Basal reptile hands: Casineria and Diplovertebron

I reexamined two fossils
via photos and found ways to improve the interpretation of both of them, Casineria (Fig. 1) and Diplovertebron (Fig. 2).

Figure 1. Manus of Casineria, a basal archosauromorph reptile. The carpals are unosssified, but left vague impressions in the matrix. Other bones overlapped the carpals and are removed here. PIls added.

Diplovertebron punctatum (Fritsch 1879, Waton 1926; DMSW B.65, UMZC T.1222a; Moscovian, Westphalian, Late Carboniferous, 300 mya) aka:  Gephyrostegus watsoni Brough and Brough 1967) and Gephyrostegus bohemicus (Carroll 1970; Klembara et al. 2014) after several name changes perhaps this specimen should revert back to its original name as it nests a few nodes away from Gephyrostegus.

Derived from a sister to Eldeceeon, Diplovertebron was basal to the larger Solenodonsaurusand the smaller Brouffia, Casineria and Westlothiana. Diplovertebron was a contemporary of Gephyrostegus bohemicus, Upper Carboniferous (~310 mya), so it, too, was a late survivor.

Overall smaller and distinct from Eldeceeon, the skull of Diplovertebron had a shorter rostrum, larger orbit and greater quadrate lean. The dorsal vertebrae formed a hump and had elongate spines. The hind limbs were much longer than the forelimbs. The tail is incomplete, but appears to have been short and deep. Seven sphere shapes were preserved alongside this specimen. They may be the most primitive amniote eggs known.

Figure 2. Diplovertebron manus in situ and reconstructed with PILs added. What appear to be displaced carpals may be something else entirely. The carpals may have been unossified, as in Casineria. See how DGS makes reconstruction less chaotic?

Casineria kiddi (Paton, Smithson & Clack 1999) Visean, Mississippean, Carboniferous, ~335 mya was a small basal archosauromorph. the oldest but not the most primitive. It was derived from a sister to Diplovertebron and Solenodonsaurus. Westlothiana was a sister taxon.

Overall smaller than and distinct from Gephyrostegus, the skull of Casineria had no otic notch. See Brouffia for more possible skull details. The cervicals of Casineria were increased in number but decreased in size. The presacral vertebral count had increased to over 30. Ribs discontinued after #22. Apparently two vertebrae formed the sacrum and were connected to the pelvis. The pectoral girdle was composed of unfused elements. The humerus had a small hourglass shape. The manus was enlarged. The ilium had no anterior dorsal process. The femur was more gracile. The pes was reduced, more nearly the size of the manus.

References
Brough MC and Brough J 1967. The Genus Gephyrostegus. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences 252 (776): 147–165.
Carroll RL 1970. The Ancestry of Reptiles. Philosophical Transactions of the Royal Society London B 257:267–308. online pdf
Fritsch A 1879. Fauna der Gaskohle und der Kalksteine der Permformation “B¨ ohmens. Band 1, Heft 1. Selbstverlag, Prague: 1–92.
Klembara J, Clack J, Milner AR and Ruta M 2014. Cranial anatomy, ontogeny, and relationships of the Late Carboniferous tetrapod Gephyrostegus bohemicus Jaekel, 1902. Journal of Vertebrate Paleontology 34:774–792.
Paton RL Smithson TR and Clack JA 1999. An amniote-like skeleton from the Early Carboniferous of Scotland. Nature 398: 508-513.
Watson DMS 1926. VI. Croonian lecture. The evolution and origin of the Amphibia. Proceedings of the Zoological Society, London 214:189–257.

wiki/Gephyrostegus
wiki/Diplovertebron
wiki/Casineria

 

Pandion, the osprey, joins the LRT

And the osprey,
Pandion haliaetus (Linneaus 1758) joins the large reptile tree (LRT, 1124 taxa) at the base of (owls + swifts) + (Old World vultures + falcons). The secretary bird, Sagittarius, and the terror birds are proximal outgroups.

Figure 1. Pandion, the osprey, nests at the base of the birds of prey, sans the secretary bird and seriema.

So the osprey is a basal
short-legged, arboreal birds-of-prey.

Wikipedia reports,
“The osprey differs in several respects from other diurnal birds of prey. Its toes are of equal length, its tarsi are reticulate, and its talons are rounded, rather than grooved. The osprey and owls are the only raptors whose outer toe is reversible, allowing them to grasp their prey with two toes in front and two behind. It has always presented something of a riddle to taxonomists, but here it is treated as the sole living member of the family Pandionidae, and the family listed in its traditional place as part of the order Falconiformes.”

References
Linnaeus C 1758. Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata.

wiki/Osprey

 

Were early pterosaurs inept terrestrial locomotors?

