird, Eocypselus, which also gives rise to the hovering seagull, Chroicocephalus. The long, mud probing beak of the stilt was adapted to probing flowers in the hummingbird. All these taxa nested close together in the LRT earlier.

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References
Chinsamy A, Codorniú L and Chiappe LM 2008. Developmental growth patterns of the filter-feeder pterosaur, Pterodaustro guinazui. Biology Letters, 4: 282-285.
Kellner AWA 2003. Pterosaur phylogeny and comments on the evolutionary history of the group. Geological Society Special Publications 217: 105-137.
Lü J and Yuan C 2005. New tapejarid pterosaur from Western Liaoning, China. Acta Geologica Sinica. 79 (4): 453–458.
Maisano JA 2002. The potential utility of postnatal skeletal developmental patterns in squamate phylogenetics. Journal of Vertebrate Paleontology 22:82A.
Maisano JA 2002.Terminal fusions of skeletal elements as indicators of maturity in squamates. Journal of Vertebrae Paleontology 22: 268–275.
Peters D 2007. The origin and radiation of the Pterosauria. In D. Hone ed. Flugsaurier. The Wellnhofer pterosaur meeting, 2007, Munich, Germany. p. 27.
Unwin DM 2003. On the phylogeny and evolutionary history of pterosaurs. Pp. 139-190. in Buffetaut, E. & Mazin, J.-M., (eds.) (2003). Evolution and Palaeobiology of Pterosaurs. Geological Society of London, Special Publications 217, London, 1-347.
Wang X and Zhou Z 2003. A new pterosaur (Pterodactyloidea, Tapejaridae) from the Early Cretaceous Jiufotang Formation of western Liaoning, China and its implications for biostratigraphy. Chinese Science Bulletin 48:16-23.
Zhang X, Jiang S, Cheng X and Wang X 2019. New material of Sinopterus (Pterosauria, Tapejaridae) from the Early Cretaceous Jehol Biota of China. Anais da Academia Brasileira de Ciencias 91(2):e20180756. DOI 10.1590/0001-3765201920180756.

wiki/Sinopterus

Was Vellbergia really a juvenile basal lepidosaur? Let’s check…

Earlier we looked at tiny Vellbergia
(Sobral, Simoes and Schoch 2020; Middle Triassic) represented by a disarticulated tiny skull (Fig. 1). The large reptile tree (LRT) nested this hatchling with the much larger adult Prolacerta (Fig. 1). The MPT was 20263 steps for 1654 taxa.

The LRT nesting ran counter to the SuppData cladogram
of Sobral, Simoes and Schoch 2020, who nested Vellbergia among basal lepidosaurs, the closest of which are shown here (Fig. 1). Earlier I did not show the competing lepidosaur candidates. That was an oversight rectified today.

Figure 1. Vellbegia compared to the lepidosaurs it would nest with if Prolacerta and all Archosauromorpha were deleted. Gray areas on Vellbergia indicate restored bone that is lost in the fossil.

To test the lepidosaur hypothesis of relationships,
I deleted all Archosauromorph taxa, including Prolacerta, from the LRT to see where among the Lepidosauromorpha Vellbergia would nest. With no loss of resolution, Vellbergia nested between Palaegama and Tjubina + Huehuecuetzpalli at the base of the Tritosauria plus Fraxinisaura + Lacertulus (Fig. 1) at the base of the Protosquamata. The resulting MPT was 20276 steps, only 13 more than the Prolacerta hypothesis of interrelationships.

That is a remarkably small number considering the great phylogenetic distance between these taxa in the LRT.

Rampant convergence
is readily visible among the competing taxa (Fig. 1). No wonder Prolacerta was named “before Lacerta“, the extant squamate. According to Wikipedia, “Due to its small size and lizard-like appearance, Parrington (1935) subsequently placed Prolacerta between basal younginids and modern lizards. In the 1970s (Gow 1975) the close link between Prolacerta and crown archosaurs was first hypothesized.” That was prior to cladistic software and suffered from massive taxon exclusion.

Allometry vs. Isometry
One of the lepidosaurs shown above, Huehuecuetzpalli (Fig. 1), is known from both an adult and juvenile. The older and younger specimens were originally (Reynoso 1998) considered identical in proportion. Such isometry is an ontogenetic trait shared with other tritosaur lepidosaur clade members, including pterosaurs. On the other hand, if Vellbergia was a hatchling of Prolacerta, some measure of typical archosauromorph allometry should be readily apparent… and it is… including incomplete ossification of the nasals, frontals and parietals along with a relatively larger orbit and shorter rostrum, giving Vellbergia a traditional ‘cute’ appearance appropriate for its clade.

