Anhanguera animation at the NHM (London)

This one started off with so much promise
as the animators at the National History Museum (NHM) in London assembled their version of the ornithocheirid pterosaur, Anhanguera, bipedally (Fig. 1), as you’ll see when you click on the video under ‘References’.

Figure 1. Animated by the NHM, Anhanguera is bipedal and flapping its literally oversize wings.

Figure 1. Animated by the NHM, Anhanguera is bipedal and flapping its literally oversize wings standing on oversize feet with an undersized skull and hyperextended elbows and unbalanced stance.

Unfortunately there were some morphology issues (compared in Fig. 2):

  1. wings too long
  2. sternal complex missing
  3. gastralia missing (but rarely preserved in ornithocheirids)
  4. feet way too big
  5. skull too small
  6. tail too short
  7. not sprawling
  8. free fingers too big
  9. wing fingers should tucked tight against elbows (in the same plane)
  10. one extra cervical
  11. anterbrachia too short and gracile
  12. elbows overextended (in Fig. 1)
  13. too much weight put on forelimbs, center of balance (wing root) should be over the toes
  14. Prepubes are extremely rare in ornithocheirds, but when present they are tiny, putter-shaped and oriented ventrally in line with the bent femora, not anteriorly
Figure 2. NHM Anhanguera compared to skeletal image from ReptileEvolution.com.

Figure 2. NHM Anhanguera compared to skeletal image from ReptileEvolution.com. There are at least 10 inaccuracies here. See text for list.

Also unfortunately, the video quickly devolved
to the invalid and dangerous quad launch, when (doggone it!) it was all set up to do a more correct and  much safer bird-like launch. The laws of physics and biomechanics are ignored here, but at least David Attenborough narrates.

Figure 3. NHM Anhanguera quad launch select frames.

Figure 3. NHM Anhanguera quad launch select frames. The laws of physics and the limitations of biomechanics are ignored here.

Attempts to convince readers and workers
that the quad-launch hypothesis cheats morphology and physics (as recounted here and at links therein) have so far failed. But I’m not giving up. So, if anyone has a connection to the NHM in London, please make this post available to alert them of their accidental foray into wishful thinking and inaccurate morphology.

References
National History Museum (NHM) in London

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Azhdarcho restored (from bits and pieces)

Earlier
we looked at the neck and skull of Azhdarcho… Today we’ll put all the bits and pieces we know (from several individuals, unfortunately) to see what we get, following the Q. sp. bauplan (Fig. 1).

Figure 1. Azhdarcho to scale with more complete smaller Quetzalcoatlus specimen and in proportion to the bauplan of Q. sp. Note the robust femur and gracile humerus. These together with the small sternal complex and short distal wing elements indicate a flightless condition.

Figure 1. Azhdarcho to scale with more complete smaller Quetzalcoatlus specimen and in proportion to the bauplan of Q. sp. Note the robust femur and gracile humerus. These together with the small sternal complex and short distal wing elements indicate a flightless condition.

Azhdarcho lancicollis (Nesov 1984, Averianov 2010) is the namesake for the clade Azhdarchidae. This species is known from several individuals of various sizes and very few complete bones. That is why reconstructions of this genus are rare. This reconstruction is based on the more complete Q. sp., but about half as tall.

Given these limitations,
(no complete long bones), the femur appears to be more robust than in other azhdarchids, while the humerus is more gracile. Only in Huanhepterus is the femur so relatively short. The sternal complex is quite small, but with a deep cristospine, distinct from other azhdarchids. (Perhaps the rest of the sternal complex is missing.) Manual 4.4 was identified by Averianov, but it appears to be the distal portion of m4.3. The scale bars for the distal femur appear to be in error, or apply to a much larger individual (see Fig. 1).

The invisible aid in this reconstruction
is the observation that in nearly all post-Huanhepterus azhdarchids, the metacarpus, manual digit 4 and tibia are similar in length (Fig. 1), no matter how small or tall… probably to facilitate terrestrial locomotion.

Unfortunately,
not enough is known of Azhdarcho to add it to the LRT. So much has to be imagined.