Witton 2015 asked:
“Were early pterosaurs inept terrestrial locomotors?” Sorry, this online paper escaped my notice until now. It’s two years old.

The answer is
an unqualified “YES” when Witton turns perfectly good bipeds (supported by morphology, outgroups (Fig. 2), ichnites and omitted citations), into stumbling quadrupeds encumbered by imaginary wing membranes (Fig. 3) that connect the ankles and lateral pedal digits to the wing tips and binds the legs together with a single uropatagium. The Unwin influence is strong in those English youngsters. He also rotates the humerus in a shoulder joint that does not permit rotation (Fig. 1), which would be very bad for a flapping reptile, bird or bat.

Figure 1. Tracings from bones (on left) compared to Witton’s freehand quads. Comments in red.

From the Witton abstract:
“Pterodactyloid pterosaurs are widely interpreted as terrestrially competent, erect-limbed quadrupeds, but the terrestrial capabilities of non-pterodactyloids are largely thought to have been poor.”

This may be true when you construct pterosaurs that don’t match footprints and you have no idea where ‘early pterosaurs’ came from, even though that has been known for 17 years. Obligate bipeds (Longisquama and Sharovipteryx) are outgroups. Basalmost pterosaur, Bergamodactylus (Fig. 2) , has longer hind limbs and shorter forelimbs (Fig. 2) than other pterosaurs, retaining these plesiomorphic traits.

Figure 2. Updated reconstruction of Bergamodactylus to scale with an outgroup, Cosesaurus. Does this look like a quadruped to anyone? All derived pterosaurs have relatively shorter legs. Outgroups, whether the invalid Scleromochlus, or the valid Sharovipteryx, have long legs like these. Uropatagia are not preserved, but they are on a related taxa one node away, Sharovipteryx. Note the tail is NOT incorporated.

Witton’s abstract continues
“This is commonly justified by the absence of a non-pterodactyloid footprint record,”

(False, see Peters 2011)

“suggestions that the expansive uropatagia common to early pterosaurs”

(False, misinterpretation of Sordes)

“would restrict hindlimb motion in walking or running, and the presence of sprawling forelimbs in some species.”

(sprawling at the top, narrow gauge on the substrate (Fig. 3).

“Here, these arguments are re-visited and mostly found problematic. Further indications of terrestrial habits include antungual sesamoids, which occur in the manus and pes anatomy of many early pterosaur species, and only occur elsewhere in terrestrial reptiles, possibly developing through frequent interactions of large claws with firm substrates.”

Or possibly by grasping branches and tree trunks, but even that possibility is not considered or argued against by Witton.

Getting back to the uropatagium found in bats…
primitive bats extend a membrane from both legs back to the tail. Only in the most derived bats, like Desmodus (Fig. 3), is the tail a vestige to absent. The resulting uropatagium without the tail extends between the legs – while completely avoiding the toes. Thus the pterosaur/bat analog, is also bogus. Final point: basal bats don’t walk or run on their hind limbs. They hang. Only in bats like the vampire do some bats reacquire the ability to actively hop around on horizontal surfaces, like cow buttocks and grassy knolls.

Witton carefully avoids
any mention of papers in which bipedal pterosaur trackways are described (Peters 2011). He fully supports the uropatagium hypothesis proposed by Sharov 1971 and further supported by Unwin and Bakhurina 1994 (disputed by Peters 2002 and here). That uropatagium, found in no other specimens of Sordes or any other pterosaur, is really a displaced wing membrane (Figs. 3–5) along with a displaced radius and ulna as shown here. Note: a few days ago Witton’s latest illustration used pedal digit 5 to frame both the uropatagium and the brachiopatagium. No one else does this. No argument or explanation is given.

Figure 3. Above, from Witton 2017 focusing on the pterosaur uropatagium. Note: even though fanciful, it does not incorporate the tail, but goes from leg to leg, UNLIKE Desmodus the bat, which incorporates what little tail is left. Besides, their is NO homology here. Witton is trying to support a bad interpretation with a bad analogy. Not a good idea to support an analogy with invalid drawings. Witton gives no support through testing to the uropatagium controversy, but accepts it with blinders on.

Witton carefully avoids
any mention of other candidate pterosaur outgroups, like fenestrasaurs (Fig. 2), and the assistance they can offer to the questions posed, but supports the basal bipedal crocodylomorph, Scleromochlus, as a potential outgroup. Ironic, isn’t it?

My first question would be, which outgroup taxon has anything resembling a leg-spanning uropatagium?Certainly not phytosaurs. Nor any archosaur. Sharovipteryx has separate uropatagia, but in Witton’s world view those are not the same, nor are they to be mentioned, because that would involve citing some academic paper from Peters, which would be antithetical to Witton’s premise. In good science, all counterarguments are considered, attacked or supported.

