Germanodactylus sp. 6592 plate and counterplate

Matching a fossil plate to a counterplate is easy.
Matching a photo of a plate to a photo of counterplate (Fig. 1) requires Photoshop, even if the differences are minute.

Figure 1. Plate and counterplate of the SMNS 6592 specimen referred to Germanodactylus matched in Photoshop.

Figure 1. Plate and counterplate of the SMNS 6592 specimen referred to Germanodactylus matched in Photoshop.

This is the first time I’ve seen
the counterplate to the SMNS 6592 specimen attributed to Germanodactylus. And I think this counterplate is composed of painted plaster. Photoshop was used to match the plate to the counterplate and to trace the resulting elements. As you can see, the pelvis is in an atypical position due to taphonomy (crash landing on its butt?), but everything else seems to be naturally posed with the exception of the displaced and overlapping femora (another results of the crash landing, perhaps).

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.

New name and a name resurrection for two Solnhofen pterosaurs

Vidovic and Martill 2017
propose new and resurrect old generic names for two Solnhofen pterosaur specimens. Both are good and needed based on an earlier abstract (Peters 2007) and tree topology published here six years ago at ReptileEvolution.com in the large pterosaur tree (LPT, 232 taxa).

Unfortunately
Vidovic and Martill remain completely in the dark regarding pterosaur ontogeny. As we learned earlier here, here and here from several adult and juvenile specimens, pterosaurs juveniles and embryos had adult proportions and that’s why they were mechanically able to fly shortly after hatching. Vidovic and Martill report, “It is difficult to distinguish ‘G. rhamphastinus’ (Fig. 3 from the holotype of D. kochi (Fig. 2) other than by using size-related criteria.” And, “juvenile pterosaurs with small crests have been identified.”

Also unfortunately,
Vidovic and Martill still consider pterosaurs to be derived archosaurs or archosauriforms. They report, “A cladistic analysis of the Pterosauria, including all the taxa discussed here, was performed. The analysis included 104 operational taxonomic units (OTUs) comprising 99 pterosaurs and five archosauriforms as an outgroup.” We have to ask ourselves, how long will pterosaur workers remain in the dark on these basic questions that were answered years ago? Look here, here (Peters 2000, 2007) and here.

Pterodactylus wastebasket
Vidovic and Martill write: “Until relatively recently, the genus Pterodactylus Cuvier, 1809 had been a wastebasket taxon that has included many diverse pterosaurs, including some that are now recognized as basal nonpterodactyloids.” We looked at the Pterodactylus wastebasket here in 2011 (Fig. `1).

The Pterodactylus lineage and mislabeled specimens formerly attributed to this "wastebasket" genus

Figure 1. Click to enlarge. The Pterodactylus lineage and mislabeled specimens formerly attributed to this “wastebasket” genus

Wellnhofer 1970
provided catalog numbers for dozens of Solnhofen specimens. Since those numbers are simpler than their museum numbers that’s how they are named (Figs. 2, 3) at ReptileEvolution.com.

basal germanodactylids

Figure 2. Basal Germanodactylia, Three taxa preceding Germanodactylus rhamphastinus: No. 6, No. 12 and No. 23, the last renamed Diopecephalus kochi. These are all adults.

No. 23 — BSP AS XIX 3 — Diopecephalus kochi (formerly Pterodactylus kochi).
(Fig. 1, left). Seeley had it right originally. Vidovic and Martill correct a century of error when they report, “The holotype of ‘P. kochi’ was considered to belong to a distinct genus by Seeley (1871), which he  unambiguously named Diopecephalus Seeley, 1871.”

No. 64 — B St AS I 745  —
Altmuehlopterus (formerly Germanodactylus) rhamphastinus

Vidovic and Martill reported, “Many phylogenetic studies demonstrate that the two species of Germanodactylus nest together (Kellner 2003; Unwin 2003; Andres & Ji 2008; Lu et al. 2009; Wang et al. 2009; Andres et al. 2014) in a monophyletic clade, but a more focussed analysis by Maisch et al. (2004) demonstrates the genus to be paraphyletic. Maisch et al. (2004) created the nomen nudum Daitingopterus, intended for the reception of ‘G. rhamphastinus’ by placing the name in a table with no specific reference to a specimen.”

Figure 3. Germanodactylus rhamphastinus, No. 64 in the Wellnhofer 1970 catalog.

