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

You heard it here first: Ichthyostega and Acanthostega were secondarily aquatic

In this YouTube video from 2018
Dr. Donald Henderson starts his online slide video presentation by repeating the traditional fin-to-finger story (Fig. 1).

Unfortunately
that story was already out-of-date in 2018 due to taxon exclusion in comparison to and competition with the phylogenetic analysis found in the large reptile tree (LRT, 1817+ taxa; subset Fig. 5).

Not surprisingly, Dr. Henderson thought it was “very peculiar”
that Middle Devonian tetrapod trackways preceded the Late Devonian fossils of tetrapods by tens of millions of years. The LRT solves this problem. Acanthostega and Ichthyostega are not transitional taxa, but dead end taxa with polydactyly not found in other tetrapod taxa. Their phylogenetic ancestors filled the gap between the Middle and Late Devonian, but those fossils have not been found yet in those strata, only in later strata as late survivors of those earlier radiations.

In the middle of the presentation
Dr. Henderson presented his alternative view: that Ichthyostega and Acanthostega were secondarily aquatic tetrapods. His YouTube video is dated January 11, 2018. Only a short month earlier the LRT recovered Ichthyostega and Acanthostega as secondarily more aquatic tetrapods, time-stamped here.

Evidently that was an idea whose time had come.
Or else Dr. Henderson read that hypothesis here and embraced it. Either way, Dr. Henderson did not employ phylogenetic analysis, but came to his solution as a notion to reconcile the Middle Devonian tracks to the late Devonian fossils.

Otherwise
Dr. Henderson’s presentation was mundane. Henderson’s customary family tree of vertebrates (Fig. 1) indicates he had no idea how clades of fish are related to one another at a species level (Fig. 2). He never tested traditional hypotheses, but accepted them without reservation.

Figure 1. Slide from Henderson's YouTube video with connections between clades highlighted in frame 2.

Figure 1. Slide from Henderson’s YouTube video with connections between clades highlighted in frame 2.

The fish phylogeny problem was resolved
here in 2019 and continues to evolve with every added taxon.

Figure 4. Shark skull evolution according to the LRT. Compare to figure 1.

Figure 2. Shark skull evolution according to the LRT. Compare to figure 1.

Dr. Henderson also presents a traditional lineup
of tetrapods (Fig. 3) that was improved by the LRT by simply adding overlooked taxa (Fig. 4).

Figure 3. Slide from Henderson YouTube presentation modified in frame 2 to reflect the order of basal tetrapods in the LRT. Missing here is Trypanognathus (Fig. 3) and kin, basal tetrapods in the LRT.

Figure 3. Slide from Henderson YouTube presentation modified in frame 2 to reflect the order of basal tetrapods in the LRT. Missing here is Trypanognathus (Fig. 4) and kin, basal tetrapods in the LRT.

Henderson’s traditional lineup is lacking several taxa,
like Trypanognathus (Fig. 4), that are also long, low and with tiny limbs, like Tiktaalik and Panderichthys, but are traditionally never included in fin-to-finger cladograms, other than here in the LRT.

Figure 6. Dorsal and ventral views of Panderichthys and several basal tetrapods demonstrating the low, flat skulls and bodies with small limbs and relatively straight ribs.

Figure 4. Dorsal and ventral views of Panderichthys and several basal tetrapods demonstrating the low, flat skulls and bodies with small limbs and relatively straight ribs.

It’s nice to have a notion, like Dr. Henderson had.
After all, that’s where all scientific inquiry has its genesis. But you can’t beat a good old, wide gamut phylogenetic analysis to make your notion into a testable hypothesis that covers all the other competing hypotheses. Let’s hope that someday PhDs will adopt a taxon list comparable to the LRT and then let the taxa and their taxonomy tell the tale.

Figure 4. Subset of the LRT focusing on basal tetrapods. Colors indicate number of fingers known. Many taxa do not preserve manual digits.

Figure 5. Subset of the LRT focusing on basal tetrapods. Colors indicate number of fingers known. Many taxa do not preserve manual digits.

Colleagues,
follow up those notions with testable analyses. It’s hard work, but it’s the professional thing to do.