Figure 4. The Sordes uropatagium is actually displaced wing material carried between the ankles by the displaced radius and ulna.

Witton supports
the invalid shrinkage hypothesis of Elgin, Hone and Frey (2011) to explain away narrow-chord wing membranes preserved in the fossil record…which would be ALL of them…

Figure 5. The hind limbs and soft tissues of Sordes. Above, color-coded areas. Below the insitu fossil. Note how insubstantial the illusory uropataigum is compared to the drawing that solidifies the area. Tsk.Tsk.

Witton reports,
“Trackways made by running pterodactyloids indirectly demonstrate how elastic their proximal membranes must have been, allowing track makers to take strides of considerable magnitude (Mazin et al., 2003) despite membranes stretching from the distal hindlimb to their hands (Elgin, Hone & Frey, 2011).” The other explanation is that the wings and hind limbs were always decoupled (as documented in all known fossils). Pterosaurs do not have a membrane extending to the ankles. Witton proposes a bounding gait for pterosaurs, even though no pterosaur tracks document this.

Figure 6. A plesiomorphic bat with the tail incorporated in the uropatagium. This bat, Myotis, cannot walk very well. Desmodus, highly derived, has required the ability to walk, but at the expense of its tail leaving a vestige uropatagium. Everything must be put into a phylogenetic context, even in analogies.

Thankfully Witton supports
“Assessments of pterosaur hindlimb muscle mechanics seem to confirm that the pterosaur pelvic and femoral musculoskeletal system is optimally configured for an erect stance.” 

But then he puts the fingers on the ground (Fig. 1). Why???

Perhaps Witton does not realize
what happens to his uropatagium when the pes is plantigrade, which is how Witton always reconstructs pterosaur pedes. Somehow he avoids drawing the lateral digit reversed toward the pelvis, as he proposed earlier.

Witton has no criticism
for one of his references, Hone and Benton 2007 (but did not cite the setup 2007 paper. Readers know, for many reasons, this is one of the worst papers ever published in this field. The facts will stun even freshmen paleontologists. 

Witton ignores the pterosaur sacrum,
which has more than the typical two sacrals found in a wide range of quadrupedal reptiles. Why does the pterosaur sacrum add and fuse vertebrae phylogenetically and more with larger taxa? For the same reason that humans, apes and theropod dinosaurs do. They are bipedal and the sacrum acts as the fulcrum to a long lever arm.

Earlier we talked about pterosaur workers wearing blinders, ignoring papers with hypotheses that conflicted with pet hypotheses. Now you see that happening in real time.

When workers, like Witton, stopped citing papers
I had published in academic journals is when I took my evidence and arguments online.

Earlier, in a multipart critique,
here, here, here, here and here we talked about Witton’s previously published work combined in a single book. I only wish someone with influence on Witton and his collaborators would remind them that their ideas and papers are going to end up like the Victorian-age cartoons they mock – unless they get back to facts and evidence.

References
Elgin RA, Hone DWE and Frey E 2011. The extent of the pterosaur flight membrane. Acta Palaeonntologica Polonica 56(1): 99-111.
Peters D 2002. A New Model for the Evolution of the Pterosaur Wing – with a twist. Historical Biology 15: 277–301.
Peters D 2011. A Catalog of Pterosaur Pedes for Trackmaker Identification
Ichnos 18(2):114-141. http://dx.doi.org/10.1080/10420940.2011.573605
Sharov AG 1971. New flying reptiles from the Mesozoic of Kazakhstan and Kirghizia. – Transactions of the Paleontological Institute, Akademia Nauk, USSR, Moscow, 130: 104–113 [in Russian].
Unwin DM and Bakhurina NN 1994. Sordes pilosus and the nature of the pterosaur flight apparatus. Nature 371: 62-64.
Witton MP 2015. Were early pterosaurs inept terrestrial locomotors? PeerJ 3:e1018 DOI 10.7717/peerj.1018

Pristidactylus: a Basiliscus sister without a crest

Figure 1. Pristidactlyus torquatus in vivo.

I got interested in the extant lizard, Pristidactylus
(Figs, 1, 2) when Bever and Norell 2017 used it as an outgroup to the clade Rhynchocephalia. The large reptile tree (LRT, 1122 taxa) using phylogenetic analysis falsifies that hypothesis of relationships.

Figure 2. Pristidactylus skull in 5 views. This iguanid lizard nests with the crested basilisk.