Figure 3. Germanodactylus rhamphastinus, No. 64 in the Wellnhofer 1970 catalog. Vidovic and Martill renamed this specimen Altmuehlopterus, which is fine and appropriate.

The LPT separates A. (G.) rhamphastinus from G. cristatus by two taxa.

Problems with the Vidovic and Martill 2017 tree:

  1. Lagerpeton nests with Marasuchus, both as proximal outgroups to the Pterosauria. Totally bogus. Tested, validated, real outgroups are listed here. The Fenestrasauria (Peters 2000) is overlooked in the text and references.
  2. Preondactylus and Austriadactylus nest as basalmost pterosaurs. Bergamodactylus, the basalmost pterosaur in the LPT, is excluded.
  3. Only one specimen each of Dorygnathus and Scaphognathus are employed. The LPT shows two clades of pterodactyloid-grade pterosaurs arise from various specimens of Dorygnathus while two others arise from tiny Scaphognathus specimens experiencing phylogenetic miniaturization.
  4. As a result (perhaps) toothy ornithocheirids nest with toothless pteranodontids. In the LPT ornithocheirids arise from equally tooth cycnorhyamphids while shartp-face pteranodontids arise from similar germanodactylids.
  5. The Darwinopterus clade nests as the proximal outgroup to the traditional Pterodactyloidea, when the LPT shows it to be a sterile clade with some pterodactyloid-grade traits.
  6. Altmuehlopterus (formerly Germanodactylus) rhamphastinus nests with G. cristatus
  7. Diopecephalus kochi nests with Pterodactylus antiquus.
  8. Those are the big problems. There are more, but I want to keep it pertinent.

Vidovic and Martill provide clues to their observational problems
when they note, “The genera Pterodactylus and Diopecephalus are remarkably similar.” No they aren’t! Species within the Pterodactylus clade are not even that similar!

Re: Germanodactylus and Pterodactylus,
Vidovic and Martill write: “We agree that some of the differences could be ontogenetically variable and perhaps vary between sexes, so in 1996 it seemed possible that the two species could be at least congeneric.” They disagree with the “common opinion” that the two are distinct genera. Let’s go with the evidence of a large gamut phylogenetic analysis — not opinion — or any analysis lacking so many pertinent taxa.

Vidovic and Martill 2017 rename G. rhamphastinus
Altmuehlopterus rhamphastinus. That’s good. It is generically distinct from its proximal relatives in the LPT. They report, “This name is presented as an alternative to the geographically significant name Daitingopterus (Maisch et al., 2004) which is a nomen nudum.” Not sure how all that falls. I’ll leave such issues to the PhDs.

If you like long nomenclature puzzles
you’ll like Vidovic and Martill 2017. They do a good job of running down all the names that prior workers gave to these century-old specimens. Beware that they are clueless as to the origin of pterosaurs, the ontogeny of pterosaurs and previous work on the phylogeny of pterosaurs based on a much larger taxon list of ingroup and outgroup taxa.

References
Peters D 2000. A redescription of four prolacertiform genera and implications for pterosaur phylogenesis. Rivista Italiana di Paleontologia e Stratigrafia 106: 293-336
Peters D 2007  The origin and radiation of the Pterosauria. Flugsaurier. The Wellnhofer Pterosaur Meeting, Munich 27. Abstract online here.
Vidovic SU and Martill DM 2017. The taxonomy and phylogeny of Diopecephalus kochi (Wagner, 1837) and ‘Germanodactylus rhamphastinus’ (Wagner, 1851). From: Hone DWE., Witton MP and Martill DM (eds) New Perspectives on Pterosaur Palaeobiology. Geological Society, London, Special Publications, 455, https://doi.org/10.1144/SP455.12
Wellnhofer P 1970. Die Pterodactyloidea (Pterosauria) der Oberjura-Plattenkalke Süddeutschlands. Abhandlungen der Bayerischen Akademie der Wissenschaften, N.F., Munich 141: 1-133.

wiki/Pterodactylus
wiki/Germanodactylus

Germanodactylus cristatus revisited

Figure 1. The Germanodactylus cristatus plate containing a scattered skeleton.

Figure 1. The Germanodactylus cristatus plate containing a scattered skeleton. Click to enlarge.