References
https://pterosaurheresies.wordpress.com/2017/12/15/ichthyostega-and-acanthostega-secondarily-more-aquatic/

Molnar et al. 2021: Forelimb function across the fish–tetrapod transition

Molnar et al. 2021
bring us a deep and complex look into the hypothetical muscles (based on muscle scars) of Eusthenopteron (Fig. 1), Acanthostega (Fig. 1) and Pederpes. The authors compare these distinctly different taxa to “show that early tetrapods share a suite of characters including restricted mobility in hurmerus long-axis rotation, increased muscular leverage of humeral retraction, but not depression/adduction, and increased mobility in elbow flexion-extension.” The authors infer the earliest ‘steps’ in tetrapod forelimb evolution were related to limb-substrate interactions. Weight support appeared later.

Figure x. The fin to finger transition in the LRT with the addition of Elpistostege.

Figure 1. The fin to finger transition in the LRT with the addition of Elpistostege.

Unfortunately, 
without a valid phylogenetic context, what these authors deliver is not quite germane to the topic of their headline. The actual fin-to-finger transition occurred between Panderichthys (Figs. 1, 2) and the extremely similar Trypanognathus (Figs. 1, 2). The former had fins. The latter had fingers and toes. Otherwise they were very much alike.

Molnar et al. looked at the wrong taxa. Neither Panderichthys nor Trypanognathus are mentioned in the Molnar et al. text.

What can we conclude given
the similarities and differences of Panderichthys and Trypanognathus?

  1. Small fins and limbs at the transition were incapable of weight bearing
  2. Elbows and knees were incapable of bending, pushing, pulling
  3. Torso much longer than tail, lots of flexible ribs
  4. Low, wide, flexible torso at the transition provided serpentine locomotion
  5. Little risk of tipping over due to low center of gravity
Figure 6. Dorsal and ventral views of Panderichthys and several basal tetrapods demonstrating the low, flat skulls and bodies with small limbs and relatively straight ribs.

Figure 2. Dorsal and ventral views of Panderichthys and several basal tetrapods demonstrating the low, flat skulls and bodies with small limbs and relatively straight ribs.

Molnar et al. conclude:
“Together, these results suggest that competing selective pressures for aquatic and terrestrial environments produced a unique, ancestral “early tetrapod” forelimb locomotor mode unlike that of any extant animal.”

Not really. Consider the moray eel chasing crabs on land without fins or fingers. Click the pic to view video on YouTube. David Attenborough is the narrator.

Now put predator and prey in a Devonian swamp setting,
with lots of growing and rotting vegetation and no rocky place to find safety. Note (in figure 2), the rather slow phylogenetic growth of the limbs relative to the torso in this sequence. Other lineages did their own thing in their own time. Ossinodus, for instance (Fig. 1), had a shorter torso and longer limbs, and was a phylogenetic ancestor to Ichthyostega and Acanthostega.


References
Molnar JL, Hutchinson JR, Diogo R, Clack JA and Pierce SE. 2021. Evolution of forelimb musculoskeletal function across the fish-to-tetrapod transition. Science Advances 2021; 7: eabd7457 22 January 2021

Latest fins-to-fingers paper stumbles due to taxon exclusion

Dickson et al. 2020 bring us their views
on the transition from fins to feet at the base of the Tetrapoda.

Unfortunately,
taxon exclusion (Fig. 1), once again, mars this study published in Nature.

Figure 1. Cladogram from Dickson et al. 2020 with an overlay indicating taxa found in the LRT and key LRT taxa with limbs lacking in Dickson et al. 2020.

Figure 1. Cladogram from Dickson et al. 2020 with an overlay indicating taxa found in the LRT and key LRT taxa with limbs lacking in Dickson et al. 2020.

Cherry-picking taxa in Dickson et al. 2020
nested Ichthyostega and Acanthostega as the first taxa to have fingers and toes. These two were highly promoted in earlier works that included co-author, Jennifer Clack, but that should not excuse the exclusion of pertinent taxa. When more taxa are added to a cladogram (Fig. 2), these two famous basal tetrapods, with their large, well-formed limbs, nest not as transitional taxa, but as derived taxa leaving no descendants (subset Fig. 2). Their polydactyl extremities were evolutionary dead-end experiments perhaps reflecting one of the first returns to a more aquatic existence.

Employing a wider gamut of taxa,
in the large reptile tree (LRT, 1766 taxa; subset Fig. 2), pre-tetrapods with flat morphologies and small fins, like Panderichthys (Fig. 3) had only four finger buds. Strongly similar taxa, like the late-surviving basalmost tetrapod, Trypanognathus, likewise had a flat morphology, small limbs and only four fingers. This finger number is typical of basal tetrapods until the advent of finger 5 in five unrelated clades (Fig. 2) leaving no living descendants, except for reptilomorphs, beginning with Tulerpeton, Utegenia and kin, the clade that ultimately evolved reptiles, mammals, primates and humans.