Pristidacatylus torquatus (Phillippi 1861, extant, snout-vent length = 6-11cm) is the extant forest lizard. It is related to Basiliscus and feeds on beetles. Image from Digimorph.org.

Figure 3. Basiliscus, the “Jesus” lizard, does not share as many traits as Draco and Chlamydosaurus do, but is related, given the short list of Iguanids currently employed.

Basiliscus basiliscus (Laurenti 1768) is the extant basilisk. It is related to Iguana but has a tall parietal crest. This frilled lizard is able to run bipedally across ponds. Skull image from Digimorph.org and used with permission.

References
Laurenti JN 1768. Specimen Medicum, Exhibens Synopsin Reptilium Emendatum cum
Experimentis Circa Venena et Antidota Reptilium Austriacorum. Wien.
Linnaeus C 1758. Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata.
Philippi RA and Landeck L 1861. Neue Wirbelthiere von Chile. Archiv für Naturgeschichte 27 (1): 289-301.

wiki/Iguana
wiki/Basiliscus_(genus)

Why do pterosaur workers ignore the most basic data?

I don’t know why,
but today’s leading pterosaur experts are actively ignoring the data from the last twenty years while inventing their own fanciful versions of what pterosaurs looked like (Fig. 1) – while claiming to be the latest word on the subject. Today we’ll be looking at a short paper from the latest Flugsaurier book by Hone, Witton and Martill 2017. And we’ll criticize the artwork that crystalizes their latest intentions. This is part 1.

For some reason
Hone, Witton and Martill like to show ancient cartoons that have little to no bearing on the present knowledge base (Fig. 1). I think it’s an English thing since most, if not all of the old engravings are indeed English in origin and easily lampooned. ‘See how far we’ve come!’, they seem to be saying. Doing so only takes up space that could otherwise go to competing current versions – which they want to avoid.

We’ve seen this
earlier when English professor D. Naish preferred to criticize work that preceded (= was not included in) ReptileEvolution.com. He employed cartoons made by others, rather than artwork that was actually posted on the website to show how bad the whole website was.

Evidently
It’s what they like to do. Someday, perhaps, they’ll look in a mirror and see some of the faults I present here… using their own artwork – which will soon enough joint their ancient engravings in a drawer full of foolish ideas they can draw upon in future decades.

Figure 1. Images from Hone, Witton and Martill 2017 showing the ‘evolution’ of our concept of Dimorphodon. Artists are credited in the text. Compare the latest color version to tracings of the several skeletons in figure 2. The long tail is based on a disassociated fossil probably belonging to a campylognathoid.

In figure 1
images of Dimorphodon through time are presented from Hone, Witton and Martill 2017.

1. Rev. GE Howman 1829. Probably based on the headless holotype BMNH R1034 (Fig. 2). The authors labeled this as ‘monstrous’ when ‘inaccurate’, ‘fanciful’ or ‘medieval’ would do.
2. Owen 1870. Probably based on the short-skull specimen, BMNH 41212 (Fig. 2), along with the disassociated tail specimen. The authors labeled this rendition as ‘ungainly, bat-like’. Odd word choice when among all the presented illustrations it is the one most like Witton’s 2017 version (#5).
3. H Seeley 1901. Probably based on the long-skull specimen, BMNH PV R 1035 (Fig. 2) In the their comment Hone, Witton and Martill report, ‘progressive interpretation of D. macronyx as an erect-limbed quadruped’, but note that a biped interpretation was also offered. They thought it best not to show that possibility. 
4. K Padian 1983. Probably based on the short-skull specimen, BMNH 41212 (Fig. 2). The authors report, ‘a highly active, bird-like digitigrade biped, a controversial interpretation that nevertheless symbolises the reinvention of pterosaurs in the late twentieth century.’ While there are minor issues associated with this figure (the orientation of fingers 1–3 and pedal digit 5, the over-extension of the metatarsophalangeal joint, the great length of the tail), it is the one that is most closely based on the skeleton (Fig. 2). BTW, when authors use the word, ‘controversial’ it usually means it does not fit their world view, but they have no evidence against it, nor any evidence to support their traditional hypothesis. 
5. M Witton 2017. Not sure which skeleton this one is based on as it appears to have been done entirely freehand from memory and imagination. The authors report, ‘Modern interpretation of D. macronyx adult and speculative juveniles reflecting contemporary interpretations of pterosaur soft tissues, muscle development and ecology.’ Ahem…we’ll run through this illustration step-by-step below.

Figure 2. Images of Dimorphodon from ReptileEvolution.com. The tail attributed to Dimorphodon is shown in figure 3.