Germanodactylus cristatus (B St 1892 IV 1) is best known from a plate of scattered fossil bones (Fig. 1), but there is also a counterplate that adds data. Today a combination of the plate and counter plate skull material (Fig. 2) refines what we know of G. cristatus, a taxon basal to Dsungaripterus, Sinopterus and Shenzhoupterus and their kin – AND ALSO – not far from the last common ancestor of Elanodactylus, Wenupteryx, Pteranodon and Nyctosaurus.

Short history
B St 1892 IV 1
 (Pterodactylus kochi Plieninger 1901, Germanodactylus cristatus Wiman 1925, No. 61 of Wellnhofer 1970) was originally considered a Pterodactylus. Bennett (1996) reported that specimens of Pterodactylus were juvenile Germanodactylus. That hypothesis was rejected by all workers including Bennett himself. Bennett (2006) reported on two tiny Solnhofen pterosaurs which he considered juveniles of Germanodactylus. Here these two tiny pterosaurs, No. 9 and No. 31 (from the Wellnhofer 1970 catalog), nest separately, as tiny adults closer to Scaphognathus.

Figure 2. GIF animation of skull elements (plate and counter plate) with DGS tracings. See figure 3 for reconstruction. Scenes change every 5 seconds. Note the rostral and mandible tip teeth, homologous with those in toothless pterosaurs related to this taxon.

Figure 2. GIF animation of skull elements (plate and counter plate) with DGS tracings. See figure 3 for reconstruction. Scenes change every 5 seconds. Note the rostral and mandible tip teeth, homologous with those in toothless pterosaurs related to this taxon. Nine frames then repeat.

This B St 1892 IV 1 specimen is fairly easy to trace,
but the palatal and occipital elements are often ignored. Some breakage requires repairs. Some soft tissue is preserved, including the location of the naris within the antorbital fenestra (small gray dot). But also look for a tiny secondary narial opening just anterior to the antorbital fenestra between the nasal and jugal. We looked at the origin of the secondary naris here, at the Scaphognathus node.

Figure 3. Reconstructiion of Germanodactylus cristatus skull, including palatal and occipital elements, plus a Y-shaped hyoid (tongue bone). The maxilla has large palatal plates. The vomers are gracile and elongated. The pterygoids are in pink They were split in situ with maxillary processes separate from posteromedial portion. Ectopterygoid + palatine = ectopalatine, slender elements ventral here to pterygoids, but actually dorsal in vivo.

Figure 3. Reconstructiion of Germanodactylus cristatus skull, including palatal and occipital elements, plus a Y-shaped hyoid (tongue bone). The maxilla has large palatal plates. The vomers are gracile and elongated. The pterygoids are in pink They were split in situ with maxillary processes separate from posteromedial portion. Ectopterygoid + palatine = ectopalatine, slender elements ventral here to pterygoids, but actually dorsal in vivo.

The standing image
of G. cristatus at reptileevolution.com has also been revised (Fig. 4). Earlier the maxilla was portrayed as preserved on one side, with a concave ventral margin, but the other side has a straight ventral margin. When matched to the mandible, the straight ventral margin is a closer match, which gives the dorsal rostral margin a concave curve when revised in vivo.

Figure 4. Germanodactylus cristatus with newly revised skull.

Figure 4. Germanodactylus cristatus with newly revised skull including the palate. Note the very thin distal wing elements.

References
Bennett SC 1996. Year-classes of pterosaurs from the Solnhofen Limestone of Germany: taxonomic and systematic implications. Journal of Vertebrate Paleontology 16 (2): 432–444. doi:10.1080/02724634.1996.10011332.
Bennett SC 2006. Juvenile specimens of the pterosaur Germanodactylus cristatus, with a review of the genus. Journal of Vertebrate Paleontology 26:872–878.
Plieninger F 1901. Beiträge zur Kenntnis der FlugsaurierPaläontographica 48, 65–90 and pls 4–5.
Wellnhofer P 1970. Die Pterodactyloidea (Pterosauria) der Oberjura-Plattenkalke Süddeutschlands. Abhandlungen der Bayerischen Akademie der Wissenschaften, N.F., Munich 141: 1-133.
Wiman C 1925. Aus dem Leber der Flugsaurier. Bulletin of the Geological Insititute of the University of Uppsala 19: 115-127

wiki/Germanodactylus

 

Germanodactylus cristatus ventral reconstruction

Every so often
old reconstructions get updated. This time Germanodactylus cristatus gets the treatment with a new ventral view, matching the exposure of the cervicals and anterior dorsals (Fig. 1).