Figure 4. Subset of the LRT focusing on basal tetrapods. Colors indicate number of fingers known. Many taxa do not preserve manual digits.

Figure 2. Subset of the LRT focusing on basal tetrapods. Colors indicate number of fingers known. Many taxa do not preserve manual digits.

Figure 6. Dorsal and ventral views of Panderichthys and several basal tetrapods demonstrating the low, flat skulls and bodies with small limbs and relatively straight ribs.

Figure 3. Dorsal and ventral views of Panderichthys and several basal tetrapods demonstrating the low, flat skulls and bodies with small limbs and relatively straight ribs.

Adding taxa 
resolves this problem and many others. Excluding taxa only perpetuates traditional myths and hobbles present research.


References
Dickson BV, Clack JA, Smithson TR et al. 2020. Functional adaptive landscapes predict terrestrial capacity at the origin of limbs. Nature (2020). https://doi.org/10.1038/s41586-020-2974-5

Woltering et al. 2020 study genes to elucidate finger origins

Woltering et al. 2020
attempted to elucidate the transition from fins to fingers by studying the genes of extant lungfish, which don’t have fingers and their ancestors never had fingers.

From the abstract
“How the hand and digits originated from fish fins during the Devonian fin-to-limb transition remains unsolved.

No. The large reptile tree (LRT; subset Fig. 1) solved that problem in 2019 following the work of Boisvert, Mark-Kurik and Ahlberg 2008. These authors found four finger buds on Panderichthys. Thereafter four fingers appear on all basalmost tetrapods in the LRT, like Trypanognathus (Fig. 2), a taxon found in Carboniferous strata with Middle Devonian origins. Taxon exclusion is once again the problem here.

Late Devonian taxa with supernumerary digits, like Acanthostega and Ichthyostega, are the traditional ‘go-to’ taxa for the fin-to-finger transition. That was supplanted in 2019 by phylogenetic analysis in the LRT (subset Fig. 1). Simply adding taxa recovers Acanthostega and Ichythyostega as terminal taxa. They have more derived skulls and bodies sporting larger limbs and more digits.

Figure 4. Subset of the LRT focusing on basal tetrapods. Colors indicate number of fingers known. Many taxa do not preserve manual digits.

Figure 4. Subset of the LRT focusing on basal tetrapods. Colors indicate number of fingers known. Many taxa do not preserve manual digits.

Woltering et al. 2020 report,
“Controversy in this conundrum stems from the scarcity of ontogenetic data from extant lobe-finned fishes. We report the patterning of an autopod-like domain by hoxa13 during fin development of the Australian lungfish, the most closely related extant fish relative of tetrapods.”

In other words, Woltering et al. looked at genes in lungfish that never had digits.

Figure 6. Dorsal and ventral views of Panderichthys and several basal tetrapods demonstrating the low, flat skulls and bodies with small limbs and relatively straight ribs.

Figure 2. Dorsal and ventral views of Panderichthys and several basal tetrapods demonstrating the low, flat skulls and bodies with small limbs and relatively straight ribs at the fin-to-finger transition. Acanthostega and Ichythyostega have more derived bodies with larger limbs and more digits.

Why study lungfish
when we have fossil taxa (Fig. 3) in the lineage of tetrapods? Why study genes when genomic studies produce false positives in deep time? Taken together the Woltering et al. study seems like a waste of effort on both fronts, but they didn’t realize this at the time. Paleontologists love genomics like Isaac Newton loved alchemy.

Figure 3. Forelimb of several basal tetrapods rearranged to more closely fit the LRT. Four fingers turns out to be the primitive number. Five is a recent mutation. Six was a short-lived experiment in Tulerpeton.

Figure 3. Forelimb of several basal tetrapods rearranged to more closely fit the LRT. Four fingers turns out to be the primitive number. Five is a recent mutation. Six was a short-lived experiment in Tulerpeton.

Woltering et al. 2020 report,”
“Differences from tetrapod limbs include the absence of digit-specific expansion of hoxd13 and hand2 and distal limitation of alx4 and pax9, which potentially evolved through an enhanced response to shh signaling in limbs. These developmental patterns indicate that the digit program originated in postaxial fin radials and later expanded anteriorly inside of a preexisting autopod-like domain during the evolution of limbs. Our findings provide a genetic framework for the transition of fins into limbs that supports the significance of classical models proposing a bending of the tetrapod metapterygial axis.”

