Carpus evolution in human ancestry back to basal reptiles

Out of 3400 prior posts
only two prior posts focused on carpals. One looked at the prepollux (radial sesamoid) of pandas and the pteroid + preaxial carpal of pterosaurs. Two looked at whale carpals here.

At present
the large reptile tree (LRT, 1825+ taxa) includes relatively few carpal traits, and none related to the migration of the pisiform and carpal 4 in mammals (see below). Crocodylomorphs elongate the proximal carpals. Many taxa do not ossify the carpals. As mentioned above, fenestrasaur centralia migrate  to become the pteroid and preaxial carpal in pterosaurs. So some carpals are more interesting than others.

FigFigure 1. Diplovertebron right manus dorsal view. Carpal elements colored.

Figure 1. (Left) Diplovertebron right manus dorsal view. Carpal elements colored. (Right) Thrinaxodon right manus dorsal view. Some elements rotated to fit reconstruction. Some phalanges are reduced to discs in Thrinaxodon on their way to disappearing in mammals.

I was also interested
in the origin of the styliform process on the human ulna. It is located where the pisiform is located in Diplovertebron (Fig. 1) a basal archosauromorph amphibian-like reptile. And thus began a look at sample taxa in the lineage of humans.

The next step
was the basal cynodont, Thrinaxodon (Fig. 1). Here the elements are larger, link closer to one another and are better ossified. Some phalanges are reduced to discs in Thrinaxodon on their way to disappearing in mammals.

Figure 2. Right manus of the platypus, Ornithorhynchus and early therian, Eomaia. Carpal elements colored.

Figure 2. Right manus of the platypus, Ornithorhynchus (left) and early therian, Eomaia (right). Carpal elements colored. Note the disappearance (or fusion) of distal tarsal 4 in Eomaia along with the centralia.

The next step in carpal evolution is represented by the basalmost mammal,
Ornithorhynchus (Fig. 2), the platypus. Here distal tarsal 5 is ventral to the lateral centralia. The pisiform is tiny. The radiale and ulnare completely cap the radius and ulna. The platypus is a highly derived monotreme, not a basal taxon.

The enlargement of the distal radius width
relative to the distal ulna width begins with Eomaia (Fig. 2), a basal therian. So does the enlargement of distal carpal 5, taking the place of distal carpal 4.

The migration of tiny distal 4 to the palmar surface
is documented in the evolution of human carpals (Fig. 4), but probably originated with Eomaia (Fig. 2) where distal tarsal 4 is not diagrammed.

At this point it is worth noting
that mammal carpals have different names than those of other tetrapods. Here are the mammal homologs (which we will ignore):

Proximal Tarsals:

    • Radiale = Scaphoid (lavendar)
    • Intermedium = Lunate (tan)
    • Ulnare = Triquetrum (dull pink)
    • Pisiform = Pisiform (yellow green)

Centralia

    • Medial Centralia = Prepollex (blue gray)
    • Lateral Centralia = Lateral Centralia (blue gray)

Distal Tarsals:

    • DT1 = Trapezium (yellow)
    • DT2 = Trapezoid (orange)
    • DT3 = Capate, magnum (green)
    • DT4+5 = Hamate, unciform (4= blue, 5=purple)
Figure 3. Right manus dorsal view of basal tree shrew, Ptilocercus (left), and basal lemur, Indri (right). Carpal elements colored.

Figure 3. Right manus dorsal view of basal tree shrew, Ptilocercus (left), and basal lemur, Indri (right). Carpal elements colored.

The next step in carpal evolution is represented by a basal placental,
Ptilocercus (Fig. 3), a tree shrew close to the base of the gliding and flying mammals. The fusion of distal tarsal 3 to the medial centrale is seen in Ptilocercus and its descendants. The ulna has a styloid process and the pisitorm extends laterally. Distal tarsal 1 is medially elongate to support a diverging thumb, further supported by the medial centralia.

Turns out the styloid process of the ulna
is not a fused carpal, but a novel outgrowth of the distal ulna appearing in basal placentals. The styloid process may have something to do with the ability of basal placentals to laterally rotate the manus for tree climbing in any orientation, including inverted, and to create a stop to prevent further rotation. Bats take this ability to its acme during wing folding.

Figure 4. Manus of human (Homo) in dorsal (left) and ventral/palmar (right) views. Carpal elements colored.

Figure 4. Manus of human (Homo) in dorsal (left) and ventral/palmar (right) views. Carpal elements colored. Carpal 4 and pisiform palmar only. Compare to Diplovertebron (Fig. 1) in which so little has changed, including relative finger length.

The final step in carpal evolution
takes us from the lemur, Indri (Fig. 3) to the human, Homo (Fig. 4). Here a ventral (palmar) view of the manus is also provided so we can finally see the ultimate destination of distal tarsal 4.

Before finishing this blog post
scroll back and forth between figures one and four to see how close the human hand and all of its proportions so greatly resembles that of a very basal ampibian-like reptile. Even the relative finger length is the same. This is probably the most important takeaway today. The LACK of change is the news story here. Dinosaurs, horses and snakes cannot make the same statement.

