AMNH 4908: at the genesis of Nyctosaurus and Pteranodon

Bennett 2017 described AMNH 4908:
“The smallest relatively complete previously known specimen of Pteranodon, AMNH 4908, consists of a partial trunk skeleton and tail, scapulocoracoid, humerus through WP2, both femora and tibiae, and a disarticulated foot (Bennett, 2001; Table 1), had an estimated wingspan in life of 3.33 m.”

AMNH 4908 is small because
it is primitive (Figs. 1–3), closer to its smaller germanodactylid ancestors (Fig. 4), not because it is ontogenetically young. That point was overlooked by Bennett 2017 who decided not to include a comprehensive phylogenetic analysis then or earlier (Bennett 1991, 1992). He thought smaller specimens, like AMNH 4908, were female.

Figure 1. The Niobrara specimen AMNH4908 in situ.

Figure 1. The Niobrara specimen AMNH4908 in situ. See figure 3 for reconstruction.

It is worth noting
AMNH 4908 has a type of pelvis with unfused ischia, a morphology common to Nyctosaurus. Earlier Bennett 1991, 1992 decided a large Nyctosaurus pelvis belonged to a female Pteranodon, which started that gender myth. Unfortunately, Bennett never used phylogenetic analysis to lump and split Pteranodon, the subject of his 1991 PhD thesis. Instead he relied on a statistical analysis, which led him astray. The large pterosaur tree (LPT, 246 taxa) is the only published pterosaur cladogram to include more than one or two Pteranodon and Nyctosaurus taxa. When you really want to know something in systermatics, use a comprehensive cladogram, not a graph.

Figure 2. Subset of the LPT focusing on pterandontids and their ancestors going back to Late Jurassic germanodactylids. AMNH 4908 nests between the Nyctosaurus clade and the Pteranodon clade.

Figure 2. Subset of the LPT focusing on pterandontids and their ancestors going back to Late Jurassic germanodactylids. AMNH 4908 nests between the Nyctosaurus clade and the Pteranodon clade.

Also worth noting
is the deltopectoral crest of the humerus in AMNH 4908, which has a round tongue shape like that of its predecessors among Eopteranodon and Germanodactylus (Fig. 4), rather than the hatchet shape found in Nyctosaurus or the warp found in Pteranodon.

Figure 3. AMNH 4908 pterosaur reconstructed. Transitional between Nyctosaurus and Pteranodon. Note the big feet, tiny sternal complex and round tip deltopectoral crest. Note the massive radius + ulna.

Figure 3. AMNH 4908 pterosaur reconstructed. Transitional between Nyctosaurus and Pteranodon. Note the big feet, tiny sternal complex and round tip deltopectoral crest. Note the massive radius + ulna.

Of course, any taxon basal to Nyctosaurus and Pteranodon
is also going to be close to the SMNK PAL 6592 specimen (Fig. 4) attributed to Germanodactylus.

Matching YPM 1179 to the post-crania of SMU 76476 (Myers 2010) and overprinted with SMNK PAL 6592. The resemblance is indeed remarkable.

Figure 4. Matching YPM 1179 to the post-crania of SMU 76476 (Myers 2010) and overprinted with SMNK PAL 6592. The resemblance is indeed remarkable.

Ever since 2003, traditional pterosaur workers (with PhDs)
have been linking toothless pteranodontids to toothy ornithocheirids. That they continue to do so, (due to taxon exclusion) is embarrassing to the profession.

Figure 2. The Tanking-Davis specimen compared to other forms. Specimen w and specimen z appear to be the closest to the Tanking-David specimen. Specimen 'w' = Pteranodon sternbergi? USNM 12167 (undescribed). Specimen 'z' = Pteranodon longiceps? Dawndraco? UALVP 24238. Click to enlarge.

Figure 5. Pteranodon and Nyctosaurus skulls. Click to enlarge.

Figure 6. AMNH 4908 specimen compared to a large Pteranodon specimen UNSM 50036.

Figure 6. AMNH 4908 specimen compared to a large Pteranodon specimen UNSM 50036.

In similar fashion and shame,
traditional paleontologists continue to insist that pterosaurs arose from Euparkeria, Scleromochlus or Erythrosuchus according to several authors, or from Macrocnemus bassaniiPostosuchus kirkpatricki and Herrerasaurus ischigualastensis according to Dalla Vecchia 2019. More closely related taxa (LangobardisaurusCosesaurus, Sharovipteryx and Longisquama) were validated as better pterosaur ancestors in four phylogenetic analyses 20 years ago.