Be wary of genetic studies over deep time. They have been shown to deliver false positives way too often to be trusted, or even attempted. Fossils and phenomic studies are better in all respects because they recover cladograms in which all taxa demonstrate a gradual accumulation of derived traits.


References
Boisvert CA, Mark-Kurik E and Ahlberg PE 2008.
 The pectoral fin of Panderichthys and the origin of digits. Nature 456:636–638.
Woltering JM et al. (5-co-authors) 2020. Sarcopterygian fin ontogeny elucidates the origin of hands with digits. Science Advances 6(34): eabc3510 DOI: 10.1126/sciadv.abc3510
https://advances.sciencemag.org/content/6/34/eabc3510

Metoposaurus gets one more finger

Konietko-Meier et al. 2020 discover digit 5
where they did not expect to find one, on Metoposaurus (Figs. 1-3).

Figure 4. Ozimek hitching a ride on top of Metoposaurus.

Figure 1. Ozimek hitching a ride on top of Metoposaurus. Note the relatively large manus and pes here compared to figure 2.

We’ve long wondered, how many fingers did the first tetrapod have? 
If more than five, when did four or five come to be?
If five, when did four or more than five come to be?
If four, when did five or more than five come to be?

Figure 3. Metoposaurus in several views.

Figure 2. Metoposaurus in several views. Smaller hands and feet on this data lacking digit 5.

From the Konietko-Meier et al. 2020 introduction:
“In contrast to crown tetrapods that rarely have more than five digits, basal tetrapod groups possessed more digits, such as Acanthostega gunnari Jarvik, 1952 which had eight in the forelimb (Coates and Clack, 1990) and Ichthyostega Säve-Söderbergh, 1932 with seven digits in the hindlimb (Säve-Söderbergh, 1932; Jarvik,1996). This fact indicates that polydactyly is the plesiomorphic condition for the tetrapod autopodium (Laurin et al., 2000).”

No. That’s a myth. The large reptile tree (LRT, 1713+ taxa; subset Fig. 4) recovered four fingers in basal tetrapods. Five fingers are derived in several convergent clades. More than five fingers occurs only in Acanthostega and kin, a derived clade without descendants. (Maybe Ichthyostega, too, but we have no hands for it).

Figure 2. Metoposaurus manus with five digits from Konietko-Meier et al. 2020. Colors and PILs added here.

Figure 3. Metoposaurus manus with five digits from Konietko-Meier et al. 2020. Colors and PILs added here. Note the foreshortening of the distal phalanges somewhat corrected here and in diagram at right. Not sure why p5.1 is so long in the diagram.

From the Konietko-Meier et al. 2020 abstract:
“Temnospondyli are commonly believed to have possessed four digits in the manus
and five in the pes. However, actual finds of articulated autopodia are extremely rare. The most important observation is the presence of five metacarpals in this specimen. This allows reconstructing the manus as pentadactyl.”

Figure 4. Subset of the LRT focusing on basal tetrapods. Colors indicate number of fingers known. Many taxa do not preserve manual digits.

Figure 4. Subset of the LRT focusing on basal tetrapods. Colors indicate number of fingers known. Many taxa do not preserve manual digits.

From the Konietko-Meier et al. introduction:
“The first known record of a pentadactyl hand belongs to the Early Carboniferous stegocephalian Casineria kiddi (Paton et al., 1999).”

Chronology does not always mirror phylogeny. Casineria nests as an archosauromorph reptile, off the bottom of the chart (Fig. 4). Many more primitive taxa had only five digits. The Late Devonian reptilomorph, Tulerpeton, had only five fingers, as we learned earlier.

Among temnospondyls in the LRT
(Fig. 4) the derived taxa leaving no descendants, Parotosuchus and Paracyclotosaurus, were illustrated with five fingers. Trematosaurus is known from skull material only. These fifth fingers are appearing de novo, not as reversals.

Proterogyrinus
developed five fingers. Fingers are not preserved in related taxa, none of which left descendants.

Dissorophids
developed five fingers without leaving descendants.

Reptilomorpha,
starting with Utegenia + Seymouriamorpha, developed five fingers and we are their descendants.

The Konietko-Meier et al. chart
(their Fig. 4) indicates the outgroup taxon, Greererpeton (Fig. 5; Godfrey 1986, 1989 had five fingers.

