Caupedactylus ybaka (Kellner 2012) enters the LPT

Kellner (2012, 2013) described
the skull of an Early Cretaceous sinopterid pterosaur, Caupedactylus ybaka (MN 4726-V, Fig. 1). The skull is about forty-six centimetres long. (Hope I got this right this time).

Earlier the same skull was posted online.

Figure x. Caupedactylus in situ and restored by sculptors.

Figure x. Caupedactylus in situ and restored by sculptors. Or a different specimen.

New Tapejarid-Tupuxuarid skull.

Figure 1. New Tapejarid-Tupuxuarid skull now named Caupedactylus.

Bones colorized in this tapejarid / tupuxuarid.

Figure 2. Bones colorized in this tapejarid / tupuxuarid, named Caupedactylus.

Abstract
“A new unusual tapejarid pterosaur from the Early Cretaceous Romualdo Formation (Araripe Basin, Brazil) is described, based on a skull, lower jaw and some postcranial elements. Caupedactylus ybaka gen. et sp. shows the typical high nasoantorbital fenestra of the Thalassodrominae but lacks a palatal ridge, and shares with the Tapejarinae several features, including a downturned rostral end, allowing its allocation to that clade.”

The new skull compared to other tapejarids. Click to enlarge.

Figure 2. Click to enlarge. The rising size of the tapejaridae.

Abstract continues
“Furthermore, the new species differs in having an anteriorly and posteriorly expanded premaxillary sagittal crest, the lacrimal process of the jugal strongly inclined, and a slit-like postpalatine fenestra, among other characters. The region of the left jugal-quadratojugal-quadrate shows a pathology that is likely the result of an infection. The lateral surface of the premaxillary crest presents grooves that were interpreted in other pterosaurs as impressions of blood vessels, corroborating growing evidence that cranial crests could have been involved in thermoregulation.”

“Also, the new species has a well-preserved palate with a large palatine forming the anterior region of the choanae and the postpalatine fenestra and a secondary subtemporal fenestra. Since the latter has been regarded as unique to non-pterodactyloids, its occurrence in Caupedactylus demonstrates that the evolution of palatal region in pterosaurs is more complex than previously thought.”

Perhaps to no one’s surprise, this specimen nested in 2013 in the large pterosaur tree (LPT) between Sinopterus dongi and Tupandactylus.


References
Campos HBN and Headden JA 2013. A review of Tupuxuara deliradamus (Pterosauria, Azhdarchoidea, Thalassodromidae) from the Early Cretaceous Romualdo Formation of Brazil. International Symposium on Pterosaurs – Rio Ptero 2013.
Elgin RA 2015. Paleobiology, Morphology and Flight Characteristics of Pterodactyloid Pterosaurs. Dissertation, University of Heidelberg.
Kellner AWA 2012. A new unusual tapejarid (Pterosauria, Pterodactyloidea) from the Early Cretaceous Romualdo Formation, Araripe Basin, Brazil. Cambridge University Press 103 (3-4) The Full Profession: A Celebration of the Life and Career of Wann Langston, Jr., Quintessential Vertebrate Palaeontologist September 2012 , pp. 409-421.
Kellner AWA 2013. A new unusual tapejarid (Pterosauria, Pterodactyloidea) from the Early Cretaceous Romualdo Formation, Araripe Basin, Brazil. Earth and Environmental Science Transactions of the Royal Society of Edinburgh 103(3-4): 409-421.
Manzig PC et al. (10 co-authors) 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.
Martill DM and Naish D 2006. Cranial Crests Development in the Azhdarchoid Pterosaur Tupuxuara, With Review of the Genus and Tapejarid Monophyly. Palaeontology 49(4): 925-941.

wiki/Caupedactylus
pterosaurheresies.wordpress.com/2013/06/06/tapejarid-or-tupuxuarid/

A quick look at the original Tapejara skull

Short one today
told mostly in pictures.