You know, you really can’t go wrong
when you strictly adhere to the bones (Figs. 2,3), soft tissue (Peters 2002) and footprints of the most closely related taxa (Peters 2011), which were made by digitigrade and bipedal pterosaur trackmakers. Unfortunately no such citations appear in this chapter. Those are purposefully omitted.

Figur 3. Dimorphodon model by yours truly. The tail is too long based on the disassociated tail.

Witton
fell under the spell of the quad-launch hypothesis (Habib 2008), then took it one step further and made Dimorphodon a galloping hunter (Fig. 4), forsaking its wings and erect, digitigrade hind limbs (according to related ichnite makers) to hunt prey on mossy logs with backward pointing fingers. The finger unguals are again too small here.

While writing this I became aware
of Sangster 2003, a PhD thesis that evidently used computer modeling to show Dimorphodon was a quadruped. I have not seen the thesis and Ms. Sangster can no longer be found online. I wonder about these conclusions because:

1. PhD theses are, by definition, the work of inexperience workers; and
2. Sangster may have had to earn her PhD by succumbing to the unveiled interests of her English professors, as we’ve seen before here and here.

Figure 4. Galloping Dimorphodon by Mark Witton.

To counter the awkward, dangerous and ultimately unproductive
quad-launch scenario, I proposed the following bipedal launch animation (Fig. 5). It combines the hind limb leap with the first flap of the large wings to provide the maximum thrust at takeoff. In the Habib proposal, you don’t get that wing flap until later in the cycle – maybe too late in the cycle. The quad launch also depends on directing the force of liftoff through the fragile free fingers. They were not strong enough for that, especialy not when there is a better option available using giant muscles in the chest and pelvis. That’s why the sacrum is so strong, to act as a fulcrum on that long, heavy lever!

FIgure 5. Dimorphodon take off (with the new small tail).

So let’s get back
to Witton’s cover illustration (Fig. 6), which they tout as our contemporary view of Dimorphodon. I will note several inaccuracies (below). See figures 2 and 3 for accurate tracings.

Figure 6. Touted as the contemporary view of Dimorphodon, this Mark Witton illustration suffers from several fancies and inaccuracies.

1. No Dimorphodon as this shape of skull.
2. Needs a longer neck.
3. Free fingers should be long and the unguals much larger.
4. Wing appears to be too short with a too narrow wing tip chord.
5. Witton wants to connect the trailing edge membrane from wing tip to ankle (or lateral toe), but look at the tremendous stretch in the membrane when that happens. Seems to be getting dangerously close to the narrow-at-the-elbow wing design of Zittel, Schaller and Peters, which they want to avoid.
6. Ouch! This is a set of elongate toe bones with butt metatarsophalangeal joints – where Witton breaks them. This is not a calcar (a novel ossification on bat ankles which enters the uropatagium). One one side of these lateral toes the wing membrane attaches. On the other side the uroropatagium attaches. This is not shown in any fossil! Related taxa, from Langobardisaurus to Sharovipteryx, to Tanystropheus, with this same sort of elongate toe morphology, do not dislocate their bones this way. See Peters 2000 for a description that fits Rotodactylus tracks.
7. No pterosaur has a uropatagium. This comes from a misinterpretation of Sordes. Pterosaur do have paired uropatagia.
8. The tail is too large. On the BMNH 41212 fossil the traditionally overlooked tail is very small (Figs. 2, 7) This is in accord with related anurognathids. An unassociated tail has been attributed to Dimorphodon (Fig. 5) but it is robust and much longer. It probably belongs to a eudimorphodontid or campylognathoid. I”m surprised the tiny tail of Dimorphodon has gone unnoticed for so long. The specimen has been in English storage for over a hundred years. It was their responsibility for discovering this, but they chose instead to use their imaginations (Fig. 6).
9. No tail vane is known for Dimorphodon. Tail vanes are found in pterosaurs like Campylognathoides and Rhamphorhynchus, both with a robust tail. Vestigial tails are unlikely to have had tail vanes.

FIgure 7. The tail of Dimorphodon (BMNH 41212 specimen). See figure 2 for reconstruction.

I’m asking my Engllsh colleagues
|to step up their game and become more professional. Otherwise chaps from across the pond are going to continue pointing out the flaws in their thinking. I’m not going to say their approach is not scientific (as they say about my work), but when you forsake accuracy for artistry, you’re treading very close to that line.

