A new cat-sized flightless azhdarchoid from Canada

Martin-Silverstone, Witton, Arbour and Currie 2019 bring us
news of a humerus and several vertebrae (some fused into a notarium) they taxonomically narrow down to a cat-sized Campanian (Late Cretaceous) azhdarchoid pterosaur. It was found on Hornsby Island, close to the much larger Victoria Island in Southwestern Canada.

The authors report,
“the individual was approaching maturity at time of death.”

In their discussion, the authors state:
“The thin bone walls, gracile bone construction and humeral morphology of RBCM.EH.2009.019.0001, indicate it clearly belonged to a volant Mesozoic animal, a pterosaur or avialan.”

There’s not much to see or reconstruct here.
The few bones found are fragments still in the matrix. The link below will take you to the online PDF.

On closer inspection,
the small triangular deltopectoral crest is smaller than in flightless pterosaurs (e.g. Sos2428 in Fig. 1) and the bone is thicker than one might expect of a volant pterosaur. The authors do not consider the possibility that their specimen had a volant ancestry, but was itself no longer volant, as happens often enough in the azhdarchid line of wading pterosaurs. Some were tiny (Fig. 1), some cat-sized, some man-sized and others much larger. Some were volant. Others were not, convergent with birds of all sizes.

Figure 2. The flightless pterosaur, Sos 2428, along with two ancestral taxa, both fully volant. Note the reduction of the wing AND the expansion of the torso. We don't know the torso of Q. northropi. It could be small or it could be very large.

Figure 1. The flightless pre-azhdarchid pterosaur, Sos 2428, along with two ancestral taxa, both fully volant. Note the reduction of the wing AND the expansion of the torso. This deltopectoral crest is at least twice the size of the new Canadian specimen

Dr. Witton has invested much time and treasure
in telling us giant azhdarchids were volant, despite the facts that weigh against that hypothesis. He also omits the data on three flightless pterosaurs, including Sos2428 (Fig. 1). Now we can add a fourth, his new cat-sized Hornby Island pterosaur.

Earlier co-author Witton
and Habib 2010 discussed hypothetical flightlessness in giant azhdarchids from many angles, but never introduced actual flightless taxa, two of which were known at the time. This online paper included infamous illustrations of an ornithocheirid manus in the process of a quadrupedal launch that had been cheated to implant the wing finger on the substrate, something that never happens according to the ichnite record. They did this by shrinking the free fingers.

Quetzalcoatlus running like a lizard prior to takeoff.

Figure 2. Quetzalcoatlus running like a bipedal lizard with no need or ability to fly.

Postscript
Interesting blog post here on an unfortunate bone misidentification on a paper earlier by one of the co-authors. Thank goodness Witton chose not to vilify his co-author.


References
Martin-Silverstone E, Witton MP, Arbour VM and Currie PJ 2019. A small azhdarchoid pterosaur from the latest Cretaceous, the age of flying giants. Royal Society open science 3: 160333. http://dx.doi.org/10.1098/rsos.160333
Witton MP, Habib MB 2010. On the Size and Flight Diversity of Giant Pterosaurs, the Use of Birds as Pterosaur Analogues and Comments on Pterosaur Flightlessness. PLoS ONE 5(11): e13982. https://doi.org/10.1371/journal.pone.0013982

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Vestigial fingers on the UNSM 93000 Nyctosaurus

The UNSM 93000 specimen attributed to Nyctosaurus
has only three wing phalanges and the tiny vestigial free fingers have never been looked at using DGS methods before. Well, here they are (Fig. 1).

Figure 1. Closeup of the UNSM 93000 specimen of Nyctosaurus focusing on three vestige free fingers.

Figure 1. Closeup of the UNSM 93000 specimen of Nyctosaurus focusing on three vestige free fingers. This is what happens when you no longer need these fingers. You can tell Nyctosaurus from Pteranodon in that the former never fuses the sesamoid (extensor tendon process) to phalanx 4.1. Other wrongly consider this a trait of immaturity.