There is no reason to continue using
the mammal specific identification of the carpals in paleontology when those bones are homologs to tetrapod wrist bones going back to the Devonian. Medical communities should also start using tetrapod homologs and let the analog identities fade into history.

Simply put:
There are five distal carpals named one through five in tetrapods. Some of them fuse with other carpals. There are three centralia. Some of these fuse with other carpals. Tetrapods have three proximal carpals. Their names are easy. The radiale is on the radius. The ulnare is on the ulna. The intermedium is intermediate between them. These tend not to fuse with other carpals, at least in basal placentals. And finally the pisiform appears by itself on the lateral margin sometimes in contact with the distal ulna sometimes not.

On a similar note,
we supported earlier efforts to provide tetrapod homologs for fish skull bones here. Make things simple. There is enough hard work out there without needlessly translating bone identities.


References
Hamrick MW and Alexander JP 1996. The Hand Skeleton of Notharctus tenebrous (Primates, Notharctidae) and Its Significance for the Origin of the Primate Hand. American Museum Novitates 3182, 20pp.
Kielan-Jaworowska Z 1977. Evolution of the therian mammals in the Late Cretaceous of Asia. Part n. Postcranial skeleton in Kennalestes and Asioryctes. In: Z. Kielan-Jaworowska (ed.) Results Polish Mongolian Palaeont. Expeds. VIII. – Palaeont, Polonica, 37, 65-84.
Peters D 2009. A reinterpretation of pteroid articulation in pterosaurs. Journal of Vertebrate Paleontology 29:1327-1330.
Salesa MJ, Antón M, Peigné S and Morales J 2005. Evidence of a false thumb in a fossil carnivore clarifies the evolution of pandas. Proceedings of the National Academy of Sciences of the United States of America. abstract and pdf

Did ALL ichthyosaurs transform manual phalanges into carpal-like elements?

Short answer to the headline question:
No.

Only a few ichthyosaurs
turned their manual phalanges into interlocking bones resembling carpal elements (Fig. 1).

Fernández et al. 2020 propose
a hypothesis in which “all marine mammals and the majority of the reptiles, the fin is formed by the persistence of superficial and interdigital connective tissues, like a ‘baby mitten’, whereas the underlying connectivity pattern of the bones does not influence the formation of the forefin. On the contrary, ichthyosaurs ‘zipped up’ their fingers and transformed their digits into carpal-like elements, forming a homogeneous and better-integrated fore fin.”

The ‘baby mitten’ is readily apparent.
The digits of many aquatic tetrapods lose their individual identities as the flesh between the fingers no longer dissolves away during their embryonic development, thereby retaining the embryonic ‘mitten’ into adulthood.

But let’s not stereotype ichthyosaur flippers
based on a few that are different from the majority (Fig. 1). Many ichthyosaurs also have a ‘baby mitten’ retaining five digits within an embryonic mitten and often extra phalanges starting with digits #2 and #5. A few ichthyosaurs have only three digits.

A few have six or seven digits.
These are the few that have the greatest number of interlocking phalanges, making their paddles harder and more rigid.  

Figure 1. A selection of ichthyosaur manus with red metacarpals and blue carpals demonstrating great variation.

Figure 1. A selection of ichthyosaur manus with red metacarpals and blue carpals demonstrating great variation. Some transformed phalanges into zipped-up carpal-elements, others did not.

When put into a phylogenetic context,
ichthyosaurs arise from the derived pachypleurosaur/mesosaur, Wumengosaurus, in the large reptile tree (LRT). So when we look at the evolution of ichthyosaur fins from a starting point that includes the proximal outgroup taxon (Fig. 1) Wumengosaurus must lead that list.

According to the LRT,
Cartorhynchus and Sclerocormus nest elsewhere, with the sauropterygian, Qianxisaurus, whenever it and other basal sauropterygians are included within the taxon list.

Confession:
Without more informative graphics, I was unable to decipher the meaning of the diagrams in Fernández et al. 2020 figure 1. The “Anatomical networks showing the forelimb-to-forefin transition in SECAD tetrapods stemming from a basic tetrapod limb, highlighting the main types of morphological changes.” I think I would have been able to decipher the diagram if an underlying diagram of the actual manus was alongside or ghosted beneath the graphic of the ‘anatomical network‘ graphics.


References
Fernández MS et al. (7 co-authors) 2020. Fingers zipped up or baby mittens? Two main tetrapod strategies to return to the sea. Biology Letters 16: 20200281.  http://dx.doi.org/10.1098/rsbl.2020.0281

 

Antarctanax: a late-surviving basal synapsid, not a dino ancestor

Please see the notes in the following comments section. Most importantly after publication the authors report an errant scale bar, nearly doubling the apparent size of one of the pedes. 

Peecook, Smith and Sidor 2019
bring us news of a Early Triassic amniote from the Transantarctic Mountains, Antarctanax shackletoni (Figs. 1, 2), “known from a partial postcranial skeleton including cervical and dorsal vertebrae, a humerus, and both pedes.” 