References
Averianov AO 2010. The osteology of Azhdarcho lancicollis Nessov, 1984 (Pterosauria, Azhdarchidae) from the Late Cretaceous of Uzbekistan. Proceedings of the Zoological Institute of the Russian Academy of Sciences, 314(3): 246-317.
Nesov LA 1984. Upper Cretaceous pterosaurs and birds from Central Asia. Archived 17 March 2012 at the Wayback Machine. Paleontologicheskii Zhurnal, 1984(1), 47-57.

wiki/Azhdarcho

Battle of the giant Quetzalcoatlus sculptures

Bigger is better
especially when it comes to pterosaurs in museum exhibits. To wit: The Field Museum (Chicago, IL, USA) is installing a flying Quetzalcoatlus (Fig. 3) and a standing Quetzalcoatlus (largely imagined and restored, based on an almost complete wing bone, Fig. 1).

The artist/professors in Southern England
have one as well (Fig. 1). And full scale Q. northropi sculptures can be found worldwide! (See links and images at the end of this post.)

Figure 1. Field Museum Quetzalcoatlus (tan) vs. English Quetzalcoatlus (gray) vs. a tracing of the real Q. sp. and Q. northropi scant remains.

Figure 1. Field Museum Quetzalcoatlus (tan) vs. English Quetzalcoatlus (gray) vs. a tracing of the real Q. sp. and Q. northropi scant remains. Yes, the skull on the Field Musuem model is too long, evidently following the invalid archosaur hypothesis of origin. Pterosaurs, like other tritosaurs, do not change their proportions during growth as fossils demonstrate.

These models arrive on the heels
of a recent post on flightless giant pterosaurs. The Field Museum model (Fig. 1) appears to have a more precisely modeled skull, though about 50% too long (but really, without a skull, who knows?). Perhaps the skull was elongated (based on the smaller ?species) based on the invalid archosaur hypothesis of pterosaur origins. We know from the evidence of fossils that hatchlings and juvenile pterosaurs had adult proportions, not longer skulls.

Both models suffer
from putting too much weight on the tiny free fingers. The feet should be beneath the shoulder joint, as in birds, to take the weight off the tiny hyperextended fingers, acting more like ski poles, not providing thrust, only some sort of support.

The folded wing membrane should tend to disappear
(Fig. 2), but it shows in both models. The wing finger should flex closer to the elbow, but it doesn’t (probably to let the wing membrane show). The wing membrane chord should be shorter, to the elbow, but in both models the brachiopatagium blends with the leg, always awkwardly and in defiance of the data preserved in all pterosaur fossils (Fig. 2) that preserve soft tissue.

Here's how the wing membrane in pterosaurs virtually disappeared when folded.

Figure 2. Here’s how the wing membrane in pterosaurs virtually disappeared when folded. This is a tiny pre-azhdarchid, CM 11426.

Over at the Carnegie Museum (Pittsburgh, PA, USA)
they have a tiny pre-azhdarchid, CM 11426 (Fig. 2), with real wing membranes, as described above, matching those of other pterosaur soft tissues.

Figure 3. Field museum flying Quetzalcoatlus model has the invalid deep chord wing that attaches to the tibia.

Figure 3. Field museum flying Quetzalcoatlus model has the traditional but invalid deep chord wing that attaches to the tibia and makes this sort of pterosaur untenably awkward.

Figure 2. Quetzalcoatlus recreated as a digital model by Henderson 2010 compared to a bone reconstruction. No wonder the results were odd. The math was wrong.

Figure 4. Quetzalcoatlus recreated as a digital model by Henderson 2010 compared to a bone reconstruction. This is a possible slightly deeper wing chord. Compare this one to figure 5, which is more typical. And look at those hind limbs, like those of Sharovipteryx, forming a horizontal stabilizer, just like a typical airplane. And it matches the evidence (Fig. 2).

Not sure why pterosaur paleontologists
keep insisting that evidence (Fig. 2) can and should be ignored. It’s disheartening to see this and leaves them open to criticism.

Quetzalcoatlus running like a lizard prior to takeoff.

Figure 5. Quetzalcoatlus running like a lizard prior to takeoff. Click to animate. Giant azhdarchids gave up flying by reducing the lenth of their wings, following the patterns of other flightless pterosaurs. There is no awkwardness with this narrow chord wing design, which follows fossils like CM 11426 (Fig. 2).