Size
Sobral, Simoes and Schoch considered Vellbergia a juvenile, but it is similar in size to the adult lepidosaurs shown here (Fig. 1). On the other hand, Vellbergia is appropriately smaller than Prolacerta, in line with its hatchling status.

Time
Remember also that Vellbergia is from the Middle Triassic. Prolacerta is from the Early Triassic. They were not found together and some differences are to be expected just from the millions of years separating them.

For comparison: another juvenile Prolacerta,
this time from Early Triassic Antarctica (Spiekman 2018; AMNH 9520), is much larger than Vellbergia from Middle Triassic Germany (Fig. 2), but just as cute. Note the relatively larger orbit and shorter rostrum compared to the adult Prolacerta (Fig. 1), traits likewise found in Vellbergia.

Figure 2. Small Prolacerta specimen AMNH 9520 from Spiekman 2018 compared to scale with Vellbergia. Sclerotic rings (SCL) identified by Spiekman 2018 are re-identified as pterygoids here.

Generally
crushed, disarticulated and incomplete juvenile specimens of allometric taxa are difficult to compare with adults. Even so, what is left of hatchling Vellbergia tends to resemble the larger juvenile and adult specimens of Prolacerta more than hatchling Vellbergia resembles the similarly-sized adult lepidosaurs it nests with in the absence of Prolacerta from the taxon list.

Phylogenetic analysis is an inexact science.
Nevertheless no other known method breaks down and rebuilds thousands of taxa more precisely. Only taxon exclusion appears to trip up workers at present.

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References
Gow CE 1975. The morphology and relationships of Youngina capensis Broom and Prolacerta broomi Parrington. Palaeontologia Africana, 18:89-131.
Parrington FR 1935. On Prolacerta broomi gen. et sp. nov. and the origin of lizards. Annals and Magazine of Natural History 16, 197–205.
Reynoso V-H 1998. Huehuecuetzpalli mixtecus gen. et sp. nov: a basal squamate (Reptilia) from the Early Cretaceous of Tepexi de Rodríguez, Central México. Philosophical Transactions of the Royal Society, London B 353:477-500.
Sobral G, Simoes TR and Schoch RR 2020. A tiny new Middle Triassic stem-lepidosauromorph from Germany: implications fro the early evolution of lepidosauromorphs and the Vellberg fauna. Nature.com Scientific Reports 10, Article number: 2273.
Spiekman SNF 2018. A new specimen of Prolacerta broomi from the lower Fremouw Formation (Early Triassic) of Antarctica, its biogeographical implications and a taxonomic revision. Nature.com/scientificreports (2018)8:17996

wiki/Prolacerta

Pangupterus: a juvenile Moganopterus

Lü et al. 2016
described a new tiny, long-snouthed pterosaur, Pangupterus liui (Jiufotang Fm., Liaoning, Aptian, Early Cretaceous; Figs. 1, 2). Lü et al. thought they had a mandible with a 30º divergence at the jaw symphysis 1/5 of the total jaw length (Fig. 1).

But then, they also report,
“The distal end of the rostrum is slightly expanded, and although it has been destroyed, it seems to have a bony process in the middle, which is similar to the case in Longchengopterus.” The paper has several authors. I don’t think they read each others input.

The color illustration of a restored Pangupterus
that was included with the paper does not follow the first description, but features extremely narrow jaws closer to the second description. The restored body was imaginatively based on a Pterodactylus bauplan.

Figure 1. Pangupterus in situ. Lü et al. had first hand access and considered this a mandible with a symphysis at 1/5 the jaw length. Here, based on this photo,  it is interpreted as a rostrum and mandible, both with parallel rami. If you’re looking at this on a 72 dpi monitor the image is 7/5 larger than life size.

Here, based on tracing
the photo in figure 1, a narrow rostrum lies at an angle to the equally narrow mandible. And the resulting reconstruction matches that of only one pterosaur, Moganopterus, except for its size. The skull of Pangupterus is only 1/4 as long as in Moganopterus (Fig. 2). A hatchling Monganoterpus, if it followed the pattern of other pterosaur hatchlings, would have been 1/8 the size of the adult (Fig. 2) or half the size of Pangupterus.

Figure 2. No other pterosaur has such narrow jaws tipped with slender teeth. Pangupterus is a good candidate to be a juvenile Moganopterus, as shown here.