Topic: Simplification. 
Get rid of the extraneous data to better see and understand the basics.

Figure 1. Just take out everything that isn't this side of Tapejara. Rearrange the parts for best fit. Make a guess for the missing parts and this is what you get.

Figure 1. Just take out everything that isn’t this side of Tapejara. Rearrange the parts for best fit. Make a guess for the missing parts and this is what you get.

In this case
the skull of the original Tapejara fossil (Fig. 1) is trimmed back to just the basics and slightly shifted to fit. Missing tips are added based on phylogenetic bracketing.

Figure 4. Click to enlarge. The Tapejaridae arise from dsungaripterids and germanodactylids.

Figure 2. Click to enlarge. Tiny Tapejaridae arise from dsungaripterids and germanodactylids, then grow larger phylogenetically.

Tapejara wellnhoferi
(Kellner 1989; 108 mya, Early Cretaceous) was immediately recognized as something quite different when first discovered. Compared to other specimens, this one appears to have no sharp premaxilla, likely due to taphonomic loss.


References
Eck K, Elgin RA and Frey E 2011. On the osteology of Tapejara wellnhoferi KELLNER 1989 and the first occurrence of a multiple specimen assemblage from the Santana Formation, Araripe Basin, NE-Brazil. Swiss Journal of Palaeontology, doi:10.1007/s13358-011-0024-5.
Kellner AWA 1989. A new edentate pterosaur of the Lower Cretaceous from the Araripe Basin, northeast Brazil. Anais da Academia Brasileira de Ciências 61, 439-446.

wiki/Tapejara

 

Huaxiapterus (Sinopterus) benxiensis enters the LPT basal to Tapejaridae

Lü et al. 2007
introduced us to the nearly complete crushed skeleton of Huaxiapterus benxiensis (Figs. 1, 2, BXGM V 0011) in a short paper with a short, three-sentence abstract.

Figure 1. Huaxiapterus benxiensis (BXGM V 0011) in situ, largely complete, crushed and articulated.

Figure 1. Huaxiapterus benxiensis (BXGM V 0011) in situ, largely complete, crushed and articulated.

From the Lü et al. abstract
“A new species of Huaxiapterus: H. benxiensis sp. nov. is erected based on the new specimen. The diagnostic characters of Huaxiapterus benxiensis are well-developed premaxillary crest and parietal spine, the crest and spine parallel and extending posterodorsally, and a shallow groove present on the dorsal surface of the anterior portion of the mandibular symphysis. The different skull morphologies of Chinese tapejarid pterosaurs indicate that they are much more diverse than the previous thought.”

As in all known specimens of pterosaurs, no two adults are alike. That fact gives us an excellent view of microevolution at work in this and other pterosaur clades.

Unfortunately, other workers refuse to add pertinent largely complete taxa shown in the large pterosaur tree (LPT, 256 taxa), nor have they added valid outgroups with correct scores. So the taxonomy and nomenclature in those smaller studies tends to get confused.

Figure 2. DGS reconstruction from the low resolution in situ image in figure 1. Note the brevity of the distal wing phalanges, the robust hind limbs and the gracile humerus.

Figure 2. DGS reconstruction from the low resolution in situ image in figure 1. Note the brevity of the distal wing phalanges, the robust hind limbs and the gracile humerus.

Huaxiapterus benxiensis (Lü et al. 2007, Early Cretaceous, BXGM V 0011, aka Sinopterus benxiensis) nests as the last common ancestor of the Tapejara clade + the Tupuxuara clade. Recently here, here and here we looked at other specimens assigned to Huaxiapterus that were later switched over to Sinopterus. This one (the BXGM specimen, Figs. 1, 2) is among those. They do nest closer to Sinopterus dongi (the holotype) than to Huaxiapterus jii, (the holotype), but H jii was also switched over to Sinopterus.