Figure 1. Antarctanax manus and pes in situ with original tracing and color added here.

Figure 1. Antarctanax manus and pes in situ with original tracing and color added here.

Unfortunately,
if the scale bars are correct, and they seem to be, the smaller ‘pes’, the one surrounded by cervicals, is really a manus (Figs. 1, 2). Furthermore, the small manus matches the small humerus and radius. Added later: The scale were not correct, as noted at top.

Figure 2. Antarctanax manus and pes compared to those of Cabarzia and Aerosaurus, two basal synapsids.

Figure 2. Antarctanax manus and pes compared to those of Cabarzia and Aerosaurus, two basal synapsids. As you can see, basal synapsids rather quickly evolved similarly sized hands and feet.

The authors mislabeled
the robust, displaced metatarsal 5 as metatarsal 1, which lies beneath it (colored orange, Figs. 1, 2). Perhaps a reconstruction would have helped expose this error before submission.

The authors report,
“Our inclusion of A. shackletoni in phylogenetic analyses of early amniotes finds it as an archosauriform archosauromorph.” Their cladogram based on Ezcurra et al. 2014 nested Antarctanax in an unresolved polytomy with the basal archosauriforms, Proterosuchus, Erythrosuchus and Euparkeria. Their cladogram based on Ezcurra 2016 nested Antarctanax in an unresolved polytomy with other basal archosauriforms, FugusuchusSarmatosuchus. I am not aware of a manus or pes preserved for these two taxa. Of the above listed taxa, Proterosuchus (Fig. 3) comes closest, but has a hooked metatarsal 5 and metacarpal 3 is the longest, distinct from Antarctanax.

Synaptichnium

Figure 3. Synaptichnium compared to a slightly altered pes of Proterosuchus. Note a reduction of one phalanx in pedal digit 4 to match one less pad in the ichnite. The last two (or three phalanges) of pedal 4 are unknown in Proterosuchus.

This time it is not taxon exclusion, but bad timing.
When the manus and pes of Antarctanax are added to the large reptile tree (LRT, 1395 taxa), Antarctanax nests with basalmost synapsids, like Cabarzia (Figs. 2, 4) and Aerosaurus (Fig. 2). Aerosaurus was included in Ezcurra et al. 2014 and tested by Peecook, Smith and Sidor 2019. You’ll have to ask the authors why Antarctanax did not nest closer to Aerosaurus. Cabarzia trostheidei (Spindler, Werneberg and Schneider 2019, Fig. 3) could have influenced their thinking and scoring, but it was published only a few weeks ago, too late to include in their submission.

Figure 1. Cabarzia in situ and tracing distorted to fit the photo from Spindler, et al. 2019. Inserts show manus and pes with DGS colors and reconstructions. Scale bar = 5 cm.

Figure 4. Cabarzia in situ and tracing distorted to fit the photo from Spindler, et al. 2019. Inserts show manus and pes with DGS colors and reconstructions. Scale bar = 5 cm.

Peecock, Smith and Sidor did not provide a reconstruction
of Antarctanax, but online Discover magazine provided an in vivo painting and crowned it, “Dinosaur Relative Antarctanax.” According to the LRT, Antarctanax was a late-surviving (Early Triassic) basal member of our own lineage, the Synapsida, with a late Carboniferous genesis.

Therapsid synapsids were plentiful in Antarctica in the Early Triassic.
The headline should have focused on the unexpected presence of this sprawling, pre-pelycosaur, basal synapsid in the Mesozoic, surviving the Permian extinction event in this Antarctic refuge, alongside a closer relative of mammals, Thrinaxodon.


References
Ezcurra MD, Scheyer TM and Butler RJ 2014. The origin and early evolution of Sauria: reassessing the Permian saurian fossil record and the timing of the crocodile-lizard divergence. PLoS ONE 9:e89165.
Ezcurra MD 2016. The phylogenetic relationships of basal archosauromorphs, with an emphasis on the systematics of proterosuchian archosauriforms. PeerJ 4:e1778.
Peecook BR, Smith RMH and Sidor C 2019. A novel archosauromorph from Antarctica and an updated review of a high-latitude vertebrate assemblage in the wake of the end-Permian mass extinction. Journal of Vertebrate Paleontology e1536664 (16 pages) DOI: 10.1080/02724634.2018.1536664
Spindler F, Werneberg R and Schneider JW 2019. A new mesenosaurine from the lower Permian of Germany and the postcrania of Mesenosaurus: implications for early amniote comparative osteology. PalZ Paläontologische Gesellschaf

The prehensile hand and foot of Caluromys

Figure 1. Pteropus and Caluromys compared in vivo and three views of their skulls. Caluromys is in the ancestry of bats and shows where they inherited their inverted posture.

Figure 1. Pteropus and Caluromys compared in vivo and three views of their skulls. Caluromys is in the ancestry of bats and shows where they inherited their inverted posture.

I could not find
and still cannot find a complete skeleton for Caluromys (Fig. 1), the transitional marsupial leading to placentals. Argot 2001 published images of the hand in vivo. Argot 2002 published images of the foot in vivo and as an incomplete set of bones (Fig. 2). I matched those bones to the foot, still wishing I had all the bones, as in an X-ray.