Moganopterus zhuiana 41HIII0419 (Lü et al. 2012) Early Cretaceous was a large sister to Feilongus and the cycnorhamphids. The skull was extraordinarly stretched out. Feeble needle-like teeth lined the anterior jaws. A long crest that did not break the rostral margin appeared posteriorly. And the neck vertebrae were very much elongated. Likely this was a very tall pterosaur.

Several other blog spots
covered Pangupterus. Some reimagine it as a hummingbird-like specimen. See other images here, here, here and here.

This specimen
further confirms the presence of tiny, long-snouted pterosaurs, some of them juveniles of larger long-snouted pterosaurs, and the isometric ontogenetic growth of all pterosaurs.

References
Lü J-C, Pu H-Y, Xu i, WuY-H and Wei X-F 2012. Largest Toothed Pterosaur Skull from the Early Cretaceous Yixian Formation of Western Liaoning, China, with Comments On the Family Boreopteridae. Acta Geologica Sinica 86 (2): 287-293.
Lü J-C, Liu C, Pan L-J and Shen C-Z 2016. A new pterodactyloid pterosaur from the Early Cretaceous of the Western part of the Liaoning Province, Northeastern China. Acta Geologica Sinica (English) 90(3):777-782.

wiki/Moganopterus
/wiki/Pangupterus

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.

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 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. 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

Allometry during ontogeny in the basal tritosaur, Huehuecuetzpalli

Huehuecuetzpalli (Reynoso 1998) is a basal tritosaur according to the large reptile tree, a lepidosaur nesting outside of the Squamata, and ancestral to Tanystropheus, Macrocnemus, drepanosaurids, fenestrasaurs and ultimately, pterosaurs. The lineage of pterosaurs is shown here.

Huehuecuetzpalli specimens are only known from the Early Cretaceous, with ghost lineage origins going back to the Late Permian. Long species survival is not uncommon among lepidosaurs, as in the extant Sphenodon with relatives in the Triassic.

Figure 1. Two specimens of Huehuecuetzpalli were found, one adult and one juvenile. Here they are shown together to scale along with manus and pes comparisons scale to a common length for metacarpal 4 and metatarsal 4.

Reynoso 1998 reported,
“The relative length of the snout, and the proportions of the skull and limbs relative to the presacral vertebral column, do not show signifcant differences between the juvenile and adult specimens, although these features usually change in ontogeny. This suggests that adult proportions were already acquired at the ontogenetic stage of the younger specimen in spite of its relatively smaller size.”

I have been repeating this observation
with regard to pterosaurs, which likewise do not show any significant differences (apart from the enlargement of any skull crests) in their morphological proportions. For examples click here, here and here and other references therein.

But is it true for Huehuecuetzpalli?
That’s why side-by-side comparisons are so useful. Sadly, I have not done so until just yesterday (Figs. 1, 2).

Figure 2. Huehuecuetzpalli, adult and juvenile skulls to scale. note the relatively shorter rostrum in the juvenile, which also had smaller teeth and a shorter set of parietals (with a smaller braincase and smaller jaw adductor chamber). In the juvenile the ascending process of the premaxilla was more robust and the tooth-bearing portion was shorter with fewer teeth.

Reynoso 1998 reported,
“The complete fusion of the cranial elements suggests that the larger specimen is of post-juvenile age, and probably an adult condition was already acquired. The olecranon process of the ulna, however, is not completely ossified and attached to the ulna, and only a ball of hard tissue (calcified cartilage or bone) is preserved. It was impossible to find information in the literature about the time when the precursor of the olecranon process become fused to the ulna.

“The age of the smaller specimen is more difficult to establish. The complete ossification of the fourth distal tarsal and the still separated astragalus and calcaneum undoubtedly suggest a post-hatchling stage. The complete fusion of the frontal, however, shows that it is older than Rieppel’s specimen number 18 and the hatchling of Cyrtodactylus pubisulcus (Gekkonidae) illustrated by Rieppel (1992a: ¢g. 1). The high degree of ossification indicates that it is close to the latest stages of development preceding complete ossification. Juvenile skull characters are the presence of a broader parietal table with short lateral processes. Compared with the adult skull, the juvenile parietal table is more than 15% broader on the narrower section excluding the ventrolateral flanges for the dorsal attachment of the jaw adductor musculature.”

We looked at olecranon ossification in tritosaurs earlier here.

As a rule, lepidosaurs don’t change much during ontogeny
as we’ve seen earlier here with Shinisaurus. But they do change… a little.

References
Reynoso V-H 1998. Huehuecuetzpalli mixtecus gen. et sp. nov: a basal squamate (Reptilia) from the Early Cretaceous of Tepexi de Rodríguez, Central México. Philosophical Transactions of the Royal Society, London B 353:477-500.