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 2017. New smallest specimen of the pterosaur Pteranodon and ontogenetic niches in pterosaurs. Journal of Paleontology. pp.1-18. 0022-3360/15/0088-0906
doi: 10.1017/jpa.2017.84
Dalla Vecchia FM 2019. Seazzadactylus venieri gen. et sp. nov., a new pterosaur (Diapsida: Pterosauria) from the Upper Triassic (Norian) of northeastern Italy. PeerJ 7:e7363 DOI 10.7717/peerj.7363

Thanks to Alex Schiller for posting this specimen on Facebook.

https://pterosaurheresies.wordpress.com/2012/04/09/female-pterosaurs/

http://www.reptileevolution.com/pteranodon-postcrania.htm

Restoring the tip of the Pteranodon rostrum in UALVP 24238

Another short one today.
Everyone in pterosaur land has wondered what shape and length the missing tip of the rostrum took on in the UALVP 24238 specimen of Pteranodon (improperly called ‘Dawndraco‘, Fig. 1). That’s because, unlike most other specimens, the upper and lower margins don’t converge to form a triangle in lateral view, but remain essentially parallel for most of their length.

Figure 1. The missing tip of the UALVP 24238 skull can be restored like the similar Tanking-David specimen skull tip. The dorsal rim is straight. The ventral rim is convex.

Figure 1. The missing tip of the UALVP 24238 skull can be restored like the similar Tanking-Davis specimen skull tip. The dorsal rim is straight. The ventral rim is convex.

The privately held
Tanking-Davis specimen of Pteranodon is the most similar to the UALVP 24238 rostrum. Yes, it converges, but at a very shallower angle. Anteriorly the ventral margin curves up to meet the very straight dorsal margin. The same can be imagined/restored for the missing UALVP 24238 rostrum, and the result appears to be reasonable.

Figure 2. The UALVP 24238 specimen of Pteranodon is largely complete and here the tip of the rostrum is restored like that of a related specimen.

Figure 2. The UALVP 24238 specimen of Pteranodon is largely complete and here the tip of the rostrum is restored like that of a related specimen.

As in most pterosaur genera,
no two specimens are identical. See the panoply of known relatively complete Pteranodon skulls to scale and in phylogenetic order here.

Flugsaurier 2018: The KUVP 967 specimen of Pteranodon

Flugsaurier 2018 part 2
Since the purpose of the symposium is increase understanding of pterosaurs, I hope this small contribution helps.

Brandão RS and Rodrigues T 2018
supports the integrity of the generic name, “Dawndraco” for a classic Pteranodon specimen. We looked at that issue earlier here and here. Bottom line: in the absence of a phylogenetic analysis that includes as many Pteranodon and outgroup specimens as in the large pterosaur tree (LPT, 233 taxa) nobody has any business lumping and separating one or two cherry-picked taxa from the long list currently attributed to Pteranodon. The evidence shows they are all congeneric (in the very stretched out pterosaur sense) and none are conspecific. The authors, like previous authors, do not realize their ‘Dawndraco‘ nests between several other Pteranodon specimens. That never happens elsewhere on anyone’s cladogram.

Brandão RS and Rodrigues T 2018 conclude:
“We regard Dawndraco kanzai as valid, and KUVP 967 as a referred specimen.”

Figure 1. The Pteranodon jaws KUVP 967.

Figure 1. The Pteranodon jaws KUVP 967.

The KUVP 967 specimen
(Figs. 1, 2) is known from a partial rostrum and partial mandible (brown and blue) graphically scaled and placed over other taxa. It was not included in the LPT because so little can be scored for it. Graphically, no two Pteranodon skulls are good matches for KUVP 967. As in Rhamphorhynchus, they are all individuals, only a few conspecific.

In figure 2
W is the undescribed USNM 12167 specimen. Z is the described UALVP 24238 specimen previously referred to Pteranodon, then to Dawndraco. The anterior rostrum  of KUVP 967 is too short to match specimen Z (UALVP 24238 ) and too tall to match specimen W (USNM 12167).

Figure 1. The KUVP 967 specimen (blue and brown) compared to two more completely known Pteranodon specimens.

Figure 2. Segment of the Pteranodon skulls page at ReptileEvolution.com. The KUVP 967 specimen (blue and brown) compared to two more completely known Pteranodon specimens. The anterior rostrum  of KUVP 967 is too short to match specimen Z (UALVP 24238 ) and too tall to match specimen W (USNM 12167).