Other online Quetzalcoatlus models/sculptures:

Low Poly Quetzalcoatlus model

https://www.alamy.com/stock-photo-muenchehagen-germany-10th-apr-2017-a-sculpture-of-the-quetzalcoatlus-138149396.html

https://blog.everythingdinosaur.co.uk/blog/_archives/2014/07/07/collecta-quetzalcoatlus-with-prey-model.html

If some of these seem to defy the ability to fly based on
a too far aft center of lift and a too far forward center of balance, or too small of a wing for such a large mass, no worries mate! If the imagination can soar, then so can these giants (NOT!) In the above YouTube video, the invalid, but traditional batwing shape of the brachiopatagium is best seen in the pterosaur’s shadow.

Quetzalcoatlus northropi

Tierra de dinos - Quetzalcoatlus Northropi

IF the wing membranes seem encumbering
awkward, liable to trip up the pterosaur or catch on some low lying shrub, no worried, mate! As these pterosaurs once wandered, let your imagination wander. There’s no need to precisely follow the evidence (Fig. 2) that shows the wing membranes essentially disappearing while flexed/folded.

Quetzalcoatlus at the Toledo Zoo

It’s going to be difficult to raise the wings for flight
given some of these awkward quadrupedal poses. Much better to have the center of balance over the toes at all static times (see below), shifting the balance forward while running at full speed (Fig. 5), like birds.

http://www.iaapa.org/DigitalShowDaily/2016/wed/Billings.asp

https://hiveminer.com/Tags/quetzalcoatlus

Why is this Houston Museum Quetzalcoatlus posed like this? Very strange.

Why is this Houston Museum Quetzalcoatlus posed like this? Very strange.

Quetzalcoatlus neck poses. Dipping, watching and displaying.

Quetzalcoatlus neck poses from David Peters Studio. Dipping, watching and displaying. Yes, the third finger is wrong here. It should be pointing posteriorly.

With what we know about pterosaurs
this should be a golden age of restoration. Instead, these models will someday be seen for what they are… near misses. They replace elegance with awkwardness, facts with fancy, and precision with tradition.

References
https://blog.everythingdinosaur.co.uk/blog/_archives/2018/06/01/pterosaur-models-go-on-display.html

Peters D 2002. A New Model for the Evolution of the Pterosaur Wing – with a twist 
Historical Biology 15: 277-301

Pterosaur brain cavities vs. posture (Unwin 2003, Witmer et al. 2003)

Unwin 2003
reviewed views on pterosaur posture (Fig.1 ) from the Witmer et al. 2003 paper on pterosaur brains and ‘smart’ wings. Unfortunately, Unwin illustrated his Nature review with freehand illustrations from Wellnhofer 1991 (Fig. 1) that bear little resemblance to traced bone sizes and proportions.

And no one raised a finger in protest.

Figure 1. Images from Unwin 2003 compared to fossil tracings and reconstructions from ReptileEvolution.com. Dashed line above toes in Rhamphorhynchus indicates center of balance, below the wing root, as in birds. Note the imaginative illustrations Unwin uses with little to no basis in reality. The skulls display the appropriate rostral tilt.

Figure 1. Images from Unwin 2003 compared to fossil tracings and reconstructions from ReptileEvolution.com. Dashed line above toes in Rhamphorhynchus indicates center of balance, below the wing root, as in birds. Note the imaginative illustrations Unwin uses with little to no basis in reality. The skulls display the appropriate rostral tilt.

Inaccuracies are permitted for some workers. 
Some have license to misrepresent, to advance bogus concepts, and to omit taxa. Then again, mistakes do happen. If so when were these mistakes (Fig. 1) noted and corrected by pterosaur workers over the past 15 years? After all… this is science and accuracy should be paramount. Fact should not be confused with fantasy.

Pterosaur workers
have advanced competing hypotheses and reconstructions, but I rarely if ever, have seen them specifically criticizing competing hypotheses and reconstructions. (Send examples or citations of this if you have them.) Instead, pterosaur workers seem to avoid criticizing the work of colleagues, leaving that to bloggers.