This is an error. Only the PhD thesis illustrates fingers and only four are illustrated (Fig. 5). Maybe the five-digit pes was accidentally added to the manus database?

Figure 5. Data for Greererpeton from Godfrey 1986.

Figure 5. Data for Greererpeton from Godfrey 1986. Only the pes has five digits.

From the Konietko-Meier et al. introduction:
“Reconstruction of the evolution of digit reduction of the most basal and post-Devonian stegocephalians is not possible because of the lack of informative fossils. It is known that reductions in the number of digits have occurred frequently during tetrapod evolution, but it is still not known exactly when or even how many times the number of digits was reduced to five or less (Laurin et al., 2000).”

The LRT clarifies this problem. Reductions in the number of digits occurred less frequently than envisioned by Konietko-Meier et al. since ‘four fingers’ is the primitive and plesiomorphic condition, even in Greererpeton.


References
Godfrey SJ 1989. The postcranial skeletal anatomy of the Carboniferous tetrapod Greererpeton burkemorani Romer, 1969. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences, 323(1213), 75–133.
Konietzko-Meier D, Teschner EM, Bodzioch A and Sander PM 2020. Pentadactyl manus of the Metoposaurus krasiejowensis from the Late Triassic of Poland, the first record of pentadactyly among Temnospondyli. Journal of Anatomy 00:1–11. DOI: 10.1111/joa.13276

How bat feet turn laterally, then upside-down

Bats are inverted bipeds.
They hang by branches and cave walls by their feet. While inverted, bat forelimbs are folded away until needed for flight. The hind limbs frame membranes linking the laterally oriented legs to the medial tail. We looked at the origin of bats from non-volant ancestors earlier here, here and at several earlier links therein (also see Fig. 3).

Figure 1. Hind limbs and closeup of ankle of Cynopterus, an extant micro bat, from Digimorph.org. Colors and diagram elements added here.

Figure 1. Hind limbs and closeup of ankle of Cynopterus, an extant micro bat, from Digimorph.org. Colors and diagram elements added here. Unlike most mammals, the knees are often above the hips in bats.

Bat experts know this, but it  comes as news to me.
A closer examination of bat hindquarters (Fig. 1) reveals two axial twists that add up to a ~180º rotated hind limb for the micro bat Cynopterus. The ankle is capable of additional rotation.

  1. The acetabulum axially rotates ~90º from ventrolateral to dorsolateral.
  2. The femur axially rotates so the distal end is ~90º rotated from the proximal head (Fig. 2).
  3. The tarsal centralia also rotate upon the tibiale (Fig. 1).
Figure 2. Bat femur animated to show untwisted typical mammal orientation of femoral head.

Figure 2. Two views of a bat femur animated to show typical untwisted orientation of femoral head as found in most mammals.

Axial torsion in proximal bones ultimately produces a pes
that is dorsal side up in flight in derived extant bats. Based on these twists, bat knees appear to bend backwards compared to other mammals.

Figure 3. The basal bat, Onychonycteris.

Figure 3. The basal bat, Onychonycteris. The feet are smaller and the hind limbs are more gracile primitively, like those of the bat precursor, Chriacus in figure 3.

In the transitional basal bat
Onychonycteris, the hind limb appears to be laterally oriented with long gracile hind limbs and the dorsal side of the tiny pes likewise oriented laterally. If you think such tiny feet seem less capable of inverted clinging compared to the relatively big feet of Cynopterus (Fig. 1), you’re being observant. But long legs and small feet are primitive for bats. So is inverted bipedal hanging. What you’re seeing is a transitional phase.

Figure 1. Hypothetical bat ancestors arising from a sister to Chriacus, which may be a large late survivor of a smaller common ancestor.

Figure 4. Hypothetical bat ancestors arising from a sister to Chriacus, which may be a large late survivor of a smaller common ancestor. Imagine stem bat 3 and Onychonycteris pinching the branch they hang from with long legs acting like pliers, an idea that did not occur to me years ago when this was illustrated.

Rather than clinging to the same side of the twig (or cave wall)
the long legs and small feet of Onychonycteris acted more like tongs or pliers, pinching both sides of a branch medially between them. We also see this in primitive micro bats, like long-tailed Rhinopoma and primitive megabats, like Balionycteris (Fig. 5).

Figure 5. Balionycteris hanging from both sides of a slender branch by laterally-twisted feet.