References
Brandão RS and Rodrigues T 2018. Reappraisal of Dawndraco kanzai as a valid taxon. Flugsaurier 2018: the 6th International Symposium on Pterosaurs. Los Angeles, USA. Abstracts: 17—18.
Kellner AWA 2010. Comments on the Pteranodontidae (Pterosauria, Pterodactyloidea) with the description of two new species. Anais da Academia Brasileira de Ciências, 82(4), 1063–1084.
Martin-Silverstone E. et al. 2017. Reassesment of Dawndraco kanzai Kellner, 2010 and reassignment of the type specimen to Pteranodon sternbergi Harksen, 1966. Vertebrate Anatomy Morphology Palaeontology, 3, 47§–59.

A Brazilian stem pteranodontid, and Brazil wants its fossils back!

Figure 1. The cf.Tupuxuara specimen is larger than sister taxa in the LPT.

Figure 1. The cf.Tupuxuara specimen is larger than sister taxa in the LPT.

cf.Tupuxuara (SMNK??? Elgin 2014, Early Cretaceous). Originally considered close to Tupuxuara, here this specimen nests between Eopteranodon and the base of the Pteranodontia. The metacarpals and antebrachium are relatively short. The large pentagonal sternal complex anchors large flight muscles. Distinct from the Pteranodontia, but like the Eopteranodon clade, the carpal and tarsal elements were not co-ossified. The ventral pelvis remained open, as in Eopteranodon and most tested nyctosaurids. In other words, this is NOT a female…necessarily.

Figure 2. Early Cretaceous cf.Tupuxuara from the Elgin 1914 dissertation. This taxon nests between the Solnhofen specimen B St 1878 VI 1 and Eopteranodontia + Pteranodontia in the LPT, far from Tupuxuara. Reconstruction from underlying in situ specimen from the Elgin 2014 dissertation available online.

Figure 2. Early Cretaceous cf.Tupuxuara from the Elgin 1914 dissertation. This taxon nests between the Solnhofen specimen B St 1878 VI 1 and Eopteranodontia + Pteranodontia in the LPT, far from Tupuxuara. Reconstruction from underlying in situ specimen from the Elgin 2014 dissertation available online. Missing parts filled in.

You might want to think of this pterosaur
as the first of the large Pteranodontia, still nesting with the Germanodactylus clade not leading to dsungaripterids, Shenzhoupterus and tapejarids, including Tupuxuara). Elanodactylus is another large member of this clade (Fig. 3).

Figure 3. Subset of the large pterosaur tree (LPT) with the addition of cf. Tupuxuara apart from Tupuxuara and at the base of the Pteranodontia.

Figure 3. Subset of the large pterosaur tree (LPT) with the addition of cf. Tupuxuara apart from Tupuxuara and at the base of the Pteranodontia.

The Elgin 2014 thesis was completed in May 2014.
Just a few months earlier, in March 2014 a paper appeared in Nature entitled, “Brazil clamps down on illegal fossil trade.” The first sentence reads, “Thirteen people are scheduled to go on trial in Brazil for smuggling fossils out of the country, apparently to private collectors and to museums in Germany and the United Kingdom.” Do you think Dr. Elgin was worried? Evidently not. In his PhD thesis Elgin wrote, The large numbers of [Chapada do Araripe] specimens that at the time of writing lacked any full or proper description was one of the major influences in the creation of this body of work, creating a catalogue of fossils that increase our understanding of this enigmatic group and permitting ready access to photographs and descriptions for future workers.” And for making those images available, Dr. Elgin, thank you!

Dr. Elgin further notes
“Brazil has banned the commercial sale of all fossil originating from its territories since 1942.” Then concludes, “The pterosaurs described within this body of work are presented for the good of the scientific community. While discouraging illicit trafficking is to be encouraged, the fact that the featured specimens are interred within a registered museum, rather than ending up within a private institution as would have certainly been their fate otherwise, guarantees the continued and universal access to any and all persons, to the benefit of the international community.”

Worried about the loss of Brazilian fossils to German museums,
Brazilian paleontologist, Alexander Kellner, cites the loss of cultural heritage. On the other hand, English paleontologist, David Martill quips, Knowing “dodgy” people is the only way to get samples, because the DNPM ignores requests to dig.” Brazilian paleontologist, Max Langer says, “Fossils must be kept in the country to help to improve Brazilian science.” And he expects fellow researchers to hold Brazil’s laws in higher regard than the private collectors who also fuel the trade.