Almost flightless?
Different from other related pterosaurs Huaxiapterus benxiensis had shorter distal wing phalanges (m4.2, 4.3 and 4.4), a slender humerus and robust hind limbs. Together these traits suggest a trend to a reduced flight ability. Other, more clearly flightless pterosaurs are documented here, here, here and here.

Figure 4. Tapejaridae in the LPT.

Figure 4. Tapejaridae in the LPT. The BXGM specimen shows in the enlargement. It was about the size of Sinopterus dongi at the genesis of the Tupuxuara clade. 

A poor flyer at the base of the flying Tapejaridae
is possible, given that H. benxiensis is probably not the real last common ancestor, but more likely close to the real last common ancestor. It likely evolved its own way. The small size of H. benxiensis is in keeping with phylogenetic miniaturization at the start of other pterosaur clades and major clades in general.


References
Lü JC, GAo YB, Xing LD, Li ZX and Ji Q 2007. A New Species of Huaxiapterus (Pterosauria: Tapejaridae) from the Early Cretaceous of Western Liaoning, China. Acta Geol Sinica – English 81: 683-687.

Sinopterus? or Huaxiapterus? It gets confusing…

A kind reader alerted me to a misidentification here.
The grayscale image (Fig. x) is the ZMNH M 8131 specimen of Huaxiapterus. When a higher resolution image becomes available I will return to this specimen and edit the copy. With that in mind… here is the original blogpost, awaiting an edit.

Figure x. Huaxiapterus ZMNH-M-8131 specimen.

Figure x. Huaxiapterus ZMNH-M-8131 specimen.

 

Thank goodness
for museum numbers.

Today the ZMNH M 8131 specimen first attributed to
Huaxiapterus corollatus (Lü et al. 2006) then renamed Sinopterus corollatus (Zhang et al. 2019; Figs. 1, 2) enters the the large pterosaur tree (LPT, 255 taxa) basal to tapejarids, derived from the Sinopterus atavismus specimen nesting basal to dsungaripterids.

Figure 1. Huaxiapterus corollatus ZMNH M 8131 reconstructed. An alternate m4.1 is provided that looks more like a m4.1 than a metacarpal 4.

Figure 1. Huaxiapterus corollatus ZMNH M 8131 reconstructed. An alternate m4.1 is provided that looks more like a m4.1 than a metacarpal 4.

Sometimes specimens are reassembled slightly wrong.
In this case several long bones were accidentally reversed end-to-end in this otherwise stunning mount. One never knows what the original fossil looked like prior to reassembly. We don’t want to call these ‘fakes’. We do want to be aware of errors and artistic reconstructions as much as is possible.

Figure 2. The ZMNH specimen in situ and somewhat corrected for original perspective issues. The correction makes the wings the same length.

Figure 2. The ZMNH specimen in situ and somewhat corrected for original perspective issues. The correction makes the wings the same length. Be wary of such wonderful-looking fossils. This specimen appears to have been reassembled. Some long bones are reversed end-to-end, which do not affect scoring.

Sinopterus – Huaxiapterus corollatus (Lü et al. 2006; Early Cretaceous, ZMNH M 8131) is another largely complete specimen with confusing nomenclature. This taxon nests at the base of the Tapejara, basal to the Aathal specimen (below). The pelvis is missing. The sternum is among the largest of all pterosaurs. The cervicals are longer creating a taller pterosaur.

From the Lü et al. abstract:
“A new species of tapejarid pterosaur, Huaxiapterus corollatus sp. nov. is erected on the basis of a nearly complete skull and postcranial skeleton from the Lower Cretaceous Jiufotang Formation of Liaoning Province, China. Huaxiapterus corollatus sp. nov. is characterized by a hatchet-shaped rectangular process on the premaxilla, whose short axis is perpendicular to the anterior margin of the premaxillae. Except for this process, other characters of the skull such as the breadth of the snout between the anterior margin of the nasoantorbital fenestra and the anterior margin of the premaxilla are similar to that of Huaxiapterus jii.”