Nyctosaurus sp. UNSM 93000 (Brown 1978, 1986) was derived from a sister to Nyctosaurus gracilis and phylogenetically preceded the crested Nyctosaurus specimens. Except for the rostral tip, the skull and cervicals are missing. Distinct from Nyctosaurus gracilis, the dorsals of the Nebraska specimen relatively shorter. The scapula and coracoid were more robust. The deltopectoral crest of the humerus most closely resembled that of Muzquizopteryx. Fingers I-III were tiny vestiges. Manual 4.1 extended to mid ulna when folded. Manual 4.4 was probably fused to m4.3 or it was missing and m4.3 became curved.

Figure 1. The UNSM specimen of Nyctosaurus, the only one for which we are sure it had only three wing phalanges.

Figure 2. The UNSM specimen of Nyctosaurus, the only one for which we are sure it had only three wing phalanges.

The pubis and ischium did not touch, as in more primitive nyctosaurs. It would have been impossible for the forelimb to develop thrust during terrestrial locomotion. It was likely elevated or used like a ski-pole.


The family tree of the Ornithocephalia and Germanodactylia is here. The expanded family tree of the Pterosauria is here.


References
Brown GW 1978. Preliminary report on an articulated specimen of Pteranodon Nyctosaurusgracilis. Proceedings of the Nebraska Academy of Science 88: 39.
Brown GW 1986. Reassessment of Nyctosaurus: new wings for an old pterosaur. Proceedings of the Nebraska Academy of Science 96: 47.

 

Another disc-head anurognathid from Jurassic China

Yesterday Yang et al. 2018 presented NJU-57003 (Figs. 1–3), a small anurognathid pterosaur with a great deal of soft tissue preservation, including feather-like filaments, said to be homologous with feathers. That was shown to be invalid by taxon exclusion here.

Today we’ll reconstruct
the crushed skull using DGS and nest this specimen in a cladogram using phylogenetic analysis (Fig. 4) in a few hours. Yang et al. were unable or unwilling to do either, even with firsthand access to the fossil and nine co-authors.

Figure 1. The NJU-57003 specimen and outline drawing, both from Yang et al. 2018. Various membranes and the overlooked sternal complex are colored in here.

Figure 1. The NJU-57003 specimen and outline drawing, both from Yang et al. 2018. Various membranes and the overlooked sternal complex and prepubes are colored in here. Clearly the uropatagia are separated here, as in Sharovipteryx. No wing membrane attaches below the knee.

Overlooked by Yang et al.
the sternal complex is quite large beneath the wide-spread ribs, a trait common to anurognathids. The torso, like the skull, would have been much wider than deep in vivo.

Figure 2. The skull elements of NJU-57003 colored to help alleviate the chaos of the crushed specimen. See figure 3 for the same elements reconstructed.

Figure 2. The skull elements of NJU-57003 colored to help alleviate the chaos of the crushed specimen. I can’t imagine betting able to interpret this skull without segregating each piece with a different color. See figure 3 for the same elements reconstructed with these colors.

As in other disc/flathead anurognathids
the palatal processes of the maxilla (red in Figs. 2, 3) radiate across the light-weight palate.  Yang et al. mislabeled these struts the ‘palatine’ (Fig. 1) following in the error-filled footsteps of other pterosaur workers who did not put forth the effort to figure things out.

The skull
is likewise supported by relatively few and very narrow struts. Contra Yang et al. 2018, who once again, mistakenly identify the toothy maxilla as an scleral ring (Fig. 1), the actual scleral rings (Figs. 2, 3) are complete and smaller within a large squarish orbit bounded ventrally by a deep jugal.

Figure 3. The skull of NJU-57003 reconstructed in animated layers for clarity. This is something the print media just cannot do as well. All elements are similar to those found earlier in other anurognathids.

Figure 3. The skull of NJU-57003 reconstructed in animated layers for clarity. This is something the print media just cannot do as well. All elements are similar to those found earlier in other anurognathids. Note the eyes, as in ALL pterosaurs, are in the back half of the skull.

Discodactylus megasterna (Yang et al. 2018; Middle-Late Jurassic, Yanlio biota, 165-160mya; NJU-57003) is a complete skeleton of a disc-skull anurognathid with soft tissue related to Vesperopterylus. The sternal complex is quite large to match the wider than tall torso. Distinct from other anurognathids, m4.1 does not reach the elbow when folded.