David Martill expressed more of his thinking
in this online report, “In an email interview, Martill said that he “doesn’t care a damn how the fossil came from Brazil”, because that is “irrelevant to the scientific significance of the fossil. I am critical of all laws that interfere with the science of paleontology; and blanket bans on fossil collecting are indiscriminatory and only hinder science, No countries existed when the animals were fossilized.”

Bottom line:
Firsthand access to fossils… can sometimes get you into trouble with Brazil. You can see how the side line up here, with Brazilians hoping to stop exports and Europeans hoping to continue exports.

More tomorrow
on the Elgin dissertation…

References
Elgin RA 2014. Palaeobiology, Morphology, and Flight Characteristics of Pterodactyloid Pterosaurs. Innaugural Dissertation. Zur Erlangung der Doktorwürde Fakultät für Chemie und Geowissenschaften Institut für Geowissenschaften Ruprecht-Karls-Universität Heidelberg. Available online here.

Smallest Pteranodon: Bennett 2017

Figure 1. The new small Pteranodon wing, FHSM 17956, compared to Ptweety and the adult NMC41-358 specimen.

Figure 1. The new small Pteranodon wing, FHSM 17956, compared to Ptweety and the adult NMC41-358 specimen. At half the size of the adult, the FHSM specimen would have been 4x the size of a hatchling.

A new small partial wing specimen of Pteranodon
discovered by Glen Rockers, was described by Bennett 2017 (Figs. 1-4). It is virtually identical to similar bones in Ptweety (Fig. 1), a specimen now lost to science and half the size of the Triebold specimen NMC41-358, which is similarly gracile. Click here to see more robust Pteranodon adults compared to the new small FHSM specimen.

Figure 2. FHSM 17956 compared to Ptweety. They are virtually identical, though Ptweety looks like a juvenile of a more robust variety of Pteranodon, thus a younger specimen because adults would be larger.

Figure 2. FHSM 17956 compared to Ptweety. They are virtually identical, though Ptweety looks like a juvenile of a more robust variety of Pteranodon, thus a younger specimen because adults would be larger.

Young (small) Pteranodon specimens
were essentially unknown prior to the Bennett paper. So this is important news.

Figure 2. Small Pteranodon, FHSM 17956, carpus insitu and reconstructed. Here several bones were reidentified.

Figure 3. Small Pteranodon, FHSM 17956, carpus insitu and reconstructed. Here several bones were reidentified. See reconstruction in figure 3. It demonstrates that all the newly identified parts fit together.

Unfortunately
a reconstruction based on Digital Graphic Segregation (DGS, Fig. 4) shows that Bennett, widely known as THE expert on Pteranodon going back to his PhD thesis, misidentified several carpal bones here. In his defense, that was easy to do. The distal carpal is beneath the other carpal bones and it has splinters that extend beyond it. Rather than using DGS, Bennett chose to outline bones the old fashioned way. This leads to problems that can be solved when you color each bone and bone splinter THEN test your colors with a reconstruction. Bennett provided no reconstruction that tested his outline tracings. Bennett also overlooked manual digit 5. The fragment (FR) probably comes from the crushed and splintered distal carpal. Bennett reported, “All carpal elements are severely deformed by compression such that they preserve little of their original morphology…” That’s because he misidentified elements that are otherwise identical to those of adult specimens.

Figure 3. Small Pteranodon (FHSM 17956) carpus reconstructed after several bones were reidentified.

Figure 4. Small Pteranodon (FHSM 17956) carpus reconstructed after several bones were reidentified.

 

Bennett also upholds several invalid paradigms

  1. Other small, short crested Pteranodon specimens represent young ones. Actually they represent taxa closer to the outgroup, Germanodacytylus
  2. Short-crested specimens are females. No male/female pairs have ever been documented. Rather short-crested taxa are closer to the crestless outgroup. 
  3. Large pelvis specimens  are females. No, they are large nyctosaurs. 
  4. Small size Rhamphorhynchus were juveniles of larger ones. No, phylogenetic analysis indicates a period of phylogenetic miniaturization followed the genesis of Rhamphorhynchus from larger Campylognathoides ancestors. Bone histology would include juvenile bone tissue in adults of these small, precocial and fast-breeding taxa. It is important that someday Bennett runs a phylogenetic analysis, something he told me decades ago was critical to understanding taxonomy. 
  5. There is no such thing as manual digit 5 in pterosaurs. He overlooked it here. 