Figure 1. Caluromys hand and foot from Argot 2002 compared to Didelphis and repaired here to match.

Figure 2. Caluromys hand and foot from Argot 2001, 2002 compared to the pes of Didelphis. The manus and pes of primates, tree shrews (in Glires) and basal arboreal Carnivorans all arise from Caluromys. These demonstrate the early appearance of the prehensile/opposable big toe and thumb, derived from the semi-opposable big toe of Didelphis, the Virginia opossum and even more so in Caluromys. 

The prehensile manus and pes of Caluromys
is primitive for the Eutheria (= Placentalia). From these arise the wings of bats, the flippers of whales, the hooves of horses as well as the fingers I just used to type this sentence.

References
Argot C 2001. Functional-Adaptive Anatomy of the Forelimb in the Didelphidae, and the Paleobiology of the Paleocene Marsupials Mayulestes ferox and Pucadelphys andinus. Journal of Morphology 247:51–79.
Argot C 2002. Functional-adaptive analysis of the hindlimb anatomy of extant marsupials and the paleobiology of the Paleocene marsupials Mayulestes ferox and
Pucadelphys andinus. Journal of Morphology 253:76–108.

Horse fingers

Here’s a paper that recovers overlooked data.
So it parallels what is done here.

Horses have only one finger and one toe,
right?

Now horses have portions of all five toes… extending to the hooves
(Solounias et al. 2017). And everybody overlooked that, until now (Figs. 1, 2).

Figure 1. Horse fingers 2 and 4 extend to the hoof, separated from the metacarpals 2 and 4 by unossified tissue. Image from Solounias et al. 2017.

Figure 1. Horse fingers extend to the hoof, separated from metacarpals by unossified tissue. Image from Solounias et al. 2017.

From the abstract:
“We revisit digit reduction in the horse and propose that all five digits are partially present in the modern adult forelimb.”

Figure 2. The modern horse, Equus, and extinct horse, Mesohippus manus. Colors over bones added here.

Figure 2. The modern horse, Equus, and extinct horse, Mesohippus manus. Colors over diagrams are from the original diagrams. Colors over proximal metacarpals added here. Image from Solounias et al. 2017.

Vestiges count.
Even when fused, bones are still there.

References
Solounias, et al. (6 co-authors) 2017. The evolution and anatomy of the horse manus with an emphasis on digit reduction. Royal Society Open Science 5:17182. http://dx.doi.org/10.1098/rsos.171782

 

Eogranivora has chicken feet and 6 fingers

Higher resolution data
and DGS color overlays reveal that the Early Cretaceous chicken, Eogranivora, has overlooked manual and pedal digits (Fig. 1). Digit zero makes an appearance here. Fusion was much less apparent than traced. Pedal digit 1 was overlooked, despite the tracing of pedal 1.1.

Figure 1. The manus and pes of the Early Cretaceous chicken, Eogranivora. Here digit 0 makes an appearance on the manus along with vestigial digits 4 and 5. On the pes pedal digits 1 (cyan) and 5 (purple) were overlooked. Here DGS reveals them. Overlay changes ever 5 seconds. The process of fusion implied by the drawings is not yet complete under DGS.

Figure 1. The manus and pes of the Early Cretaceous chicken, Eogranivora. Here digit 0 makes an appearance on the manus along with vestigial digits 4 and 5. On the pes pedal digits 1 (cyan) and 5 (purple) were overlooked. Here DGS reveals them. Overlay changes ever 5 seconds. The process of fusion implied by the drawings is not yet complete under DGS.

Earlier we looked at Eogranivora and nested it with Gallus the extant chicken using low-rez data. Here even the skull is updated with plate and counter plate revealing data overlooked by original authors (Figs. 2,3 for those who don’t review updated blog posts).

Figure 1b. Eogranivora skull in situ (plate and counterplate) in higher resolution.

Figure 2. Eogranivora skull in situ (plate and counterplate) in higher resolution.

Figure 1c. Skull of Eogranivora in situ and reconstructed using DGS, replacing a lower resolution attempt. Some details added for the palate here.

Figure 3. Skull of Eogranivora in situ and reconstructed using DGS, replacing a lower resolution attempt. Some details added for the palate here.

Eogranivora edentulata (Zheng et al. 2018; Early Cretaceous, Yixian Fm. Aptian, 125 mya; STM35-3) was earlier referred to Hongshanornis by (Zheng et al. 2011) who found evidence for an avian crop, along with feathers, gastroliths and seeds in the present specimen. Distinct from the holotype of HongshanornisEogranivora is toothless. This specimen is a direct link from the Early Cretaceous to the present day. With larger wings and a smaller body Eogranivora would have been a better flyer than extant chickens.

Figure 2. Gallus, the chicken, nests as a sister to the Early Cretaceous, Eogranivora, also a seed-eater.

Figure 4.. Gallus, the chicken, nests as a sister to the Early Cretaceous, Eogranivora, also a seed-eater. Note the length of the robust scapula.