Figure 1. Germanodactylus cristatus in ventral view, wings outspread.

Figure 1. Germanodactylus cristatus in ventral view, wings and legs outspread. in that configuration the legs act as horizontal stabilizers and create their own lift. The torso appears to have been wider than deep. The prepubes were fused medially. The pelvis and three toes are conjectural based on matrix impressions, perhaps buried bones.

Note
the femoral heads are nearly at right angles to the femoral shafts. This gives the hind limbs more of an erect configuration, which may have aided this taxon in terrestrial stalking, something we talked about earlier here.

Figure 2. Germanodactylus cristatus in lateral view, bipedal/quadrupedal configuration.

Figure 2. Germanodactylus cristatus in lateral view, bipedal/quadrupedal configuration. The pelvis and three toes are based on matrix impressions.

References
Plieninger F 1901. Beiträge zur Kenntnis der FlugsaurierPaläontographica 48, 65–90 and pls 4–5.
Wellnhofer P 1970. Die Pterodactyloidea (Pterosauria) der Oberjura-Plattenkalke Süddeutschlands. Abhandlungen der Bayerischen Akademie der Wissenschaften, N.F., Munich 141: 1-133.
Wiman C 1925. Aus dem Leber der Flugsaurier. Bulletin of the Geological Insititute of the University of Uppsala 19: 115-127.

wiki/Germanodactylus

At last! Two pterosaur specimens that may be conspecific!

Phylogenetic lumpers
like to consider several closely related specimens conspecific and/or congeneric.

Phylogenetic splitters
do the opposite. Rarely are specimens conspecific and/or congeneric in their eyes.

In my studies
of pterosaurs I have never seen two specimens collected at different times and in different places that were conspecific. So I guess that makes me a splitter, but I have to work within the confines of traditional naming paradigms.

In Witton et al. 2015,
I believe I have found a germanodactylid (NMING F15005) pictured in Witton et al. 2015, that may be conspecific with a previously discovered germanodactylid (SMNK PAL 64592, Fig. 1), based on comparative skull anatomy only.

Figure 1. Two germanodactylids that may be conspecific, the more complete SMNK PAL 6592 specimen and the skull only NMIING-F15005 specimen. These two score almost identically in the large pterosaur tree. Can you see the subtle differences?

Figure 1. Two germanodactylids that may be conspecific, the more complete SMNK PAL 6592 specimen and the skull only NMIING-F15005 specimen. These two score almost identically in the large pterosaur tree. Can you see the subtle differences?

The two skulls are almost identical… almost.
The question is… are the differences enough to create a new species? Or not?

So what are the differences in the NMING specimen? 

  1. The anterior dentary is deeper
  2. More teeth are present in the dentary
  3. The antorbital fenestra is longer (further anteriorly)
  4. The anterior jugal is taller and more gracile
  5. The lower antorbital fenestra bar (mx + ju) is more gracile
  6. The quadrate leans further posteriorly
  7. The postorbital appears to be more gracile and lower on the cranium
  8. The base of the orbit is not pointed, but flat creating a larger orbit area

So, are these two specimens conspecific?
Depends if you’re a lumper or a splitter. Do you judge these trait differences as individual variation? or just a little beyond that?

Figure 2. the NMING F15005 specimen with arrows pointing to the upper and lower procumbent teeth that characterize this clade.

Figure 2. the NMING F15005 specimen with arrows pointing to the upper and lower procumbent single teeth that characterize this clade. These teeth are also found in all pteranodontids, dsungaripterids and tapejarids. The blue part of the mandible is also part of the dentary.

I hope you’ll also take note
that at first glance these two skulls appear identical. Putting them side-by-side helps bring out the differences. Always try to make your comparative presentations side by side.

References
Witton MP, O’Sullivan M and Martill DM 2015. The relationships of Cuspicephalus scarfi Martill and Etches, 2013 and Normannognathus wellnhoferi Buffetaut et al., 1998 to other monofenestratan pterosaurs.

Wenupteryx – a newly named South American germanodactylid

MOZ-3625P

Figure 1. Reconstruction of Wenupteryx based on in situ figure from Codorniu et al 2006, simply moving bone images to restore the standing pterosaur using DGS. They were unable to identify this specimen, but the large pterosaur tree nested it precisely with the germanodactylids.