References
Habib MB 2008. Comparative evidence for quadrupedal launch in pterosaurs. Zitteliana B28:159-166.
Hone DWE, Witton MP and Martill DM 2017. New perspectives on pterosaur paleobiology in Hone DWE, Witton MP and Martill DM (eds) New Perspectives on Pterosaur Palaeobiology. Geological Society, London, Special Publications, 455, https://doi.org/10.1144/SP455.18
Peters D 2000. Description and Interpretation of Interphalangeal Lines in Tetrapods 
Ichnos, 7: 11-41.
Peters D 2002. A New Model for the Evolution of the Pterosaur Wing – with a twist 
Historical Biology 15: 277-301
Peters D 2011. A Catalog of Pterosaur Pedes for Trackmaker Identification.
Ichnos 18(2):114-141. http://dx.doi.org/10.1080/10420940.2011.573605
Sangster S 2003. The anatomy, functional morphology and systematics of Dimorphodon macronyx (Diapsida: Pterosauria)..Unpublished PhD thesis, University of Cambridge.

Harpagolestes uintensis is a mesonychid. Harpagolestes macrocephalus is not.

Welcome to the wonderful world of convergence!
Harpagolestes uintensis (Fig. 2) and H. macrocephalus (Fig. 1) look similar enough to be considered similar, but they are not congeneric in the LRT. One of them needs a new generic name.

Of the several heresies
recovered by the large reptile tree (LRT, 1120 taxa) the latest is the separation of some former mesonychids (Fig. 1, Andrewsarchus, Sinonyx, Hapalodectes) from current and traditional mesonychids (Fig. 2, Mesonyx and Harpagolestes uintensis). The clade of former mesonychids now nests as giant tenrecs. This clade produced odontocete whales and transitional taxa. The latter group of true mesonychids gave rise to mysticete (baleen) whales and the following transitional taxa: hippos, anthrobunids and desmostylians.

Figure 1. Harpagolestes macrocephalus compared to sisters Sinonyx and Andrewsarchus to scale.

Today the addition of Harpagolestes macrocephalus
(Fig. 1) to the LRT nests it not congenerically with Harpagolestes uintensis (Fig. 2), but between Andrewsarchus and Sinonyx. So the two are not congeneric.

Figure 1. Andrewsarchus, Sinonyx, Mesonyx and Harpagolestes to scale for direct comparison of these two tenrecs with these two mesonychids.

We’ve seen convergence many times
in the LRT. This is just one more example of convergence that has been traditionally overlooked.

Deleting nine tenrecs
on either side of Sinonyx + Andrewsarchus + H. macrocephalus changes nothing in the LRT. The above taxa still nest with odontocetes far from mesonychids through mysticetes, though some loss of resolution occurs in the mammal subset of the LRT.

Deleting hippos and anthracobunids
from the mesonychid clade changes nothing.

References
O’Leary MA and Rose KD 1995. Postcranial skeleton of the early Eocene mesonychid Pachyaena (Mammalia: Mesonychia). Journal of Vertebrate Paleontology 15(2):401-430.

Pelagornis is a giant gannet in the LRT.

Quickly becoming one of the most famous of all birds
because of its great size and teeth, Pelagornis is, evidently, still a bird of mystery in the world of paleo-taxonomy.

Figure 1. Pelagornis skeletal elements.

Wikipedia reports,
“Pelagornis [was] probably rather close relatives of either pelicans and storks, or of waterfowl, and are here placed in the order Odontopterygiformes to account for this uncertainty. Like many pseudotooth birds, it was initially believed to be related to the albatrosses in the tube-nosed seabirds (Procellariiformes), but subsequently placed in the Pelecaniformes where it was either placed in the cormorant and gannet suborder (Sulae) or united with other pseudotooth birds in a suborder Odontopterygia.

Figure 2. Skull of Morus bassanus the Northern gannet. This taxon is most similar to Pelagornis in the LRT.

If experts can’t nest Pelagornis with certainty
based on rather complete morphology, then we have a problem. With the addition of the Northern gannet, Morus bassanus, Pelagornis nests with it with certainty in the large reptile tree (LRT, 1032 taxa).

Figure 3. Skeleton of Morus bassanus, the Northern gannet.

Pelagornis chilensis (Lartet 1857, Mayr and Rubilar-Rogers 2010; Miocene; MNHN SGO.PV 1061) is an extinct giant soaring bird related to the gannet Macronectes. Bony, not true teeth, developed along the jaw margins. The external naris was divided by bone. Mayr and Rubilar-Rogers reported, “We finally note that the phylogenetic affinities of bony-toothed birds still have not been convincingly resolved.”

Figure 4. The gannet (genus: Morus) in vivo. Note the diving pose below.

Morus bassanus (Linneaus 1758, extant; 100cm long) is the Northern gannet. Here the nasal has extended over the external naris to prevent water from entering the nose of this plunge diver. Secondary nostrils appear inside the mouth. The keratin at the jaw rims appears to form tiny teeth (Fig. 2). Compare to the much larger Pelagornis (Fig. 1).