Figure 4. Subset of the LPT nesting Discodactylus with Vesperopterylus within the Anurognathidae.

Figure 4. Subset of the LPT nesting Discodactylus with Vesperopterylus within the Anurognathidae.

This specimen was introduced without a name
in a paper that incorrectly linked pterosaur filaments to dinosaur feathers (Yang et al. 2018), rather than with their true ancestor/relatives, the filamentous fenestrasaurs, Sharovipteryx and Longisquama, taxa omitted in Yang et al. and all workers listed below. Details here. The authors were unable to score traits for the skull and did not mention Vesperopterylus in their text.

Apparently the same artist
who originally traced the skull of Jeholopterus in 2003 (Fig. 5) also traced the present specimen (Fig. 1) with the same level of disinterest and inaccuracy. Compare the original image (Fig. 5 left) to a DGS image (Fig. 5 right). 

Figure 5. The original 2003 tracing of Jeholopterus (upper left) was inaccurate, uninformed and uninformative despite first hand access compared to the more informative and informed tracing created using DGS methods.

Why did these anurognathids have such long filaments?
Owls use similar fluffy feathers to silence their passage through air, first discussed earlier here.

The pterosaur experts weigh in the-scientist.com/news:
“I would challenge nearly all their interpretations of the structures. They are not hairs at all, but structural fibers found inside the wings of pterosaurs, also known aktinofibrils,” says pterosaur researcher David Unwin at the University of Leicester in the UK who was not part of the study. “They discovered lots of hair-like structures, but [don’t report any] wing fibers. I find that problematic.” Unwin suspects these fibers are likely to be present but have been mislabeled as feathers.  

This is a very important discovery,” says Kevin Padian, a palaeontologist at the University of California, Berkeley, “because it shows that integumentary [skin] filaments evolved in both dinosaurs and pterosaurs. That’s not surprising because they are sister groups, but it is good to know.”  

Padian draws attention to the pycnofibers’ “hair-like structure” as illustrating that they served as insulation. This is yet another characteristic of dinosaur and pterosaurs, along with high growth rate, pointing to their common ancestor as warm blooded.  “I wish the illustrations in the paper were better, but there is no reason to doubt them,” he adds.

Dr. Padian knows better.
He’s keeping the family secret by not mentioning fenestrasaurs (Peters 2000).

“The thing that is cool is that it bolsters the idea that pterosaurs and dinosaurs are sister taxa, if they are correct in interpreting these structures as a type of feather,” writes paleobiologist David Martill of the University of Plymouth in the UK, in an email. 

Dr. Martill knows better.
He’s keeping the family secret by not mentioning fenestrasaurs.

The specimens described in the paper are very interesting, agrees Chris Bennett, a palaeontologist at Fort Hays State University in Kansas, but in an emailed comment he describes the interpretation of the structures as problematic. “The authors’ characterization of the integumentary structures as ‘feather-like’ is inappropriate and unfortunate,” he writes. Some of the structures look like they could be from fraying or other decomposition, rather than feathers. Bennett adds that filamentous structures for insulation and sensation are fairly common, from hairy spiders to caterpillars to furry moths. “It seems to me to be premature to use filamentous integumentary structures to support a close phylogenetic relationship between pterosaurs and dinosaurs,” says Bennett. 

Dr. Bennett knows better.
He’s keeping the family secret by not mentioning fenestrasaurs.

Benton stands by his conclusion that pterosaurs wore plumage. Asked about the suggestion that the feathers could be wing fibers, he writes in an email, “Actinofibrils occur only in the wing membranes, whereas the structures we describe occur sparsely on the wings, but primarily over the rest of the body.”

Dr. Benton knows better.
He’s keeping the family secret by not mentioning fenestrasaurs. More details here.