Wenupteryx uzi is the new name (Codorniú and Gasparini 2007) given to the second Jurassic pterosaur discovered in Argentina (Codorniu et al. 2006, Figs 1-4). Formerly it was known from its specimen number, MOZ 3625. Nearly a complete skeleton, sans the skull, is known from scattered and crushed remains. A recent closeup of the lower hind leg  (Codorniú et al. 2013) permitted a DGS reconstruction of the pes (Fig. 2) in line with its previous phylogenetic nesting within the germanodactylids, between the BMM private specimen and all higher germanodactylids, seen here (Fig. 3).

Codorniú et al. 2013) considered this a tibiatarsus and the loose tarsals distal tarsals. They did not realize there are no tibiotarsi known among pterosaurs. The loose tarsals here (Fig. 2) are proximal tarsals (astragalus and calcaneum) plus one distal tarsal (likely #4).

Figure 2. Colorized bones of the tibia, tarsus and pes of Wenupteryx (from Codorniu and Garrido 2013) permitted identification of the elements. Not a tibiotarsus, but a fused tibia/fibula and proximal tarsals plus distal tarsal 4 are identified here.

Figure 2. Colorized bones of the tibia, tarsus and pes of Wenupteryx (from Codorniu and Gasparini 2013) permitted identification of the elements. Not a tibiotarsus, but a fused tibia/fibula and proximal tarsals plus distal tarsal 4 are identified here. Gray and outline areas are restored.

Germanodactylus and kin

Figure 3. Click to enlarge. Germanodactylus and kin. Wunepteryx (MOZ 3625) is third from right.

Wenupteryx is an important taxon that nests at the base of two higher Germanodactylus clades. One branch leads to dsungaripterids, shenzhoupterids and tapejarids. Another branch leads to elanodactylids, eopteranodontids and pteranodontids. So, it is a very plesiomorphic specimen. No wonder it was hard to originally identify (Codorniu et al. 2006).

Figure 4. Wenupteryx in dorsal view as figured by Codorniu et al. 2006

Figure 4. Wenupteryx in dorsal view as figured by Codorniu et al. 2006. The hypothetical skull in shape and size is a very good guess based on phylogenetic bracketing. Note the nearly right angle femoral heads. This germanodactyloid pterosaur was therefore likely a good runner.

The outgroup taxon, the BMM private specimen sometimes mistakenly considered a large, robust Pterodactylus, is likewise plesiomorphic.

Someday…
We are going to have to take a serious look at all specimens given the genus Germanodactylus and all of the specimens, like Wenupteryx, nesting between them but given other generic names (Fig. 3).

References
Codorniú L, Gasparini Z and Paulina-Carabajal A 2006. A late Jurassic pterosaur (Reptilia, Pterodactyloidea) from northwestern Patagonia, Argentina. Journal of South American Earth Sciences 20:283-389.
Codorniú L and Gasparini Z 2007. Pterosauria. Pp. 143-166 in Patagonian Mesozoic Reptiles Gasparini, Salgado and Coria editors. Indiana University Press.
Codorniú L and Gasparini Z 2013. The Late Jurassic pterosaurs from northern Patagonia, Argentina. Earth and Environmental Science Transactions of the Royal Society of Edinburgh 103:1-10.

Herbstosaurus – What is it?

Herbstosaurus pigmaeus CTES-PZ-1711 (late Jurassic) was originally described as a small dinosaur (Casamiquela 1975). All that is known of this specimen is a pelvis, limb and prepubis (ahhh, there’s a clue!). Ostrom (1978) more accurately identified Herbstosaurus as a pterosaur.

Figure 1. Herbstosaurus on the left compared to Coloborhynchus on the right. The sacrum is similar, but more swept back in Herbstosaurus. The femoral head is disssimilar.

Figure 1. Herbstosaurus on the left compared to the similarly-sized Coloborhynchus on the right to the same scale. The sacrum is similar, but the transverse processes are more swept back in Herbstosaurus. The anterior extent of the ilium and the femoral head/neck/shaft angle are disssimilar. Ornithocheirids are not known for a large prepubis. So, no match here.

But what kind?
Galton 1981 considered Herbstosaurus a member of the “Pterodactyloidea.”  Carroll 1988 narrowed that down to the Pterodactylidae. Wellnhofer 1991 suggested it was a more basal pterosaur, in view of the form of the pelvis. In Unwin 1996 considered it a basal member of the Dsungaripteroidea.

So, with so little consensus, what is Herbstosaurus?