Earlier we looked at the plunge diving possibility of the outwardly similar Late Jurassic pterosaur, Germanodactylus.

References
Lartet E 1857. Note sur un hum´erus fossile d’oiseau, attribu ´e `a un tr `es-grand palmip`ede de la section des Longipennes. Comptes rendus hebdomadaires des S´eances de l’Acad´emie des Sciences (Paris) 44:736–741.
Linnaeus C 1758. Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata.
Mayr G and Rubilar-Rogers D 2010. Osteology of a new giant bony-toothed bird from the Miocene of Chile, with a revision of the taxonomy of Neogene Pelagornithidae. Journal of Vertebrate Paleontology 30(5):1313-1340.

wiki/Macronectes
wiki/Pelagornis
wiki/Gannet

Early Cretaceous Longicrusavis houi under DGS

A complete and articulated,
except for a skull suspiciously emerging from/near the cloaca, Early Cretaceous ornithuromorph bird, Longicrusavis houi (O’Connor, Gao and Chiappe 2010; PKUP V1069), gets the DGS treatment today. That involves segregating the bones by coloring them, then rearranging them to an in vivo pose. This is done without seeing the specimen firsthand. You can see what little the authors saw in figure 1.

Figure 1. Skull of Longicrusavis houi in situ, as originally traced and colorized using the DGS method. Below is a reconstruction based on the colored bones. The palate was not reconstructed, but palatal bones are colored. Quite a bit more data is gleaned without having seen the fossil firsthand here. The occiput is well exposed here. Possible squamosal/quadratojugal contact here. The lacrimal was displaced to the dorsal frontal. The caudals intersect the skull and push the jugal (cyan) dorsally.

After analysis,
Longicrusavis nests in the large reptile tree (LRT, 1027 taxa) at the base of the Yanornis + Changzuiornis clade and these are sisters to Apsaravis + Ichthyornis and Gansus + Hesperornis among the basalmost neognaths. The O’Connor team recovered a similar nesting with a different list of taxa.

Figure 2. It’s always valuable to see what the taxon looks like with scale bars. This is a tiny specimen, but rather completely known. If you have a typical 72 dpi screen, the images is 1.5x life size.

O’Connor et al. report,
“There are no teeth preserved in PKUP V1069, though alveoli do appear to be present in the premaxilla and maxilla.” I see tiny teeth (Fig. 1). Phylogenetic bracketing indicates it could go either way as derived members of this clade redevelop teeth. The jugal, lacrimal, quadratojugal and several other bones were also overlooked by those who had firsthand access. We’ll see as time goes by and better data comes in.

References
O’Connor JK, Gao K-Q and Chiappe LM 2010. A new ornithuromorph (Aves: Ornithothoraces) bird from the Jehol Group indicative of higher-level diversity. Journal of Vertebrate Paleontology 30(2):311–321.

PKUP, Peking University Paleontological Collection, Beijing, China

Patagopteryx: it’s a hen-sized ostrich sister in the LRT

Patagopteryx deferrariisi (Late Cretaceous, 80mya; Alvarenga and Bonaparte 1992, Chiappe 1996, Chiappe 2002, MACN-N-03, 10, 11, 14 and others) was a hen-sized bird originally considered a ratite, but later (Chiappe 1996) nested it between Enantiornithes and Hesperonis. Back then Patagopteryx was one of only a few Cretaceous birds known. Here, with more included taxa, Patagopteryx nests with Struthio, the ostrich, back among the ratites.

FIgure 1. Patagopteryx compared to Struthio to scale and scaled to a similar shoulder height. This Late Cretaceous taxon retained four toes and a robust tail.

Four toes are present.
As in Casaurius pedal ungual 2 is elongate. The pubis tip bends ventrally, like a boot. The anterior skull is unknown but otherwise is similar to Struthio.

Figure 2. Patagopteryx skull from Chiappe 2002 with color restoration show the skull to be very ostrich like. Note the tiny squamosal barely overlapping the quadrate, smaller than in Struthio.

Aepyornis, the elephant bird, moves over the the tinamous with this taxon addition.

Figure 3. Struthio skull with a long maxilla.

References
Alvarenga and Bonaparte 1992. A new flightless land bird from the Cretaceous of Patagonia; pp. 51–64 in K. E. Campbell (ed.), Papers in Avian Paleontology, Honoring Pierce Brodkorb. Natural History Museum of Los Angeles County, Science Series 36.
Chiappe LM 1996. Early avian evolution in the southern hemisphere: Fossil record of birds in the Mesozoic of Gondwana. Memoirs of the Queensland Museum 39:533–556.
Chiappe LM 2002. Osteology of the flightless Patagopteryx deferrariisi from the late Cretaceous of Patagonia (Argentina) pp.281–316 in Mesozoic Birds, Above the Heads of Dinosaurs, Chapter: 13, Editors: Chiappe LM and Witmer LM, University of California Press.

http://www.reptileevolution.com/struthio.htm

The origin of hummingbirds

Updated November 17, 2020
with new data nesting the stilt, Himanopterus, with hummingbirds, both descending from Eocene Eocypselus.