Figure 3. Huaxiaptrus iii and Huaxiapterus corollatus to scale. These two do not nest next to one another in the LPT.

Figure 3. Huaxiaptrus iii and Huaxiapterus corollatus to scale. These two do not nest next to one another in the LPT.

The Lü et al. abstract continues
“Huaxiapterus and a second Chinese tapejarid, Sinopterus, share several unique cranial characters in common with Tapejara and these three genera appear to be more closely related to each other than to other azhdarchoids.

In the LPT azhdarchids nest with dorygnathids, not tapejarids. Adding these taxa missing from prior studies makes this inevitable.

“The Chinese tapejarids (Sinopterus and Huaxiapterus) have relatively elongate skulls and weakly developed cranial crests and seem to be less derived than Tapejara, with its shorter, deeper skull and large cranial crest. Tupuxuarids (Tupuxuara and Thalassodromeus) have often been associated with tapejarids in the family Tapejaridae, but this relationship is controversial because some phylogenetic analyses have supported the pairing of tupuxuarids with Azhdarchidae.”

Adding taxa moves Azhdarchidae away from tupuxuarids.

Figure 4. Tapejaridae in the LPT.

“We propose that Tapejaridae be restricted to Tapejara, Sinopterus and Huaxiapterus.”

The LPT does not support that proposal (Fig. 4). The Tapejaridae remains a monophyletic clade in the LPT derived from dsungaripterids, shenzhoupterids and earlier, germanodactylids… not azhdarchids.


References
Lü JC, Jin XS, Unwin DM, Zhao LJ, Azuma Y and Ji Q 2006. A new species of Huaxiapterus Pterosauria: Pterodactyloidea from the Lower Cretaceous of western
Liaoning, China with comments on the system atics of tapejarid pterosaurs. Acta Geol Sinica English 80: 315-326.
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
wiki/Huaxiapterus
reptileevolution.com/tapejaridae.htm

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.

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.

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 comparison

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 3. Sinopterus atavismus skull restored (gray areas).

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

Figure 4. Sinopterus atavisms compared to Dsungaripterus to scale.

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.

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 1. Gene studies link swifts to hummingbirds. Trait studies link swifts to owlets. Trait studies link hummingbirds to stilts.

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.


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

The Aathal specimen enters the LPT at the base of the Tapejaridae

Updated November 18, 2020
with the realization that this is not Sinopterus jii, as previously noted. I believed an online image that labeled it such and did not reference earlier comments. Apologies for the confusion.

Earlier I found this image of a sinopterid with broken wings
as I sought the specimen number in order to nest it in the large pterosaur tree (LPT, 253+ taxa). A kind reader (TG) replied it was from the Sauriermuseusm in Aathal, Germany.  Today it enters the LPT.

Figure 1. Sinopterus jii in situ. Note the broken arms.

Figure 1. Sinopterus jii in situ. Note the broken arms.

Reconstructions of the body and skull
enabled scoring of the Aathal specimen as it nested basal to tapejarids. Two Sinopterus specimens nested basal to the tupuxuarids. Outgroups include other China pterosaurs in the Dsungaripteridae and Shenzoupteridae.

Figure 1. The Sauriermuseum Aathal specimen. Scale remains unknown.

Figure 1. The Sauriermuseum Aathal specimen. Scale remains unknown.

Figure 3. Sauriermuseum Aathal skull in situ and reconstructed.

Figure 3. Sauriermuseum Aathal skull in situ and reconstructed.

The frontal + parietal crest on the Aathal specimen
is distinctive and points to a wider variation than typically imagined for this genus.

Phylogenetic Note
The antorbital fenestra is not taller than the orbit in the Aathal specimen (Fig. 3). Traditional phylogenies 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. 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 4. Click to enlarge. The Tapejaridae arise from dsungaripterids and germanodactylids.

Figure 4. Click to enlarge. The Tapejaridae arise from dsungaripterids and germanodactylids.