Ironically
Unwin is of the mind set that pterosaurs were dinosaur relatives — but gave Rhamphorhynchus a sprawling, lizard-like posture (Fig. 1), with fingers pointing anteriorly. I gave Rhamphorhynchus an erect posture in the knowledge that some lepidosaurs were occasional bipeds while others, like Sharovipteryx (Fig. 2), Longisquama and Bergamodactylusdid not employ their forelimbs at all during terrestrial locomotion.

Figure 2. Cosesaurus was experimenting with a bipedal configuration according to matching Rotodactylus tracks and a coracoid shape similar to those of flapping tetrapods. Long-legged Sharovipteryx was fully committed to a bipedal configuration.

Figure 2. Cosesaurus was experimenting with a bipedal configuration according to matching Rotodactylus tracks and a coracoid shape similar to those of flapping tetrapods. Long-legged Sharovipteryx was fully committed to a bipedal configuration.

Caption to Unwin’s 2003 figures:
“The horizontal alignment of the lateral semi-circular canal, indicated by the red line, is consistent with a crouching posture and forward-directed head in basal pterosaurs, represented by Rhamphorhynchus. b, In derived forms such as Anhanguera, the reorientation of the canal can be interpreted in terms of an upright position and a downward-pointing head. (Pterosaurs redrawn from Wellnhofer 1991 and not to scale.)”

Back to the abstract of Witmer et al. 2003.
“Comparison of birds and pterosaurs, the two archosaurian flyers, sheds light on adaptation to an aerial lifestyle. The neurological basis of control holds particular interest in that flight demands on sensory integration, equilibrium, and muscular coordination are acute. Here we compare the brain and vestibular apparatus in two pterosaurs based on high-resolution computed tomographic (CT) scans from which we constructed digital endocasts. Although general neural organization resembles birds, pterosaurs had smaller brains relative to body mass than do birds. This difference probably has more to do with phylogeny than flight, in that birds evolved from nonavian theropods that had already established trends for greater encephalization. Orientation of the osseous labyrinth relative to the long axis of the skull was different in these two pterosaur species, suggesting very different head postures and reflecting differing behaviours. Their enlarged semicircular canals reflect a highly refined organ of equilibrium, which is concordant with pterosaurs being visually based, aerial predators. Their enormous cerebellar floccular lobes may suggest neural integration of extensive sensory information from the wing, further enhancing eye- and neck-based reflex mechanisms for stabilizing gaze.”

References
Unwin DM 2003. Smart-winge pterosaurs. https://www.nature.com/articles/425910b.pdf
Witmer LM, Chatterjee S, Franzosa J. and Rowe T 2003. Neuroanatomy of flying reptiles and implications for flight, posture and behavior. Nature 425, 950–953.

 

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 giant Romanian pterosaur mandible fragment

FIgure 1. LPB R 2347 largest pterosaur mandible compared to Bakonydraco.

FIgure 1. LPB R 2347 largest pterosaur mandible compared to Bakonydraco.

Vremir et al. 2018
describe a pterosaur mandible fragment (Figs. 1, 2), “This specimen represents the largest pterosaur mandible ever found and provides insights into the anatomy of the enigmatic giant pterosaurs.”

Figure 2. LPR pterosaur mandible compared to related taxa, like Eopteranodon, and to the largest known pterosaur, Quetzalcoatlus. Figure 2. LPR pterosaur mandible compared to related taxa, like Eopteranodon, and to the largest known pterosaur, Quetzalcoatlus.

Figure 2. LPR pterosaur mandible compared to related taxa, like Eopteranodon, and to the largest known pterosaur, Quetzalcoatlus to scale.

It’s worthwhile
to place the jaw fragment in context with other pterosaurs. We don’t have a similar jaw fragment for the big Quetzalcoatlus (Fig. 2), which likely stood twice as tall as the giant eopteranodontid owner of the jaw fragment. Bakonydraco is a likely eopteranodontid, larger than Eopteranodon, but much smaller than the jaw fragment owner.

Earlier this jaw fragment was used as the basis for restoring the rest of this pterosaur as a giant azhdarchid nicknamed, ‘Dracula’ (with beaucoup errors, Fig. 2).