With robust ribs
and a scapula extending back to the pelvis, Gallus, the chicken stands out from most birds. Eogranivora, if I have this right, also has robust ribs and an extended scapula (Fig. 5). Preservation is a funny thing when plates are split from counter plates. Sometimes we see the bone. Sometimes we see an impression of bone. Sometimes the bone splits down the middle and we see the inside of the bone. Here the parts of the scapula appear to be below and above the ribs, hence, my trepidation.

Figure 5. The crop, gizzard, sternum and scapulae of Eogranivora with DGS color overlays. Some guesswork here.

Figure 5. The crop, gizzard, sternum and scapulae? of Eogranivora with DGS color overlays. Some guesswork here. Some vertical bones apparently cross over and under the horizontal ribs. 

References
Zheng X, O’Connor JK, Wang X, Wang Y and Zhou Z 2018. Reinterpretation of a previously described Jehol bird clarifies early trophic evolution in the Ornithuromorpha. Proceedings of the Royal Society B 285: 20172494
Zheng X-T, Martin LD, Zhou Z-H, Burnham DA, Zhang F-C and Miao D 2011. Fossil evidence of avian crops from the Early Cretaceous of China. Proceedings of the National Academy of Sciences. USA 108: 15 904–907

wiki/Eogranivora
wiki/Gallus

Long carpals on crocodylomorphs = quadrupedal stance?

Figure 1. Terrestrisuchus is a bipedal basal crocodylomorph with elongate proximal carpals.

Figure 1. Terrestrisuchus is a bipedal basal crocodylomorph with elongate proximal carpals.

Long proximal carpals,
like the radiale and ulnare in Terrestrisuchus (Figs. 1, 2; Crush 1984), distinguish most crocodylomorphs from all basal dinosaurs (Fig. 2).

The question is:
why did long carpals develop? A recent comment from a reader suggested they enabled quadrupedal locomotion. But looking at the proportions of Terrestrisuchus does not inspire great confidence in that hypothesis. Terrestrisuchus has elongate carpals AND it seems to be comfortably bipedal with hands that only descend to the knees. And the pectoral girdle is relatively gracile.

Figure 2. Manus of several crocodylomorphs compared to the basal dinosaur, Herrerasaurus. Not sure what those two bones are on Junggarsuchus as they were cut off as shown when published.

Figure 2. Manus of several crocodylomorphs compared to the basal dinosaur, Herrerasaurus. Not sure how long those two proximal carpals are on Junggarsuchus. They were cut off as shown when published. Since the distal carpals are labeled (dc) I assume  proximal carpals are cut off below them. Oddly the radiale is much smaller than the ulnare if so, or rotated beneath it, unlike the other crocs.

Looking back toward more primitive taxa
provides only one clue as to when the proximal carpals first started elongating: with Terrestrisuchus. The following basal and often bipedal croc taxa unfortunately do not preserve carpals.

  1. Lewisuchus
  2. Gracilisuchus
  3. Saltopus
  4. Scleromochlus
  5. SMNS 12591
  6. Litargosuchus
  7. Erpetosuchus

Phylogenetic bracketing suggests that all
were bipedal or facultatively bipedal. Post-crania is missing or partly missing in several of these specimens.

Gracilisuchus

Figure 3. Gracilisuchus does not preserve the hands or carpals, but was possibly experimenting with bipedal locomotion based on its proximity to taxa that were obligate bipeds. Note the tiny pectoral girdle.

The distal carpals,
wherever preserved (Figs. 2, 3), appear to be small, scarce and flat, the opposite of a supple flexible wrist. So the proximal carpals of crocs comprise the great majority of the wrist, distinct from dinosaurs (Fig. 2).

Figure 3. Alligator carpals.

Figure 3 Alligator carpals. Of course, this is a quadruped that has inherited long carpals from bipedal ancestors in the Triassic.

So… what do other bipedal taxa do with their hands?
Cosesaurus, a bipedal ancestor to pterosaurs, probably flapped, based on the shape of its  stem-like coracoid and other traits. Herrerasaurus, a bipedal ancestor to dinosaurs had elongate raptorial unguals (claws) lacking in any basal crocodylomorph (Fig. 2). Such claws were probably used in grasping prey in dinos… not so much in crocs.

The elongate proximal carpals in crocodylomorphs
appear to extend the length of the slender antebrachium (forearm) of Terrestrisuchus for only one reason at present. The offset lengths of the shorter radius and longer ulna become subequal again with the addition of the longer radiale and shorter ulnare. So there is no simple hinge joint at the antebrachium/proximal carpal interface. So that joint was relatively immobile. The lack of deep distal carpals also suggests a lack of mobility at the metacarpal/distal carpal interface in basal taxa. However in extant crocs, that hinge appears to be more flexible.

Figure 5. Trialestes parts. Note the much larger ulna relative to the radius and the much longer forelimb relative to the bipedal basal crocs.

Figure 5. Trialestes parts. Note the much larger ulna relative to the radius and the much longer forelimb relative to the bipedal basal crocs.