Figure 5. Balionycteris, the smallest megabat, hanging from both sides of a slender branch by laterally-twisted small feet.

Is there any new process on the bat pelvis that facilitates such adduction?
Yes. The pubis often develops a bump or rod, a prepubic process, analogous to the prepubis in pterosaurs. This process anchors muscles of femoral adduction.

Colugos and pangolins
also hang inverted from branches like that, with feet on both sides of a supporting branch.

The outgroup to bats in the LRT,
Chriacus, (Fig. 4) does not preserve evidence of long bone axial torsion (the mid-portion of the femur is not preserved). The acetabulum does not open dorsally.

Hanging upside-down
is something nearly all small arboreal mammals (e.g. squirrels, tree shrews, monkeys, tree opossums) can do facilitated by a flexible ankle that ensures the claws attach to the bark at any angle. Only bats and their immediate ancestors had such a firm toe grip while inverted they no longer needed their hands to grip. That freed the forelimbs to evolve into infant nurseries and parachute-like wings, not quite like those of birds and pterosaurs (Fig. 5), which were bipedal the conventional way: right side-up.

Figure 6. Click to enlarge. The origin of the pterosaur wing and the migration of the pteroid and preaxial carpal. A. Sphenodon. B. Huehuecuetzpalli. C. Cosesaurus. D. Sharovipteryx. E. Longisquama. F-H. Bergamodactylus.

The axial rotation of long limb bones is rare in tetrapods,
but it also can be seen in metacarpal 4 of Sharovipteryx, Longisquama and basal pterosaurs like Late Triassic Bergamodactylus (Fig. 6). That twist facilitates wing (finger 4) folding in the lateral plane of the wing (Peters 2002) rather than against the palm as in other tetrapods, including bats. Apparently the storage of long wings was just as important as the evolution of the long wings themselves in all volant tetrapods.


References
Peters D 2002. A New Model for the Evolution of the Pterosaur Wing – with a twist. – Historical Biology 15: 277–301.
Simmon NB, Seymour KL, Habersetzer J, Gunnell GF 2008. Primitive Early Eocene bat from Wyoming and the evolution of flight and echolocation. Nature 451 (7180): 818–21. doi:10.1038/nature06549. PMID 18270539.

wiki/Onychonycteris

 

Perhaps Tulerpeton had only 5 fingers (and five toes)

Let’s get right to it.
Tulerpeton (Fig. 1) was originally described with six fingers. If not six fingers, where did that sixth finger come from?

The other hand.
Specifically, the tip of finger 4 from the left hand (Fig. 1) provides a suitable match.  The left hand is otherwise buried in the matrix beneath the well-exposed right hand.

Figure 1. Tulerpeton manus with digit 6 re-identified as the top of digit 4 from the other hand.

Figure 1. Tulerpeton manus with digit 6 re-identified as the top of digit 4 from the other hand. The drawing at left is the in situ presentation. The diagram at right is the traditional six-finger interpretation. The manus in the middle represents the new hypothesis of digit identity.

Tulerpeton sisters in the LRT
don’t have a digit 6. So, maybe the original description was a mistake.

Likewise, the pes of Tulerpeton
was also originally described with six digits (Fig. 2). However, a new interpretation first discussed here indicated only five toes were present. That sixth digit was created to fill a perceived space produced by broken and displaced phalanges.

Figure 1. Tulerpeton pes reconstruction options using published images of the in situ fossil.

Figure 2. Tulerpeton pes reconstruction options using published images of the in situ fossil.

References
Coates MI and Ruta M 2001 2002. Fins to limbs: What the fossils say. Evolution & Development 4(5): 390–401.
Lebedev OA 1984. The first find of a Devonian tetrapod in USSR. Doklady Akad. Navk. SSSR. 278: 1407–1413.
Lebedev OA and Clack JA 1993. Upper Devonian tetrapods from Andreyeva, Tula Region, Russia. Paleontology36: 721-734.
Lebedev OA and Coates MI 1995. postcranial skeleton of the Devonian tetrapod Tulerpeton curtum Lebedev. Zoological Journal of the Linnean Society. 114 (3): 307–348.
Mondéjar-Fernandez J, Clément G and Sanchez S 2014. New insights into the scales of the Devonian tetrapods Tulerpeton curtum Lebedeve, 1984. Journal of Vertebrate Paleontology 34:1454-1459.

wiki/Tulerpeton

 

In memoriam: Professor Jennifer Clack

If you never met her,
here’s your second chance, via YouTube videos.