References
Bennett SC 1996. The phylogenetic position of the Pterosauria within the Archosauromorpha. Zoological Journal of the Linnean Society 118:261-308.
Hone DWE and Benton MJ 2007.
An evaluation of the phylogenetic relationships of the pterosaurs to the archosauromorph reptiles. Journal of Systematic Palaeontology 5:465–469.
Hone DWE and Benton MJ 2009.
Contrasting supertree and total evidence methods: the origin of the pterosaurs. Zitteliana B28:35–60.
Peters D 2000. 
A Redescription of Four Prolacertiform Genera and Implications for Pterosaur Phylogenesis. Rivista Italiana di Paleontologia e Stratigrafia 106 (3): 293–336.
Yang et al. (8 co-authors) 2018. Pterosaur integumentary structures with complex feather-like branching. Nature ecology & evolution doi:10.1038/s41559-018-0728-7

 

SVP 2018: Resolving the Mesadactylus complex of individual bones

Figure 1. Mesadactylus restored from available data as an anurognathid.

Figure 1. Mesadactylus restored from available data as an anurognathid.

The BYU specimens attributed to 
Mesadactylus 
nest with anurognathids like ?Dimorphodon weintraubi (coeval and also from North America) in the large pterosaur tree (LPT). The specimen is Late Jurassic (Morrison Formation) in age.

Sprague and McLain 2018 conclude, “the taxonomic affinity of the genus is uncertain.” So why did they title their talk “Resolving…”

All they had to do
was Google “Mesadactylus.” Or run a phylogenetic analysis. The affinities of this specimen have been known here at the LPT since 2012.

References
Spratue M and McLain MA 2018. Resolving the Mesadactylus complex of Dry Mesa Quarry, Morrison Formation, Colorado. SVP Abstracts.

Mistralazhdarcho: a new pterosaur, but not an azhdarchid

Vullo et al. 2018 bring us a new small ‘azhdarchid’
known from a few 3D bones. In the large pterosaur tree (LPT, 236 taxa) Mistralazhdarcho nests with tiny Nemicolopterus and mid-sized Shenzhoupterus (Fig. 1). Mistralazhdarcho is more than twice as tall as Shenzhoupterus with similar gracile cervicals, a longer radius and shorter metacarpus. Distinct from Shenzhoupterus, the mandible is gracile, more like that of Nemicolopterus.

Figure 1. Mistralazhdarcho compared to reconstructions of Shenzhoupterus and Nemicolopterus.

Figure 1. Mistralazhdarcho compared to reconstructions of Shenzhoupterus and Nemicolopterus. A longer antebrachium is found in Mistalazdarcho.

A downturned dentary
is a trait found in this clade of pterosaurs, and to a lesser extent in sister sinopterids.

The small prominence at the ‘bend’ of the mandible
in Mistralazhdarcho is a curious trait not visible in Shenzhoupterus due to closed jaws in situ. Nemicolopterus might preserve that trait, but a humerus is under the mandible exactly at that point, making it difficult to determine in photos.

A warped deltopectoral crest,
like the one found in Mistalazdarcho (Fig. 1), is not found in azhdarchids. And look at the size range in this clade!

Having reconstructions for direct comparisons,
and a large cladogram that is regularly adding new taxa are tools the LPT and www.ReptileEvolution.com offer freely online to paleontologists worldwide. Best to test here rather than trust your hunch elsewhere.

References
Vullo R, Garcia G, Godefroit P, Cincotta A, and Valentin X 2018. Mistralazhdarcho maggii, gen. et sp. nov., a new azhdarchid pterosaur from the Upper Cretaceous of southeastern France. Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2018.1502670.

Fish nibbles on Pteranodon metacarpal

Figure 1. Fish teeth compared to grazed Pteranodon metacarpal

Figure 1. Fish teeth compared to grazed Pteranodon metacarpal

Ehret and Harrell 2018
bring us news from Alabama of two distinct sets of tooth marks on a Pteranodon (Fig. 2) metacarpal (Fig. 1). They report:

“The Pteranodon specimen exhibits serrated teeth marks on the surface of the bone and a second set of larger, unserrated teeth marks unlike those of any contemporary shark species. These feeding traces compare favorably with the tooth spacing and morphology of Squalicorax kaupi, and a small to moderate-sized saurodontid fish, such as Saurodon or Saurocephalus, respectively. In both instances, feeding traces appear to be scavenging events due to the lack of any healing or bone remodeling. The specimen represents a pterosaur that either fell into marine waters or was washed out from nearshore areas and then scavenged by both a chondrichthyan and osteichthyan.”