Small pterosaurs are usually small adult pterosaurs, 
not juveniles, contra traditional thinking. There are a few notable exceptions herehere and here.


References
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.
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.

wiki/Sinopterus

New basal tapejarid with broken wings needs specimen number, citation

Updated April 1, 2020
The specimen number is SMA 0154 / 02. Kind readers reported the location of this specimen: Sauriermuseum, Aathal, Switzerland. I can now reveal the phylogenetic nesting of this specimen is between Sinopterus and Tapejara. I know of no citation yet.

Figure 1. Complete basal tapejarid without identification. Please provide a museum number or citation if possible.

Figure 1. Complete basal tapejarid without identification. Please provide a museum number or citation if possible.

This image above (Fig. 1) appears on the website,
Tapejaraluv.weebly.com” under the headline “The Tapejara,” created by Jordyn Rosen and Teya Good. There are no ‘contact us‘ or ‘comments‘ links on their website and all attempts at finding them elsewhere on the ‘net don’t seem to be leading to any Tapejara fans. 

I will forego posting any more information on this specimen
pending the acquisition of a citation or museum number on the chance that it is currently under study and awaiting publication. Even so, it has been added to the large pterosaur tree (LPT) as the 243rd taxon, but not yet posted online.

Sinopterus IVPP V 23388 in detail

Zhang et al. 2019 bring us
a beautiful, virtually complete pterosaur specimen from the Early Cretaceous of China: Sinopterus atavismus (Fig. 1). Despite a thorough description, some small parts here and there were overlooked.

Figure 1. Sinopterus atavismus, matrix removed. Snout tip, fingers and toes enlarged to show detail.

Figure 1. Sinopterus atavismus, matrix removed. Snout tip, fingers and toes enlarged to show detail. The lack of fusion in the scapula + coracoid has nothing to do with ontogeny, just phylogeny, as in other lepidosaurs

‘Toothless’ pterosaurs retain one tooth
at the tip of their premaxilla and dentary (Fig. 1). We looked at that trait earlier here in 2011. Mainstream paleontologists still don’t look for it. The evolution of these procumbent teeth are documented in tiny Solnhofen ancestors.

Manual digit 5 is not lost,
but it remains as a tiny vestige in pterosaurs. In this case manual digit 5 was so prominent Zhang et al. traced it without labeling it. Here it is (Fig. 1) in situ and unfolded. We looked at that trait earlier here in 2011. Mainstream paleontologists still don’t look for it.

Pedal digit 5 likes to tuck itself behind the ankle
in situ as it does here (Fig. 1). It was not hard to find, if you don’t set out to ignore it. We looked at that trait earlier here in 2012 (Peters 2011). Mainstream paleontologists still don’t look for it.

Extended tail vertebrae are easy to overlook,
but if you look for them, you will find them on this pterodactyloid-grade tapejarid. We looked at that trait earlier here in 2013. Mainstream paleontologists still don’t look for it.

Figure 2. The new IVPP Sinopterus specimen in situ with original interpretation and DGS colors used to find the 'missing' pteroids. The cervicals are numbered anew. The sternal complex (not just the sterum) borders are identified or reidentified.

Figure 2. GIF animation of the new IVPP Sinopterus specimen in situ with original interpretation and DGS colors used to find the ‘missing’ pteroids. The cervicals are numbered anew. The sternal complex (not just the sterum) borders are identified or reidentified.

Where are the missing pteroids?
Zhang et al. report the pteroids and all sesamoids are missing from this specimen. Digital Graphic Segregation (DGS) is helpful by eliminating all the other bones by color until the pteroids appear by default (Fig. 2).

Figure 3. Sinopterus skulls presented by Zhang et al. 2019.

Figure 3. Sinopterus skulls presented by Zhang et al. 2019.

Juvenile?
The new IVPP V 23388 specimen is considered a juvenile, despite being one of the largest Sinopterus specimens (Fig. 3). The lack of fusion in the scapula + coracoid has nothing to do with ontogeny, just phylogeny, as in other lepidosaurs, as we learned earlier here.