This week marks the passing of Professor Jennifer Clack (1947-2020),
a renown specialist in Devonian tetrapods, especially Acanthostega (Fig. 1). In the above 4-minute YouTube video from 2017, Clack introduces her concept that the first tetrapods, like her discovery of Acanthostega, had more than five manual digits. This is confirmed by Middle Devonian tetrapod tracks (Fig. 3) with more than five digits.

Figure 4. Acanthostega does not have much of a neck.

Figure 1. Acanthostega does not have much of a neck. Note the narrow torso, taller than wide, distinct from lobefin fish that phylogenetically led to basal tetrapods, like Trypanognathus in figure 4.

But not
according to the large reptile tree (LRT) which recovers Acanthostega as a terminal taxon, not a transitional one, far from the main line of tetrapod origins. Four digits are found in Panderichthys, Greererpeton and many other basal tetrapods, as we learned earlier here, here and here. More than five digits are found in only a few derived taxa, including the stem reptile, Tulerpeton, far from the origin of digits.

A more complete and technical account
of basal tetrapod traits is provided by Clack in this 20-minute YouTube lecture video from 2016 (above).

It may be that Clack only saw evolutionary progress
without considering the possibility of evolutionary reversal, as happens when taxa return to a more aquatic niche from a less aquatic niche, reducing the importance of their digits and limbs. In the above video, Clack does not provide a phylogenetic analysis, like the LRT (subset Fig. 2) that includes more primitive, but late-surviving basal tetrapods, all of which follow the pattern of a wider than deep torso, as in ancestral fish with embedded arm bones in their lobefins. Rather, she concentrates on individual traits, which while valuable, set her up for ‘Pulling a Larry Martin‘, rather than concentrating efforts on determining a phylogeny that minimizes taxon exclusion and lets the software determine (= mirror) evolutionary events, as the LRT does while minimizing taxon inclusion bias.

Figure 4a. Subset of the LRT focusing on basal tetrapods. Note the displaced positions of Acanthostega and Ichthyostega.

Figure 2. Subset of the LRT focusing on basal tetrapods. Note the displaced positions of Acanthostega and Ichthyostega.

Only after a phylogeny is documented and validated
can one then discuss the various traits and their uses by the creature that possessed them.

Lest we forget
the first tetrapod tracks (Fig. 1, Niedźwiedzki et al. 2010) predate fossil tetrapods, including Acanthostega, by 20 to 30 million years, as we looked at here. And even they had more than five toes. Thus the phylogenetic origin of tetrapods goes back even further. The early Devonian must have provided quite a few niches for such rapid evolution to take place.

Figure 3. Best Devonian Valentia track with various overlays.

Figure 3. Best Devonian Valentia track with various overlays.

We need to look more closely at
Trypanognathus (Fig. 4; latest Carboniferous), which is the most primitive, but by far not the earliest, taxon in the LRT to document fingers and limbs, rather than lobe fins. Note the anterior eyes, wide flat skull and body, and primitive sprawling limbs. Can someone count the fingers and toes on this specimen? I find no more than four digits. Some may be hiding here.

Figure 1. Trypanognathus in situ, colorized to bring out ribs and limbs.

Figure 4. Trypanognathus in situ, colorized to bring out ribs and limbs is the most primitive, but not the earliest taxon with limbs and toes, not lobe fins.

We’ve seen the chronology of several fossil finds
at odds with their phylogeny in the LRT (e.g. multituberculates, bats, Gregorius). That keeps it interesting, but only a wide gamut phylogenetic analysis based on traits will deliver a valid tree topology. As time goes by and more discoveries are made the competing hypotheses will someday converge.

Figure 2. Silvanerpeton from the Upper Viséan (331 mya) is the outgroup taxon for Gephyrostegus and the Amniota.

Figure 5. Silvanerpeton from the Upper Viséan (331 mya) is the outgroup taxon for Gephyrostegus and the Amniota.

And one more thing,
Clack 1994 described Silvanerpeton (Fig. 5, Viséan, 335 mya) first as an anthrcosauroid and later (Ruta and Clack 2006) as a stem tetrapod, all without recovering it as the basalmost reptile, as shown in the LRT. Adding taxa, creating a wider gamut phylogenetic analysis, would have brought even more fame to this well-respected paleontologist.