“Many fossils from late Cretaceous Alabama appear to have been nibbled by sharks, including sea turtles and dinosaurs, which are often ‘covered in predation marks,’ says Ehret.”

NatGeo publicized the find by talking to some pterosaur experts, “Pterosaurs actually had a lot of meat on their skeletons,” says Michael Habib, a pterosaur expert at the University of Southern California who was not involved with the latest find. “They were not the skinny animals often depicted in films and art. The flight muscles in particular would have made a great meal.”

Pterosaur metacarpals,
like all metacarpals, actually are sinewy and have little to no associated muscle.

Habib adds,
“Pteranodon also inhabited this coastal environment during the late Cretaceous, making a living snatching smaller fish from the shark-filled waters. Pterosaurs could float, but being less buoyant than birds, they probably didn’t sit on the surface for long. Some species, including Pteranodon, did likely plunge into the water for prey. “They could then quickly take back off from the surface. But these diving pterosaurs might have been vulnerable to sharks just after they entered the water,” he says.

M. Witton concluded,
“It’s nice to know what species were interacting in this way.”

Ehret corrected the pterosaur experts,
“It’s also possible that the animal died near the shore and was scavenged when it washed out to sea.”

Figure 3. Triebold Pteranodon in floating configuration. Center of balance marked by cross-hairs.

Figure 2 Triebold Pteranodon in floating configuration. Center of balance marked by cross-hairs.

Contra Habib’s statement
Pteranodon was at least as buoyant as a pelican. It has been widely known for over a century that pterosaur bones are thinner than bird bones and Pteranodon metacarpals, in particular, were hollow like pontoons (Fig. 2).

Ultimately
the bite marks represent curiosity, not predation, a point understood by Ehret and Harrell.

References
Ehret DJ and Harrell TL Jr. 2018. Feeding traces of a Pteranodon (Reptilia: Pterosauria) bone from the late Cretaceous (Campanian) Mooreville Chalk in Alabama, USA. Palaios 33(9):414–418.

www.natgeo.com/

Big pterosaurs: big or little wing tips

Earlier and below (Fig. 2) we looked at large and giant pterosaur wings comparing them to the largest flying birds, including one of the largest extant flying birds, the stork, Ciconia, and the extinct sheerwater, Pelagornis, the largest bird that ever flew.

FIgure 2. A basal pteranodotid, the most complete Pteranodon, the largest Pteranodon skull matched to the largest Pteranodon post-crania compared to the stork Ciconia and the most complete and the largest Quetzalcoatlus

FIgure 1. A basal pteranodotid, the most complete Pteranodon, the largest Pteranodon skull matched to the largest Pteranodon post-crania compared to the stork Ciconia and the most complete and the largest Quetzalcoatlus. Note the much reduced distal phalanges in the complete and giant Quetzalcoatlus, distinct from the Pteranodon species.

Today
we’ll look at how the largest Pteranodon (Figs. 1, 4) compares to much larger pterosaurs, like Quetzalcoatlus northropi (Figs. 1, 2) that have vestigial wingtips similar to those of the  much smaller flightless pre-azhdarchid, SOS 2428 (Fig. 3).

Note the tiny three distal phalanges
on the wing of the largest Quetzalcoatlus, distinct from the more typical elongate and robust distal phalangeal proportions on volant pterosaurs of all sizes. Much smaller definitely flightless pterosaurs, like SOS 2428, shrink those distal phalanges, too. That’s the pattern when pterosaurs lose the ability to fly.

Figure 2. Q. northropi and Q. sp. compared to Ciconia, the stork, and Pelagornis, the extinct gannet, to scale. That long neck and large skull of Quetzalcoatlus would appear to make it top heavy relative to the volant stork, despite the longer wingspan. Pteranodon and other flying pterosaurs do not have such a large skull at the end of such a long neck (Fig. 1). The longer wings of pelagornis show what is typical for a giant volant tetrapod, and Q. sp. comes up short in comparison.