References

Peters D 2011. A Catalog of Pterosaur Pedes for Trackmaker Identification Ichnos 18(2):114-141.
Zhang X-J, Jiang S-X, Cheng X and Wang X-L 2019.
New Material of Sinopterus (Pterosauria, Tapejaridae) from the Early Cretaceous Jehol Biota of China. Anais da Academia Brasileira de Ciências (2019) 91(Suppl. 2): e20180756 (Annals of the Brazilian Academy of Sciences). Printed version ISSN 0001-3765 / Online version ISSN 1678-2690
http://dx.doi.org/10.1590/0001-376520192018756

SVP 7 – Montanazhdarcho, not an azhdarchid

Carroll 2015 reports
on the North American Late Cretaceous pterosaur, Montanazhdarcho.

From the abstract:
“The latest Cretaceous fossil record of pterosaurs is dominated by azhdarchids, despite various reports of non azhdarchid material from the Late Campanian and Maastrichtian. However, the only indisputable non-azhdarchid pterosaur material from the latest Cretaceous has been the single nyctosaurid humerus from the Gramame Formation of Brazil*. This study presents evidence that Montanazhdarcho minor is a nonazhdarchid member of the Azhdarchoidea**. M. minor was assigned to the Azhdarchidae based on diagnostic features of the humerus, shoulder girdle, and the partial cervical vertebra of the holotype MOR 691. The initial description focused mainly on the diminutive size (2.5 m wingspan). Subsequent discoveries of postcranial material from thallasadromines, tapejarines, and azhdarchids have revealed that the postcranial features initially used to assign M. minor to the Azhdarchidae are synapomorphies for the more inclusive Azhdarchoidea clade. Phylogenetic analysis reveals that Montanazhdarcho possesses multiple characters that are shared by the Tapejarinae and Thallasadrominae: (1) a broad and well-developed tubercle at the ventroposterior margin of the coracoid; (2) a massive, distinct ulnar crest with a developed proximal ridge; (3) a strong boot-like ventral margin of the humeral head; (4) an ulna/radius as long or longer than metacarpal IV; an (5) a phalanx IV-1 that is as long or longer than metacarpal IV. The results of this study show that the Late Cretaceous pterosaur fauna was not entirely dominated by azhdarchids and recognizes important post-cranial characters that better define Azhdarchoidea. The reappraisal of M. minor as a non-azhdarchid member of theAzhdarchoidea also recognizes M. minor as the first known pterosaur of that clade found in North America, as well as one of the latest occurrences of the group.”

*Actually there is a large rostrum that belongs to another tupuxuarid from the Latest Cretaceous of southern Texas (Fig. 1), so it is not surprising to find tapejarid material then and there.

TMM 42489-2, the tall crested Latest Cretaceous large rostrum and mandible. It's a close match to that of Tupuxuara, otherwise known only from Early Cretaceous South American strata.

Figure 1. TMM 42489-2, the tall crested Latest Cretaceous large rostrum and mandible. It’s a close match to that of Tupuxuara, otherwise known only from Early Cretaceous South American strata.

** Azhdarchoidea improperly includes tapejarids. In the large pterosaur tree the azhdarchids are derived from dorygnathids while the tapejarids are derived from germanodactylids.

References
Carroll N 2015.
Reassignment of Montanazhdarcho minor as a nonazhdarchid member of the Azhdarchoidea. Journal of Vertebrate Paleontology abstracts.
Padian, K., Horner, J.R., and de Ricqlès, A.J 1993. A new azhdarchid pterosaur from the Two Medicine Formation (Late Cretaceous, Campanian) of Montana, identified on the basis of bone histology. Journal of Vertebrate Paleontology 13: 52A.
Padian K, de Ricqlès AJ and Horner JR 1995. Bone histology determines identification of a new fossil taxon of pterosaur (Reptilia: Archosauria)”, Comptes Rendus de l’Academie des Science Serie II (320): 77-84.
McGowen MR, Padian K, de Sosa MA and Harmon RJ 2002. Description ofMontanazhdarcho minor an azhdarchid pterosaur from the Two Medicine Formation (Campanian) of Montana. PaleoBios 22(1): 1–9.