Bennett now realizes:
“A new juvenile specimen of Pteranodon collected from the Smoky Hill Chalk Member is so small that it challenges the interpretation of rapid growth to large size before flying and feeding (Bennett, 2014a).” As everyone knows now, hatchling pterosaurs were able to fly shortly after hatching. To his credit, Bennett continues, “The interpretation of rapid growth while under parental care is rejected.”

Bennett examined the specimen under stereo microscope
and made mistakes here re-identified on a computer monitor applying colors to each bone to visually segregate one from another and facilitate accurate reconstruction. This is something that cannot take place using old-fashioned stereo microscopes.

Bennett occasionally
misidentifies small pterosaur bones. This was documented here dealing with the flat-headed anurognathid SMNS 81928, in which he considered the mandible a giant sclerotic ring in the front half the skull, different from all other pterosaurs. Bennett 2008 promoted an invalid hypothesis on the origin of the pterosaur wing based on imagination rather than taxa, documented here. Bennett’s (2007) interpretation of pteroid articulation against the preaxial carpal. was invalidated by Peters 2009 who nested it on the anterior radiale (Fig. 4).

Note
The extensor tendon process is articulated with the rest of m4.1, as in all Pteranodon specimens. Bennett once considered unfused  extensor tendon processes a sign of immaturity. This is not correct. As reported earlier, since pterosaurs are lepidosaurs they display lepidosaur fusion traits, typically not ontogenetic, but phylogenetic. As an example, in Nyctosaurus the extensor tendon process remains unfused, distinct from Pteranodon. Bennett insists that the extensor tendon process in the juvenile specimen is unfused but notes that the fragile cortical bone was lost during preparation. And just think about it.. the carpals, typically wrapped tightly in ligaments were scattered while the extensor tendon process didn’t move during taphonomy. By contrast, in Nyctosaurus the extensor tendon process popped off before the toes disarticulate.

Bennett avoid mentioning or citing
work by Peters 2009, which disputed Bennett 2007, who articulated the pteroid with the preaxial carpal. In order to do so, Bennett 2017 did not cite Bennett 2007, but did manage to cite nearly every other one of his papers. Kids.. sometimes you have to look for what’s not mentioned.

Pteranodon variety
is best seen and appreciated by direct comparison of the skulls and the post-crania. FHSM 17956 is a juvenile of a gracile form, similar to the Triebold specimen NMC41-358 (Fig. 1), a short-crested gracile variety. By contrast, Ptweety appears to be more similar to the more robust long-crested taxa. 

Bennett describes ontogenetic niches
for hatchling, juvenile and adult Pteranodon. This is necessary for 8x smaller hatchlings incapable of handling adult-sized prey.

In Bennett’s Acknowledgements he reports, 
“Constructive reviews from M. Witton and L. Codorniú led to improvements in the manuscript, and an anonymous reviewer disagreed with everything.” That anonymous reviewer was not me. That would be blackwashing. I always try to find something of value in any manuscript I review, even if I disagree with some of what is presented.

Bennett first described this taxon
in a 2014 SVP abstract. See how long traditional studies take to get published? I was just about to call Chris to see if he was okay. I’m glad to see he is still out there publishing important specimens.

References
Bennett SC 2007. Articulation and Function of the Pteroid Bone of Pterosaurs. Journal of Vertebrate Paleontology 27(4):881–891.
Bennett SC 2008. Morphological evolution of the forelimb of pterosaurs: myology and function. Pp. 127–141 in E Buffetaut and DWE Hone eds., Flugsaurier: pterosaur papers in honour of Peter Wellnhofer. Zitteliana, B28.
Bennett SC 2017. New smallest specimen of the pterosaur Pteranodon and ontogenetic niches in pterosaurs. Journal of Paleontology. pp.1-18. 0022-3360/15/0088-0906
doi: 10.1017/jpa.2017.84
Peters D 2009. A reinterpretation of pteroid articulation in pterosaurs. Journal of Vertebrate Paleontology 29:1327-1330.

There’s a Pteranodon wing at the University of Missouri

No doubt
it was reassembled into its present position, despite the in situ appearance.

Figure 1. There is just no way to avoid reflections on this stairwell specimen of Pteranodon, if you want to capture the whole specimen in one shot from a distance.

Figure 1. There is just no way to avoid reflections on this stairwell specimen of Pteranodon, if you want to capture the whole specimen in one shot from a distance.