Figure 2. A previously published GIF animation. Q. northropi and Q. sp. compared to Ciconia, the stork, and Pelagornis, the extinct gannet, to scale. That long neck and large skull of Quetzalcoatlus would appear to make it top heavy relative to the volant stork, despite the longer wingspan. Pteranodon and other flying pterosaurs do not have such a large skull at the end of such a long neck (Fig. 1). The longer wings of pelagornis show what is typical for a giant volant tetrapod, and Q. sp. comes up short in comparison.Today we’ll compare the wingspan of the largest Quetzalcoatlus to the largest and more typical Pteranodon species (Fig. 2).

Unfortunately
pterosaur workers refuse to consider taxa known to be flightless, like SOS 2428 (Peters 2018). It’s easy to see why they would be flightless (Fig. 3). Scaled to similar snout/vent lengths with a fully volant pterosaur like n42 (BSPG 1911 I 31) the wing length and chord are both much smaller in the flightless form.

Lateral, ventral and dorsal views of SoS 2428

Figure 3. Lateral, ventral and dorsal views of the flightless SoS 2428 (Peters 2018) alongside No. 42, a volant sister taxon.

Comparing the largest ornithocheirid,
SMNK PAL 1136, to the largest Pteranodon (chimaera of largest skull with largest post-crania in Fig. 4) shows that large flyers have elongate distal phalanges, distinct from body and wing proportions documented in the largest azhdarchids, like Quetzalcoatlus.

Figure 5. Largest Pteranodon to scale with largest ornithocheirid, SMNS PAL 1136.

Figure 4. Largest Pteranodon to scale with largest ornithocheirid, SMNS PAL 1136. Note the long distant wing phalanges on both of these giant flyers. This is what pterosaurs evolve to if they want to continue flying. And this is how big they can get and still fly. Giant azhdarchids exceed all the parameters without having elongate wings. Note: the one on the left has a longer wingspan whir the one on the right has a more massive torso and skull together with more massive proximal wing bones and pectoral girdle. On both the free fingers are tiny, parallel oriented laterally and slightly tucked beneath the big knuckle of the wing finger. The pteroid points directly at the deltopectoral crest. 

As the largest Pteranodon and largest ornithocheirid (SMNS PAL 1136)
(Fig. 4) demonstrate, as flying pterosaurs get larger, they retain elongate distal wing phalanges. And big, robust phalanges they are.

By contrast in azhdarchids and pre-azhdarchids
there is a large size bump after n42 (BSPG 1911 I 31) the fourth wing phalanx either disappears (see Microtuban and Jidapterus) or shrinks to a vestige. Then there’s Zhejiangopterus (Fig. 5), with a big pelvis, gracile forelimbs and a giant skull on a very long neck. Just that neck alone creates such a long lever arm that the pterosaur is incapable of maintaining a center of balance over or near the shoulder joints.

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 5. 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 mentioned earlier, becoming flightless permitted, nay, freed azhdarchid pterosaurs to attain great size. They no longer had to maintain proportions that were flightworthy. Instead they used their shortened strut-like forelimbs to maintain a stable platform in deeper waters. And when they had to move in a hurry, their wings could still provide a tremendous amount of flurry and thrust (Fig. 6) for a speedy getaway.

Quetzalcoatlus running like a lizard prior to takeoff.

Figure 6. Quetzalcoatlus running without taking off, using all four limbs for thrust. That long lever arm extending to the snout tip in front of the center of gravity is not balanced in back of what would be the center of lift over the wings

For the nitpickers out there…
some specimens of Nyctosaurus (UNSM 93000, Fig. 7) also have but three wing phalanges, but they are all robust. The distal one is likely the fourth one because it remains curved. Phalanges 2 and 3 appear to have merged, or one of those was lost. Compare that specimen to a more primitive Nyctosaurus FHSM VP 2148 with four robust wing phalanges.

Figure 5. Cast of the UNSM 93000 specimen of Nyctosaurus. Missing parts are modeled here.

Figure 5. Cast of the UNSM 93000 specimen of Nyctosaurus. Missing parts are modeled here.

References
Peters D 2018. First flightless pterosaur (not peer-reviewed). PDF online.