Introducing Sinopterus(?) liui

Figure 1. Sinopterus liui reconstructed from the original tracing of Meng Xi thesis for Masters degree.

Figure 1. Sinopterus liui reconstructed from the original tracing of Meng Xi thesis for Masters degree. Note the long cervicals not found in Sinopterus.

A new, virtually complete and articulated pterosaur specimen (IVPP V14188, Fig. 1), Sinopterus liii (Meng 2015; Jiufotang Formation, 120 mya), is considered a new species of a well known genus, Sinopterus. According to Meng, distinct traits include:

  1. large body size
  2. short beak
  3. high and short skull
  4. cranially sloping crest on premaxilla
  5. well developed hatchet-shaped deltopectoral crest of humerus

Figure 2. Tapejaridae. Sinopterus liui is indeed larger than other Sinopterus specimens.

Figure 2. Tapejaridae. Sinopterus liui is indeed larger than other Sinopterus specimens. Click to enlarge. This clade evolves to greater size and greater crest size.

Only the abstract is in English. Meng also modifies the description of the genus Sinopterus (Fig. 3).

Figure 1. Sinopterus and purported juvenile, but note the skull is relatively smaller with smaller eyes in the smaller specimen. The feet are also distinct. This appears to be a smaller adult of another species, not a juvenile.

Figure 3. Sinopterus and purported juvenile, but note the skull is relatively smaller with smaller eyes in the smaller specimen. The feet are also distinct. This appears to be a smaller adult of another species, not a juvenile.

From the Meng abstract
“Studies on the cranial crest have resulted in the suggestion that the bony crest was ossified by a crest composed of soft tissue during the ontogeny. As a result, the variation of size and shape among various Sinopterus specimens may be related to ontogeny and sexual behavior. Consequently, the sole difference in the cranial crest morphology between Huaxiapterus and Sinopterus is not taxonomically diagnostic, thus we propose that Huaxiapterus and Sinopterus are synonymy. Frequent volcanic activity in this region has caused the mass mortality of vertebrates.”

So Meng is a lumper.
Unfortunately we have not been able to verify variation in size and shape (except for enlargement of a cranial crest) in any pterosaur during ontogeny — and none, so far, have been able to assigned to a gender unless an aborted egg is nearby. These observations, of course, are contra traditional pterosaur paradigms, but they do follow the data.

Figure 4. Sinopterus iui skull in situ and reconstructed using DGS. That small orbit is distinct from Sinopterus and similar to Tupuxuara.

Figure 4. Sinopterus iui skull in situ and reconstructed using DGS. That small orbit is distinct from Sinopterus and similar to Tupuxuara.

I would have given this specimen a new genus. 
Based on the low resolution images provided and the great size difference, the new specimen (Fig. 4) appears to be sufficiently distinct to warrant its own genus — but then again, all those Pteranodon specimens are lumped into one genus.

Will the lumpers and splitters please get together?

TMM 42489-2, the tall crested Latest Cretaceous large rostrum and mandible. It's a close match to that of Tupuxuara, otherwise known only from Early Cretaceous South American strata.

Figure 4. TMM 42489-2 and the Tupuxuara longicristatus, to tupuxuarids with skulls similar, but not too similar, to the new Sinopterus.

 

References
Meng X 2015. A New Species of Sinopterus from Jehol Biota and Reconstraction of Stratigraphic Sequence of the Jiufotang Formation. A Thesis for the Master Degree of Science in the Graduate School of Chinese Academy of Sciences, directed by Wang X-L. (in Chinese)