Mizzou has very few other vertebrate fossils.
The University of Missouri (Mizzou) Geology Department has a wonderful and complete small ichthyosaur from the Holzmaden and they have a partial parasuchian skull from the Petrified Forest. I don’t think the Mizzou Pteranodon wing has a number. Small portions, like the wrist and free fingers are restored.

Figure 1. The Mizzou Pteranodon wing is average in size and not very robust or gracile compared to others shown here. Click to enlarge.

Figure 2. The Mizzou Pteranodon wing is average in size and not very robust or gracile compared to others shown here. Click to enlarge.

If we take the wing at face value
and place it in context with other Pteranodon wings (Fig. 2), we find that it is not the largest, nor the smallest, not the most robust, nor the most gracile. The scapulocoracoid is relatively small. This could be a chimaera.

And while we’re on the subject of variation,
it is worthwhile to consider the post-cranial variation in Pteranodon, a subject we touched on earlier here and in figure 2, but has not been adequately addressed elsewhere.  The lack of more than a few skulls matched to post-crania (Fig. 2) has hampered efforts, but a decent cladogram of Pteranodon interrelationships can still be managed.

Figure 2. The Tanking-Davis specimen compared to other forms. Specimen w and specimen z appear to be the closest to the Tanking-David specimen. Specimen 'w' = Pteranodon sternbergi? USNM 12167 (undescribed). Specimen 'z' = Pteranodon longiceps? Dawndraco? UALVP 24238. Click to enlarge.

Figure 2. The Tanking-Davis specimen compared to other forms. Specimen w and specimen z appear to be the closest to the Tanking-David specimen. Specimen ‘w’ = Pteranodon sternbergi? USNM 12167 (undescribed). Specimen ‘z’ = Pteranodon longiceps? Dawndraco? UALVP 24238. Click to enlarge.

Pushback on ‘Dawndraco’ (Pteranodon UALVP 24238)

Figure 1. Pteranoodn (Dawndraco) UALVP 24238 in situ, with Martin-Silverstone tracing applied, with mandible moved and missing parts colorized. The putative rostral tip looks more like displaced manus elements.

Figure 1. Pteranoodn (Dawndraco) UALVP 24238 in situ, with Martin-Silverstone tracing applied, with mandible moved and missing parts colorized. The putative rostral tip looks more like displaced manus elements. The crest and distal wing finger do not belong to the original specimen.

A new paper by Martin-Silverstone et al. 2017
disputes the earlier study by Kellner 2010, giving a new generic name to a well-preserved putative Pteranodon specimen, UALVP 24238, Figs. 1-3). They also write: “The re-evaluation of Pteranodon sensu lato by Kellner (2010) is troubling for pterosaur palaeontology, as so much of our understanding of pterosaur ontogeny and growth stem from Bennett’s work on Pteranodon and the conclusion that Pteranodon specimens can be divided into two closely and perhaps anagenetically related species.” Bennett’s conclusions were disputed earlier here, here and here, and are nowhere in evidence here (Fig. 2). Praise for Bennett’s work needs to be limited to those items that stand the tests of closer scrutiny and analysis, Gender and ontogenetic differences recovered in Bennett’s statistical analyses are not recovered in phylogenetic analysis.

Figure 2. The Tanking-Davis specimen compared to other forms. Specimen w and specimen z appear to be the closest to the Tanking-David specimen. Specimen 'w' = Pteranodon sternbergi? USNM 12167 (undescribed). Specimen 'z' = Pteranodon longiceps? Dawndraco? UALVP 24238. Click to enlarge.

Figure 2. The Tanking-Davis specimen compared to other forms. Specimen w and specimen z appear to be the closest to the Tanking-David specimen. Specimen ‘w’ = Pteranodon sternbergi? USNM 12167 (undescribed). Specimen ‘z’ = Pteranodon longiceps? Dawndraco? UALVP 24238. Click to enlarge.

As you can see (Fig. 2) NONE
of the known Pteranodon-grade skulls would be considered conspecific in the modern world, and few would be considered congeneric. Size and crest size differences are without a doubt phylogenetic (contra Bennett and Martin-Sivlerstone et al.) as demonstrated in the large pterosaur tree. You can’t get large or have a large crest without evolving from smaller progenitors. It may also be the case that Pteranodon, like pterosaurs in general were extremely individually variable within a genus, but we’d need a time machine or a mass fossil assemblage for that.