In Trialestes
(Fig. 5) the elongate fore limbs more closely match the hind limbs. So the elongate carpals in Trialestes do appear to enhance a secondarily evolved quadrupedal stance.

Also take a look at
Hesperosuchus, Dromicosuchus, Protosuchus. Saltoposuchus, Dibrothrosuchus, Baurusuchus, Simosuchus, and Pseudhesperosuchus. After long carpals first appeared in Terreistrisuchus, they do not change much despite the many other changes in the morphology of derived taxa. Bipeds have them. Quadrupeds have them. Long-bodied taxa have them, Short-bodied taxa have them.

Some thoughts arise
when considering the first crcoc with elongate carpals, Terrestrisuchus.

  1. At some point in the day Terrestrisuchus probably rested on its elongate pubis bone (the first in this lineage), flexing its long hind limbs beneath itself to do so. In that pose elongate carpals may have been useful in steadying the animal as it balanced on the pubis tip and whenever it rose to a bipedal stance.
  2. A male Terrestrisuchus may have used its hands to steady itself while riding on the back of a female while mating. The carpals were elongated as part of the balancing act performed during this possibly awkward bipedal conjugation.
  3. Coincidentally, the coracoids in crocodylomorphs begin to elongate in this taxon. So freed from quadrupedal locomotion duties, basal crocs may have done some early form of flapping as part of a secondary sexual behavior, long since lost in extant taxa.

So, in summary
I think the elongate carpals developed in crocs with a really long pubis to steady it while resting. Very passive. Not sure what other explanation explains more.

Did I miss anything?
Has anyone else promoted similar or competing hypotheses?

References
Crush PJ 1984. A late upper Triassic sphenosuchid crocodilian from Wales. Palaeontology 27: 131-157.

wiki/Terrestrisuchus

The forgotten clade: the REAL proximal ancestors to Dinosauria

Ignored by Baron et al. 2017, and everybody else
the Junggarsuchus clade (including Pseudhesperosuchus, Carnufex and Trialestes in order of increasing quadrupedality, Figs. 1–4) nests as the proximal ancestors to Herrerasaurus (Fig. 1) and the rest of the Dinosauria (Fig. 5) in the large reptile tree (LRT). That cladogram tests a wider gamut of taxa in greater detail than any other reptile cladogram ever published, attempting to not overlook anything. The Junggarsuchia is a sister clade to the Crocodylomorpha with both arising from a taxon near Lewisuchus (Fig. 1). Traditional paleontology (see Wikipedia) nests this largely ignored clade with the sphenosuchian crocodylomorphs (Fig. 4)… and for two good reasons!

Figure 1. Members of the Junggarsuchus clade were derived from a sister to the basal crocodylomorph, Lewisuchus and produced one line that includes Pseudhesperosuchus and Trialestes. The other line produced dinosaurs. These taxa are shown to scale. Note the evolution from a bipedal configuration to a quadrupedal stance.

Figure 1. Members of the Junggarsuchus clade were derived from a sister to the basal crocodylomorph, Lewisuchus and produced one line that includes Pseudhesperosuchus and Trialestes. The other line produced dinosaurs. These taxa are shown to scale. Note the evolution from a bipedal configuration to a quadrupedal stance.

One: Paleontologists never seem to include Dinosauria
in their smaller gamut croc analyses because they’re looking at crocs!~. So once again, taxon exclusion is holding some workers back from seeing ‘the big picture’. ReptileEvolution.com and the blog you are currently reading is all about examining ‘the big picture.’

Figure 2. Skulls of the Junggarsuchus clade not to scale. Herrerasaurus is the basalmost dinosaur.

Figure 2. Skulls of the Junggarsuchus clade not to scale. Herrerasaurus is the basalmost dinosaur, closely related to Junggarsuchus.

Two: Junggarsuchians ALSO have elongate proximal wrist bones
Elongate proximal carpals are found in both sphenosuchian crocs and derived members of the Junggarsuchus clade. Paleontolgists wrongly assumed such odd wrist bones were homologous. It’s an easy mistake to make. However, the LRT makes clear that intervening taxa, including Junggarsuchus, do not have elongate wrist bones.

Among taxa that preserve the manus,
(Fig. 3) it is Junggarsuchus that nests closest to Herrerasaurus and the Dinosauria.

Figure 3. Hands of Lewisuchus, Herrerasaurus, Junggarsuchus, Pseudhesperosuchus and Trialestes. The proximal carpals (radiale and ulnare) were elongate by convergence with a line of crocodylomorphs. This has confused paleontologists and mentally removed them from possible ancestry to the Dinosauria. Note the very short proximal carpals in Junggarsuchus.

Figure 3. Hands of Lewisuchus, Herrerasaurus, Junggarsuchus, Pseudhesperosuchus and Trialestes. The proximal carpals (radiale and ulnare) were elongate by convergence with a line of crocodylomorphs. This has confused paleontologists and mentally removed them from possible ancestry to the Dinosauria. Note the very short proximal carpals in Junggarsuchus.