Unfortunately,  
there is not enough here to run Herbstosaurus through phylogenetic analysis using the large pterosaur tree. So Let’s see where trait analysis leads us.

Ilial processes obliquely angled:
Most derived pterosaurs have this except basal pterosaurs up to MCSNB 8950.

Five+ swept back sacrals not coosified:
Most basal pterosaurs have five sacrals not coossified. This does not mean they are juveniles, unless the fossil record is seriously skewed. There are several clades of chiefly larger pterosaurs that coossify the sacrals. Other large forms don’t. Apparently only germanodactylids and their many descendants have swept back sacral transverse processes (Fig. 2) and basal forms, like Germanodactylus rhamphastinus and kin, do not coossify the sacrals.

Figure 2. Swept back sacrals and right angle femoral heads on Germanodactylus rhamphasitinus. Herbstosaurus was much larger overall.

Figure 2. Swept back sacrals and right angle femoral heads on Germanodactylus rhamphastinus. Herbstosaurus was much larger overall. Many bones here are difficult to gauge. The counterplate was also referenced.

Figure 3. The BMM Germanodactylus with an ilium, femur and prepubis shaped like Herbstosaurus, but much smaller.

Figure 3. The BMM Germanodactylus with an ilium, femur and prepubis shaped like Herbstosaurus, but overall the BMM specimen is much smaller.

Prepubis fan shape perhaps without perforation:
The fan shape appears in several clades The perforation is seen in most pterosaurs, but absent in an unassociated few. Germanodactylus often has a fan-shaped prepubis.

Femur with right angle head and neck, but head axis not aligned with neck axis.
This is a tough one. I wondered, is there some breakage that would, if repaired, align the femoral head axis with the femoral neck axis in Herbstosaurus? I’ve only seen drawings (Fig. 1). In most pterosaurs the axis of the head of the femur is very closely aligned with the axis of the neck of the femur. Not so in Herbstosaurus, hence the concern.

Size:
Substantial. No tiny pterosaurs.

Chronology:
Late Jurassic

Summary:
From the available evidence, Unwin 1996 was the most correct. Herbstosaurus is likely a large and primitive Germanodactylus-type pterosaur. Large taxa are already known from that clade. They are the elanodactylids (Fig. 4). Unfortunately what is known from the partial skeletons of those specimens does not overlap with the partial skeleton of Herbstosaurus. In any case, at present with these clues, this seems to be a “best match.”

Germanodactylus and kin

Figure 4. Click to enlarge. Germanodactylus and kin.

References
Casamiquela RM 1975. Herbstosaurus pigmaeus (Coeluria, Compsognathidae) n. gen. n. sp. del Jurásic medio del Neuquén (Patagonia septentrional). Uno de los más pequeños dinosaurios conocidos [Herbstosaurus pigmaeus (Coeluria, Compsognathidae) n. gen. n. sp. from the Middle Jurassic of Neuquén (northern Patagonia). One of the smallest known dinosaurs]. Actas del Primer Congreso Argentino de Paleontologia y Bioestratigrafia, Tucumán 2:87-103.
Galton PM 1981. A rhamphorhynchoid pterosaur from the Upper Jurassic of North America. Journal of Paleontology 55(5):1117-1122.
Ostrom JH 1978. “The osteology of Compsognathus longipes Wagner”, Zitteliana4: 73-118.
Unwin DM 2003. On the phylogeny and evolutionary history of pterosaurs. In E. Buffetaut, J.-M. Mazin (eds.), Evolution and Palaeobiology of Pterosaurs, Geological Society of London, Special Publications 217:139-190

A Big Germanodactylus in the Late Cretaceous Niobrara of Kansas

Germanodactylus is chiefly known from Solnhofen formation (Late Jurassic, Germany), but there’s one Germanodactylus that was found in the Niobrara formation of Kansas in the Late Cretaceous. And it was a big one (Fig. 1)!

No. 13 and its sisters

Figure 2. No. 13 and its sisters, Germanodactylus, Muzquizopteryx and Eopteranodon. Click to Enlarge. Note the resemblance between YPM 1179 and SMNK PAL 6592. The other taxa shown here do not come as close, but have evolved distinct traits.