Moreover,
UALVP 24238 had a tiny cranium, very different from the large cranium of P. sternbergi (FHSM VP 339, Fig. 2).

Figure 3. The UALVP specimen of Pteranodon. Note the lack of taper in the rostrum along with the small size of the orbit.

Figure 3. The UALVP specimen of Pteranodon. Note the lack of taper in the rostrum along with the small size of the orbit.

From the Martin-Silverstone et al. 2017 abstract:
“The previous most comprehensive study on Pteranodon [Bennett 1991, 1992m 19994, 2001] recognized two species: P. longiceps and P. sternbergi, but complete skeletons of Pteranodon are rare. One of the best preserved (UALVP 24238) has been identified as both P. sternbergi and as a new genus and species, Dawndraco kanzai. Here, the specimen is redescribed, additional portions of the rostrum are identified for the first time, new details of the specimen’s provenance and preparation history are presented, and its taxonomic placement is discussed. Whereas the shape of the rostrum appears at first glance to distinguish it from known Pteranodon, this feature is more parsimoniously interpreted in the context of sexual dimorphism; a male has a longer and therefore more shallowly tapering rostrum. Metrics from this specimen, and from published photographs and illustrations, support the conclusion that the rostrum of UALVP 24238 is not unique, and so provides no grounds for recognition of a taxon distinct from Pteranodon sternbergi. Other putatively unique features of UALVP 24238 are examined and found unconvincing.”

The rostrum is not the key trait that separates
UALVP 24238 from P. sternbergi (Fig. 2). It’s the cranium (among comparable elements preserved). The two species are related, but not conspecific. A phylogenetic analysis would have been helpful here. A set of skull reconstructions would have made things clear. Both are lacking from the new Martin-Silverstone study.

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 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.
Kellner AWA 2010. Comments on the Pteranodontidae (Pterosauria, Pterodactyloidea)
with the description of two new species. Anais da Academia Brasileira de Ciências 82(4): 1063-1084.
Martin-Silverstone E, Glaser JRN, Acorn JH, Mohr S and Currie PJ 2017. Reassesment of Dawndraco kanzai Kellner, 2010 and reassignment of the type specimen to Pteranodon sternbergi Harksen, 1966.  Vertebrate Anatomy Morphology Palaeontology 3:47-59.
Marsh OC 1876a. Notice of a new sub-order of Pterosauria. American Journal of Science, Series 3, 11:507-509.
Miller HW 1971. A skull of Pteranodon (Longicepia) longiceps Marsh associated with wing and body parts. Kansas Academy of Science, Transactions 74(10):20-33.

wiki/Pteranodon

Pterodactyls Alive! 1984 BBC video with David Attenborough on YouTube

Pterodactyls Alive! on YouTube click here

It’s more than 30 years old.
It’s not HD. It still supports the notion of a inverted hanging pterosaur. It precedes the discovery of eggs. Even so, it features a gliding Pteranodon model, lots of great sea bird scenes (including diving pelicans, soaring, dipping frigate birds and skimming skimmers) along with several great bat scenes (including grounded bats taking off).

And Mr. Attenborough was just starting to get gray hair back then.  :  )

The video also stars a bipedal animated Dimorphodon following the then recent release of Padian’s early work on that pterosaur.

 

Fun cardboard Pteranodon costume and model

Figure 1. Pteranodon costume with wings that fold in the plane of the wing.

Figure 1. Pteranodon costume with wings that fold in the plane of the wing with a distal membrane that extends to the anterior femur! Or is this a giant woodpecker team mascot?

Lisa Glover is a very creative person, doing great things with cardboard. Although this could be a woodpecker, Glover promotes it as a Pteranodon. More pix on her website where she writes, “what started as a homework assignment, quickly became a time machine to the cretaceous period. glover originally created a walking, wearable velociraptor and has now progressed to something a bit more challenging.”

Figure 2. Smaller more complete cardboard model of Pteranodon.

Figure 2. Smaller more complete cardboard model of Pteranodon. I like the widespread hind limbs and narrow chord wing membrane no deeper than the knee. 

On a similar note
If you’re in the mood for a cardboard cut out model, a few years earlier I offered this version of Pteranodon which you can download as a pdf then print on cover stock, cut out, fold, glue and hang from a string.

Build Your Own Pteranodon Paper Model

Click to download pdf. Build Your Own Pteranodon model on 8.5×11 inch paper.