Like the basal members of the Crocodylomorpha
the Junggarsuchus clade (the Prodinosauria here) transition from bipedal basal members to quadrupedal derived members, with the longest forelimbs belonging to the most derived member, Trialestes (Fig. 3). Distinct from the others and contra the original interpretation, I think Trialestes may have had a larger ulnare than radiale, to match its larger ulna.

Figure 4. Crocodylomorph manus and carpus samples including Terrestrisuchus, Erpetosuchus, Hesperosuchus and Dibothrosuchus along with Scleromochlus documenting the elongate radiale and ulnare on derived taxa. Ticinosuchus is the closest example of an ancestral/plesiomorphic manus in the LRT.

Figure 4. Crocodylomorph manus and carpus samples including Terrestrisuchus, Erpetosuchus, Hesperosuchus and Dibothrosuchus along with Scleromochlus documenting the elongate radiale and ulnare on derived taxa. Ticinosuchus is the closest example of an ancestral/plesiomorphic manus in the LRT.

Let’s not forget
PVL 4597 (Fig. 6) which was mistakenly considered a specimen of Gracilisuchus by (Lecuona and Desojo 2011), but under phylogenetic analysis in the LRT, still nests as the proximal outgroup to Herrerasaurus. It is tiny specimen, supporting the hypothesis of phylogenetic miniaturization at clade origin. And it retains a small proximally oriented calcaneal tuber, as found in other Junggarsuchians.

Figure 1. Subset of the LRT focusing on the Archosauria (Crocodylomorpha + Dinosauria and kin). Gray areas document specimens with elongate proximal carpals (radiale and ulnare).

Figure 5. Subset of the LRT focusing on the Archosauria (Crocodylomorpha + Dinosauria and kin). Gray areas document specimens with elongate proximal carpals (radiale and ulnare).

We looked at
phylogenetic miniaturization at the origin of several pterosaur clades. Well, it happens here too, at the base of the Dinosauria (Fig. 1) with PVL 4597 (Fig. 6), easily overlooked, easily mistaken for something else.

One should not ‘choose’ outgroup taxa
based on paradigm, tradition, guessing, convenience or opinion. Rather outgroup taxa should ‘choose themselves’ based on rigorous testing of a large gamut of outgroup candidates in phylogenetic analysis. To minimize selection bias, the LRT provides 858 outgroup taxa the opportunity to nest close to dinosaurs.

Figure 6. The closest known taxa to the Dinosauria, PVL 4597, is a tiny taxon (phylogenetic miniaturization) with erect hind limbs, a large and deep pelvis and a tiny calcaneal tuber.

Figure 6. The closest known taxa to the Dinosauria, PVL 4597, is a tiny taxon (phylogenetic miniaturization) with erect hind limbs, a large and deep pelvis and a tiny calcaneal tuber.

 

References
Baron MG, Norman DB, Barrett PM 2017. A new hypothesis of dinosaur relationships and early dinosaur evolution. Nature 543:501–506.
Bonaparte JF 1969. 
Dos nuevos “faunas” de reptiles triásicos de Argentina. Gondwana Stratigraphy. Paris: UNESCO. pp. 283–306.
Butler RJ. et al. 2014. New clade of enigmatic early archosaurs yields insights into early pseudosuchian phylogeny and the biogeography of the archosaur radiation. BMC Evol. Biol. 14, 128.
Clark JM et al. 2000. A new specimen of Hesperosuchus agilis from the Upper Triassic of New Mexico and the interrelationships of basal crocodylomorph archosaurs. Journal of Vertebrate Paleontology 20 (4): 683–704.
doi:10.1671/0272-4634(2000)020[0683:ANSOHA]2.0.CO;2.
Clark JM, Xu X, Forster CA and Wang Y 2004. A Middle Jurassic ‘sphenosuchian’ from China and the origin of the crocodilian skull. Nature 430:1021-1024.
Lecuona A and Desojo, JB 2011. Hind limb osteology of Gracilisuchus stipanicicorum(Archosauria: Pseudosuchia). Earth and Environmental Science Transactions of the Royal Society of Edinburgh 102 (2): 105–128.
Nesbitt SJ 2011. The early evolution of archosaurs: relationship and the origin ofmajor clades. Bull. Amer. Mus. Nat. Hist. 352, 1–292.
Novas FE 1994. New information on the systematics and postcranial skeleton of Herrerasaurus ischigualastensis (Theropoda: Herrerasauridae) from the Ischigualasto
Reig OA 1963. La presencia de dinosaurios saurisquios en los “Estratos de Ischigualasto” (Mesotriásico Superior) de las provincias de San Juan y La Rioja (República Argentina). Ameghiniana 3: 3-20.
Sereno PC and Novas FE 1993. The skull and neck of the basal theropod Herrerasaurusischigualastensis. Journal of Vertebrate Paleontology 13: 451-476. doi:10.1080/02724634.1994.10011525.
Zanno LE, Drymala S, Nesbitt SJ and Schneider VP 2015. Early Crocodylomorph increases top tier predator diversity during rise of dinosaurs. Scientific Reports 5:9276 DOI: 10.1038/srep09276.

wiki/Pseudhesperosuchus
wiki/Junggarsuchus
wiki/Carnufex
wiki/Herrerasaurus
wiki/Sanjuansaurus

 

Diplovertebron and amphibian finger loss patterns

Updated June 13, 2017 with the fact that Diplovertebron is the same specimen I earlier illustrated as Gephyrostegus watsoni. And the Watson 1926 version of Diplovertebron (Fig. 1) was so inaccurately drawn (by freehand) that the data nested is apart from the DGS tracing. Hence this post had deadly errors now deleted.