Hummingbirds are the tiniest of living birds.
They are famous for hovering with wings beating so rapidly they essentially blur from view. Today hummingbirds live only in the New World.

Prum et al. 2015
based on DNA, nested hummingbirds with swifts and these nested with the nocturnal nightjars. That is the traditional nesting.

In the LRT 2015
based on morphology, hummingbirds nest with the extinct Eocypselus (Fig. 4, 50 mya) and the sea gull, Chroicocephalus (Figs. 1, 3; extant). Mayr 2004 reported on an Old World hummingbird, Eurotrochilus inexpectatus (Fig. 4; 30 mya), from the early Oligocene. Elsewhere on the cladogram, swifts nest with owls and owlets in the large reptile tree (LRT, 1129 taxa then, 1760+ taxa now).

In the LRT 2020
the stilt, Himantopus (Fig. x), nests closer to the hummingibird, Archilochus.

Figure x. Photo and skeletal drawing of Himantopus, the black-necked stilt.

Figure 1. A sea gull hovering. Many birds can do this for short periods, but sea gulls are phylogenetic sisters to hummingbirds, so this is where it all began for hummers.

It should be noted
that swifts do not feed while hovering. They speed through the air snatching insects in flight. On the other hand, gulls do hover (Figs. 1, 2). Gulls appear to hover only in a breeze, which is often present at shorelines. Thus gulls represent the awkward origin of hummingbird hovering, which improved with faster wingbeats. a deeper sternum and a smaller size. Stilts probe mud like hummingbirds probe flowers with their long slender beak.

Figure 2. The smallest gull, Hydrocoloeus_minutus, hovering while feeding.

Fossils tell us
that hummingbird-sized specimens, like Eocypselus (Figs. 3, 4), lived 50 mya and probably originated much earlier. One-sixth the size of the small gull, like Hydrocoloeus (Figs. 2, 3), Eocypselus had a relatively short, small beak and shorter legs, though still longer than the wings.

Figure 3. Chroicocephalus, the smaller Hydrocoloeus (the smallest living gull), the much smaller Eocypselus and the ruby-throated hummingbird, Archilochus to scale.

Of course, small size is key to hummingbird evolution.
At this point, I’m not aware of any gulls smaller than Hydrocoloeus, whether extant or in the fossil record. Stilts are taller, but that is due to those extreme legs and feet.

Figure 4. Eocypselus from 50 mya, Eurotrochilus, from 30 mya and Archilochus, the extant ruby-throated hummingbird to scale.

The fossil Eutrochilus
(Fig. 4, Mayr 2004) bridges the time gap between Eocypselus and extant hummingbirds and would appear to be a complete and fully realized hummingbird itself, living some 30 mya, while originating much earlier. Eocypselus (Fig. was not much different in size or morphology.

Figure 5. Gene studies link swifts to hummingbirds. Trait studies link swifts to owlets. Trait studies link hummingbirds to stilts.

Old World vs. New World
So, based on Eutrochilus, hummingbirds used to be in Europe. Now they are restricted to the New World. Why? There is a long list of hummingbird eaters online here. Something killed European hummingbirds in the Old World.

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References
Mayr G 2004. Old World fossil record of modern-type hummingbirds. Science 304:861–864,
Ksepka DT, Clarke JA, Nesbitt SJ, Kulp FB and Grande L. 2013. Fossil evidence of wing shape in a stem relative of swifts and hummingbirds (Aves, Pan-Apodiformes). Proceedings of the Royal Society B: Biological Sciences 280 (1761): 20130580. doi:10.1098/rspb.2013.0580. Supplementary materials here.
McGuire JA et al. (7 coauthors) 2014. Molecular Phylogenetics and the Diversification of Hummingbirds. Current Biology, 2014; DOI: 10.1016/j.cub.2014.03.016
Zhang Z, Feduccia A and James HF 2012. A Late Miocene Accipitrid (Aves: Accipitriformes) from Nebraska and Its Implications for the Divergence of Old World Vultures. PLoS ONE7(11): e48842. https://doi.org/10.1371/journal.pone.0048842

https://wordpress.com/post/pterosaurheresies.wordpress.com/10805
https://www.livescience.com/44593-first-hummingbird-evolutionary-tree.html