Figure 1. Dracula the giant azhdarchid pterosaur museum mount. Hopefully it's not too late to fix the problems here.

Figure 2. Dracula the giant pterosaur model built and based on the jaw fragment in today’s post. That’s a lot of imagination!

References
Vremir M et al. 2018. Partial mandible of a giant pterosaur from the uppermost Cretaceous (Maastrichtian) of the HaÈeg Basin, Romania. Lethaia doi: https://doi.org/10.1111/let.12268 https://onlinelibrary.wiley.com/doi/abs/10.1111/let.12268

Elgin 2014 PhD dissertation: nits and picks, part 2

Yesterday we looked at
a misidentified pterosaur specimen from the State Museum of Natural History Karlsruhe (SMNK) that wasn’t a Tupuxuara, as originally considered by then PhD candidate, now Dr. Ross Elgin. Consider that blogpost an unlabeled part 1.

Today, in part 2,
we’ll take a look at several other aspects of the Ross Elgin 2014 PhD dissertation that deserve credit and discredit. 

FIgure 1. GIF animation of image from Elgin 2014 comparing wing membrane configurations and a more accurate rendition of what Peters 2002 proposed. See text for list of issues here.

FIgure 1. GIF animation of image from Elgin 2014 comparing wing membrane configurations and a more accurate rendition of what Peters 2002 proposed and Elgin distorted. See text for list of issues here. In reality, like birds, the wings and hind limbs are decoupled. Most pterosaur workers don’t believe the data.

Elgin mistakenly reported,
“The primary flight membrane is reconstructed with an ankle attachment of the trailing edge, a configuration that was never fundamentally altered throughout the evolutionary history of the group.” Elgin ignored the bipedal origin of the Pterosauria, and ignored the narrow-chord wing membrane data that attends every pterosaur fossil that preserves wing membranes, perhaps on the advice of his mentors. And he ignored, or distorted, what Peters 2000 actually proposed for a wing membrane and bone orientation in pterosaurs (Fig. 1).  This may be what happens when you have to feed the fever dreams of a teacher rather than recording actual data. Elgin reported, “I am indebted to both David Hone and Eberhard “Dino” Frey for the opportunity to undertake this project and continue my work on the enigmatic and curious group of animals known as pterosaurs.”

Solutions to problems with Figure 1.

  1. The scapula/coracoid should rotate laterally, as in all articulated fossils
  2. The elbow should angle further posteriorly, as in all articulated fossils (+ in birds and bats)
  3. The pteroid should point to the deltopectoral crest, as in all articulated fossils
  4. The free fingers should point ventrally (in flight). Crushing typically rotates the narrow claws anteriorly.
  5. Metacarpals 1-3 should line up anteriorly, not stack themselves against mc4
  6. Manual 4.4 should articulate at an angle to m4.3, deepening the wing tip
  7. The propatagium should extend only to the deltopectoral crest
  8. The brachiopatagium should stretch between the wingtip and elbow with a fuselage fillet to distal thigh, as in all articulated fossils
  9. The thigh should be more meaty based on the long anterior ilium
  10. The femur should extend laterally to form a horizontal stabilizer (and note the sprawling lepidosaur orientation!)
  11. The femur should be flipped, as shown in Elgin’s figure 3 below.
  12. The uropatagia do not extend to the tiny tail, as in all articulated fossils. Elgin’s essentially creates a single uropatagium, which is a decades-old false paradigm.

If the above illustration by Elgin 2014 looks familiar
it’s because we looked at it earlier here, after publication of Elgin, Hone and Frey 2011. Also check out all the images on this ReptileEvolution page. You might remember these authors employed the fiction of ‘wing membrane shrinkage’ to explain the data, instead of just accepting the data, as is. We have evidence of pterosaur ancestors with wings decoupled from the hind limbs, so there was never a gliding transitional phase. Flapping preceded flying.

Along the same lines
Hone and Benton (2007, 2008); and Nesbitt and Hone (2010a, b) preferred to see things their own way, rather than strict adherence to the data. So, maybe young Elgin was unduly influenced by his professors and mentors.