Figure 2. The gradual loss of basal tetrapod fingers. Unfortunately fingers are not known for every included taxon.

Figure 2. The gradual loss of basal tetrapod fingers. Unfortunately fingers are not known for every included taxon. Odd Tulerpeton with 6 fingers may result from taphonomic layering of the other manus peeking out below the top one. See figure 6. Mentally delete Diplovertebron from this chart. 

The presence of five manual digits
in Balanerpeton (Figs. 4, 5) sheds light on their retention in Acheloma + Cacops. There is a direct phylogenetic path between them (Fig. 2). Note that all other related clades lose a finger or more. Basal and stem reptiles also retain five fingers.

Figure 2. Utegenia nests as a sister to Diplovertebron.

Figure 3. Utegenia nests as a sister to Diplovertebron.

Distinct from the wide frontals
in Utegenia and Kotlassia,  Balanerpeton (Fig. 4) had narrower frontals like those of Silvanerpeton, a stem reptile.

Figure 4. The basal amphibian, Balanerpeton apparently has five fingers (see figure 5).

Figure 4. The basal amphibian, Balanerpeton apparently has five fingers (see figure 5).

As reported
earlier, finger five was lost in amphibians,while finger one was lost in temonospondyls. Now, based on the longest metacarpal in Caerorhachis and Amphibamus (second from medial), apparently manual digit one was lost in that clade also, distinct from the separate frog and microsaur clades. In summary, loss from five digits down to four was several times convergent in basal tetrapods.

Figure 5. DGS recovers five fingers in Balanerpeton with a Diplovertebron-like phalangeal pattern.

Figure 5. DGS recovers five fingers in Balanerpeton with a Diplovertebron-like phalangeal pattern. Two 5-second frames are shown here.

Finally, we have to talk about
Tulerpeton (Fig. 6). The evidence shows that the sixth manual digit is either a new structure – OR – all post-Devonian taxa lose the sixth digit by convergence, since they all had five fingers. Finger 6 has distinct phalangeal proportions, so it is NOT an exposed finger coincident rom the other otherwise unexposed hand in the fossil matrix.

Figure 2. Tulerpeton manus and pes in situ, reconstructed by Lebdev and Coates 1995 and newly reconstructed here.

Figure 6. Tulerpeton manus and pes in situ, reconstructed by Lebdev and Coates 1995 and newly reconstructed here. Digit 6 is either a new structure, or a vestige that disappears in all post-Devonian taxa.

References
Fritsch A 1879. Fauna der Gaskohle und der Kalksteine der Permformation “B¨ ohmens. Band 1, Heft 1. Selbstverlag, Prague: 1–92.
Kuznetzov VV and Ivakhnenko MF 1981. Discosauriscids from the Upper Paleozoic in Southern Kazakhstan. Paleontological Journal 1981:101-108.
Watson DMS 1926. VI. Croonian lecture. The evolution and origin of the Amphibia. Proceedings of the Zoological Society, London 214:189–257.

wiki/Diplovertebron

Better data for the manus of Eryops

Just found this reference
Dr. David Dilkes (2015) provides photo data (Fig. 1) on the carpus and manus of Eryops the giant temnospondyl. Earlier the best data I had was a decades old (Romer era) reconstruction and based on that manus and those of its sister taxa. With that data it appeared that the four digits preserved were 2–5, not 1–4 as traditionally considered. Dilkes likewise follows tradition in listing the fingers as 1–4.

Figure 1. Forelimb of Eryops from Dilkes 2005. Here freehand drawings of the manus cannot compete with a taking a tracing of the photo and restoring the digits and carpal elements to their in vivo positions. Note the subtle differences that happen in the freehand drawing by Dilkes and the Romer era illustrator.

Figure 1. Forelimb of Eryops from Dilkes 2005. Here freehand drawings of the manus cannot compete with  a tracing of the photo and restoring the digits and carpal elements to their in vivo positions (middle). Note the subtle differences that happen in the freehand drawing by Dilkes (above) and the Romer era illustrator (below).

The present data further cements
the hypothesis that the fingers of Eryops are 2–5, not 1–4.

And further cements
the hypothesis that freehand drawing is not as accurate as tracing a photo of the bones.

Today’s post also demonstrates
that better data, no matter where it comes from or makes your hypothesis go, must be incorporated. And finally…

Today’s post also demonstrates
that good Science can take place with second-hand data.

References
Dilkes D 2015. Carpus and tarsus of Temnospondyli. Vertebrate Anatomy Morphology Palaentology 1(1):51-87.