The Tanking-Davis Pteranodon (private collection)

Americo Michael Minotti
recently let me know of a new Pteranodon skull in his collection that he considered the largest Pteranodon skull yet discovered (Figs. 1-3). Discovered by fossil collector David Tanking, within the Niobrara formation at Gove Co. Kansas and was added to the private collection of  Stanley Davis. Mr. Davis introduced the skull to Dr. Bennett, Associate professor at Fort Hays State University, who did an onsite detailed analysis of it. According to the reports of Davis and Tanking, Dr. Bennett stated that the skull had significant scientific value and was the largest and most complete sample of the species he had ever seen.

Sometime after the senescence of this animal, both cranial and mandibular sections of the skull may have sunk together in a nose-down position causing the rostrum to be splintered on impact with the sea floor and the tip of the mandible lost. The post-crania, a primary food source, was likely scattered by scavengers, etc.

Figure 1. Tanking-Davis Pteranodon in situ and with parts traced out and restored. The rostrum indicates a probable shorter rostrum, but the mandible does not taper as quickly as would be expected for the relatively short rostrum in a conventional linear digression model. It is conceivable that the distal maxillary and mandibular beaks may have therefore taken a less conventional configuration in this specimen. Click to enlarge. Photos courtesy of A. M. Minotti.

Figure 1. Tanking-Davis Pteranodon in situ and with parts traced out and restored. The rostrum indicates a probable shorter rostrum, but the mandible does not taper as quickly as would be expected for the relatively short rostrum in a conventional linear digression model. It is conceivable that the distal maxillary and mandibular beaks may have therefore taken a less conventional configuration in this specimen. Crest and ventral mandible are restored. The two gray maxillary triangles were retrieved from deeper matrix. Click to enlarge. Photos courtesy of A. M. Minotti.

The rostrum indicates a possible shorter rostrum, but the mandible does not taper as quickly as would be required for that short rostrum. In Pteranodon the rostrum extends beyond the mandible. In Nyctosaurus just the opposite.

Figure 2. The Tanking-Davis specimen compared to other forms. Specimen w and specimen z appear to be the closest to the Tanking-David specimen. Specimen 'w' = Pteranodon sternbergi? USNM 12167 (undescribed). Specimen 'z' = Pteranodon longiceps? Dawndraco? UALVP 24238. Click to enlarge.

Figure 2. The Tanking-Davis specimen compared to other forms. Specimen w and specimen z appear to be the closest to the Tanking-David specimen. Specimen ‘w’ = Pteranodon sternbergi? USNM 12167 (undescribed). Specimen ‘z’ = Pteranodon longiceps? Dawndraco? UALVP 24238. The Tanking-Davis specimen may not have been the largest Pteranodon of all, but it currently represents the longest skull of preserved elements. Click to enlarge

The reconstructed specimen (Fig. 3) appears to be largely complete, lacking only the crest and the rostral tooth (an empty space remains at the root) and the mandible tip.

Figure 1. Tanking-Davis Pteranodon reconstructed and with bones identified.

Figure 3. Tanking-Davis Pteranodon reconstructed and with bones identified. Click to enlarge.

The size of the skull, the type of taper in the rostrum, the small size of the orbit, and the placement of the anterior most portion of the crest just superior to the orbit all suggest that this specimen is most closely related to the UALVP 24238 specimen, but distinct. Note that in the UALVP 24238 specimen (Fig. 3, specimen z) the orbit is slightly higher relative to the dorsal crest margin and the Tanking-David specimen locates the orbit below this margin. The orbit is also slightly larger in the Tanking-Davis specimen. The  USNM 12167 specimen (specimen w) is also similar.

Figure 3. The UALVP specimen of Pteranodon. Note the lack of taper in the rostrum along with the small size of the orbit.

Figure 4. The UALVP specimen of Pteranodon. Note the lack of taper in the rostrum along with the small size of the orbit. UALVP 24238 (Dawndraco kanzai according to Kellner 2010)

 

With regard to females vs. males,
I still don’t see any male/female pairings here, and certainly none associated with pelves. The deep pelvis formerly attributed to a female Pteranodon actually belongs to a rare giant Nyctosaurus, as discussed here earlier.

Missing anterior tooth?
We looked at that subject earlier here.

References
Kellner AWA 2010. Comments on the Pteranodontidae (Pterosauria, Pterodactyloidea) with the description of two new species. Anais da Academia Brasileira de Ciências 82(4): 1063-1084.

wiki/Pteranodon