Okay, I may have been a little misleading.
The Niobrara specimen has been formally named, Pteranodon occidentalis YPM 1179 (Marsh 1876a).  The skull was twice the length (and therefore 8x the size of is nearest Germanodactylus sister, SMNK PAL 6592. But in the panoply of Pteranodon skulls (Fig. 2K), YPM 1179 had the shortest rostrum and ranks among the smallest otherwise. We don’t know the post-crania of YPM 1179. Did extreme metacarpals and shorter hind limbs evolve coincidentally with extreme rostral lengths? No one knows (but see below for clues). Those traits are the basic post-cranial attributes of Pteranodon that separate it from Germanodactylus (other than size). If YPM 1179 had longer legs and shorter metacarpals than a typical Pteranodon, then there would be even more argument to calling it a big Germanodactylus.

Pteranodon skulls

Figure 2. Click to enlarge. A family tree of Pteranodon. There’s YMP 1179 (K) close to the SMNK PAL 6592 Germanodactylus (B).

Did you know…
There’s more morphological variation among the various specimens of Germanodactylus (Fig. 3) than there is between SMNK PAL 6592 and  YPM1179?  Someday someone will sort out the real Germanodactylus from the non-Germanodactylus. It’s nearly as bad of a situation as the Pterodactylus sorting problem, reported on earlier here. And someone, hopefully, will confirm the separation of the eopteranodontids from the very similar azhdarchids seen here in the large pterosaur family tree. Experts are still confused about that one.

Germanodactylus and kin

Figure 3. Click to enlarge. Germanodactylus and kin. There’s YPM 1179 on the right.

So, where do you draw the line?
Morphology? Size? Location? Tradition? They all have impact factors here in lumping or splitting these two sister taxa. YPM 1179 will probably remain a Pteranodon due to its separation in time and location. But now you and I both know, YPM 1179 is closer to Germanodactylus and forms a great transitional taxon to the Pteranodon clade (Fig. 1).

Possible Post-Crania?
Mated with SMU 76476, the “oldest Pteranodon” (Myers 2010) we may have a clue as to the post-crania of YPM 1179 (Fig. 4). The humerus is relatively shorter in the SMU specimen. We don’t know how long the metacarpus and m4.1 were due to end breaks, but the metacarpus was at least slightly longer, as in Muzquizopteryx and Eopteranodon (Fig. 1).

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.

The Deltopectoral Crest Warp
Pteranodon has a deltopectoral crest warp and thicker anteriorly. Myers (2010) reports, “The anterior edge of the terminal expansion of the deltopectoral crest is flat, and its long axis is oriented at an angle to the base of the crest, creating the distinctive warped appearance described by Padian (1984) and Bennett (1989).” Actually the warp in SMU 76476 is not as apparent and there is no thickness in the depth of the anterior deltopectoral crest found in other Pteranodon specimens. So, this humerus has a thin deltopectoral crest more like that of Germanodactylus.

References
Marsh OC 1876a. Notice of a new sub-order of Pterosauria. American Journal of Science, Series 3, 11:507-509.
Myers TS 2010. Earliest occurrence of the Pteranodontidae (Archosauria: Pterosauria) in North America: new material from the Austin Group of Texas. Journal of Paleontology 84(6): 1071-1081. PDF online

 

Is Kepodactylus a Germanodactylid?

Kepodactylus insperatus (Harris and Carpenter 1996, DMNH 21684) was described as a large pterodactyloid. Only a few pieces are known (Fig. 1, left). The humerus (here scaled to half the published scale bar) most closely resembles that of Germanodactylus cristatus.

Kepodactylus

Figure 1. Kepodactylus (left) compared to Germanodactylus cristatus (greyed out, right). The humerus is a close match. The cervical is larger. What Harris and Carpenter (1996) described as a possible distal wing phalanx is much too large. I’m wondering if it is a distal radius?

Kepodactylus gives us not much to go on, but the best match I was able to find is with Germanodactylus cristatus. If so, this continues the matches seen in Late Jurassic Solnhofen pterosaur fauna now starting to add up in Late Jurassic North America. Perhaps from such as Kepodactylus we would eventually get Eopteranodon, Pteranodon and Nyctosaurus.

As always, I encourage readers to see specimens, make observations and come to your own conclusions. Test. Test. And test again.

Evidence and support in the form of nexus, pdf and jpeg files will be sent to all who request additional data.

References
Harris JD and CarpenterK 1996. A large pterodactyloid from the Morrison Formation (Late Jurassic) of Garden Park, Colorado. Neues Jahrbuch für Geologie und Paläontologie Monatshefte 1996(8):473-484.

wiki/Kepodactylus