Elgin introduced us to
Microtuban altivolan, which he described as a non-azhdarchid azhdarchoid. In the large pterosaur tree (LPT, 1189 taxa) Microtuban indeed nests at the base of the azhdarchid clade, arising from certain dorygnathids, a relationship Elgin never tested.

Elgin suggested a possible sexual dimorphism,
“where the pelvic girdle lacks a symphysis and remains open even in large adults is observed within Coloborhynchus robustus.” (Fig. 2) Let’s blame this one partly on his teachers, and partly on sloppiness. The base of the pterosaur pelvis opens and closes in phylogenetic patterns, not gender patterns. That tricked up Bennett, too, and Elgin’s teachers followed Bennett’s mistakes. On the sloppiness point, the open pelvis of C. robustus is missing its ventral and posterior borders, exactly the bones needed to join the ischia together (Fig. 2). It doesn’t look like Elgin was cheating. He must have thought he was not cheating. But he was cheating by changing points of view, changing scales and not adding back missing bone to follow generic patterns. This was all resolved with a little tracing in Adobe Photoshop, the paleontologists’ best friend.

Figure 1. Elgin compared these two Coloborhynchus pelves together, but failed to align them, scale them and add back missing bone.

Figure 2. Elgin compared these two Coloborhynchus pelves together, but failed to align them, scale them and add back missing bone.

Elgin was also tricked by
traditional archosaur patterns and paradigms in ontogeny. He expected sutures to close at adulthood. This tricked up Bennett, too. Instead, since pterosaurs are lepidosaurs, sometimes they do, sometimes they never do, and sometimes they fuse sutures before maturity and reading their final size. It’s all phylogenetic, not ontogenetic with lepidosaurs, including pterosaurs.

Figure 2. Coloborhynchus robustus (bones and outlines from Elgin 2014) compared to C. spielbergi (b&w). Notes added.

Figure 3. Coloborhynchus robustus (bones and outlines from Elgin 2014) compared to C. spielbergi (b&w). Notes added.

Elgin described and did not illustrate the missing wing finger:
“The wing finger phalanges in almost all pterosaurs are similar in form with expanded proximal and distal margins, the shafts of which show various degrees of curvature. Those preserved in SMNK PAL 1133 are not exception and agree well with other descriptions of pterodactyloids.” This description can only be the result of naiveté and inexperience. In reality phalanx proportions change between genera and species. It would have been helpful to see the wing finger of C. robustus since the C. spielbergi wing finger is missing.

Elgin 2014 mistakenly considered

  1. pterosaurs to be archosauromorphs.
  2. anurognathids to be basal pterodactyloids
  3. Darwinopterus to be ‘an animal intermediate’ linking basal to derived pterosaurs

These issues are resolved and settled
here and here when you add more taxa. It’s good for science to be critical. If nothing else this blog will hopefully show all readers that published scientific text and figures can sometimes include errors that can be exposed and corrected by colleagues.

It’s not okay
to disfigure the figures of other workers and then claim that’s the essence of their work (contra Fig. 1). We’ve seen this before with other PhDs.

References
Elgin RA, Hone DWE and Frey E 2011. The extent of the pterosaur flight membrane. Acta Palaeontologica Polonica 56 (1), 2011: 99-111. doi: 10.4202/app.2009.0145
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.
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 2008. Contrasting supertree and total evidence methods: the origin of the pterosaurs. Zitteliana B28:35–60.
Nesbitt SJ and Hone DWE 2010a. An external mandibular fenestra and other archosauriform character states in basal pterosaurs. Palaeodiversity 3: 225–233
Nesbitt SJ and Hone DWE 2010b. An external mandibular fenestra and other archosauriform character states in basal pterosaurs. Palaeodiversity 3: 225–233

https://pterosaurheresies.wordpress.com/2012/04/13/a-supertree-of-pterosaur-origins-hone-and-benton-2007-2009/

http://www.reptileevolution.com/pterosaur-wings.htm

https://pterosaurheresies.wordpress.com/2011/11/18/did-dimorphodon-have-an-external-mandibular-fenestra/

https://pterosaurheresies.wordpress.com/2013/01/16/a-closer-look-at-the-antorbital-fossa-in-two-pterosaurs-raeticodactylus-and-dimorphodon/