References
Clack JA 1994. Silvanerpeton miripedes, a new anthracosauroid from the Visean of East Kirkton, West Lothian, Scotland. Transactions of the Royal Society of Edinburgh: Earth Sciences 84 (for 1993), 369–76.
Niedźwiedzki G, Szrek P, Narkiewicz K, Narkiewicz M and Ahlberg PE 2010. Tetrapod trackways from the early Middle Devonian period of Poland Nature 463, 43-48. doi:10.1038/nature08623
Ruta M and Clack, JA 2006 A review of Silvanerpeton miripedes, a stem amniote from the Lower Carboniferous of East Kirkton, West Lothian, Scotland. Transactions of the Royal Society of Edinburgh: Earth Sciences, 97, 31-63.

https://www.zoo.cam.ac.uk/news/professor-jenny-clack-frs-1947-2020

http://www.theclacks.org.uk/jac/Biography.html

https://www.pbs.org/wgbh/nova/link/clack.html
(make sure to click on the parts 2-4 links therein)

 

Late Devonian origin of four-fingered hand revisited and tweaked

Earlier we looked at the origin of fingers
in basal tetrapods (Fig. 1). The primitive number then, as now, was 4 fingers, 5 toes.

Figure 1. Graphing the presence of fingers and toes in basal tetrapods, updated today with the addition of 4 digits in Panderichthys.

Figure 1. Graphing the presence of fingers and toes in basal tetrapods, updated today with the addition of 4 digits in Panderichthys. Sharp-eyed readers will note the switching of Panderichthys with the Tiktaalik clade here.

Unfortunately,
I overlooked a paper (Boisvert, et al., 2008) that found four proto-digits in the lobefin of Panderichthys (Fig. 2) and provided good data for the Tiktaalik manus that I did not have. With those corrections, a quick review is in order.

Figure 1. From Boisert et al. 2008, colors added. This is their ordering for the evolution of manual digits. Compare to figure 2.

Figure 2. From Boisert et al. 2008, colors added. This is their ordering for the evolution of manual digits. Compare to figure 3 where Panderichthys and Tiktaalik switch places and several taxa are inserted transitional to Acanthostega.

For some reason, fingers are rarely preserved in basal tetrapods,
but most continue to have four. Proterogyrinus (Fig. 2) is an early exception with five. So is Acanthostega (Fig. 2) with eight. See chart above  (Fig. 1) for all tested taxa in the LRT.

Tradition holds that eight is a primitive number,
later reduced to five or four. The large reptile tree (LRT, 1590 taxa, subset Fig. 1) flips that around. Eight is a derived number on a terminal taxon (Acanthostega) leaving no descendants. The primitive number is four (subset Fig. 1).

Boisvert, Mark-Kurik and Ahlberg 2008 used a CT-scanner
to find four proto-digits on the manus of Panderichthys (Fig. 2) and compared those to the traditional basal tetrapod taxa: Eusthenopteron, Tiktaalik and Acanthostega. Note the big phylogenetic leap they show between Tiktaalik and Acanthostega. And note the apparent reversal in Tiktaalik as the metacarpal seems to revert to a ray. These problems are corrected in figure 3.

Figure 3. Forelimb of several basal tetrapods rearranged to more closely fit the LRT. Four fingers turns out to be the primitive number. Five is a recent mutation. Six was a short-lived experiment in Tulerpeton.

Figure 3. Forelimb of several basal tetrapods rearranged to more closely fit the LRT. Compare to figure 2. As discovered here earlier, four fingers turns out to be the primitive number. Five is a recent mutation. Six was a short-lived experiment in Tulerpeton.

The large reptile tree
(LRT, 1590 taxa) recovers a different topology (subset Fig. 1). In the LRT Acanthostega is not a transitional taxon, but an aberrant one returning to a more aquatic lifestyle and leaving no descendants. On the other hand, the four digits in Panderichthys are retained by a wide variety of basal tetrapods. The number jumps to five with the addition of a lateral digit in only a few taxa (Fig. 1). Importantly, in Utegenia (Fig. 4) we see the most primitive appearance of five digits in our lineage despite its late appearance in the fossil record. More derived, but earlier, Late Devonian Tulerpeton had six fingers representing a failed experiment leaving no descendants in the lineage of reptiles, represented by Silvanerpeton in the Early Carboniferous, also pre-dating Utegenia.

So frogs did not lose a finger.
They retained the four that Panderichthys provided them. Four, not five or eight, is the primitive number of digits for basal tetrapods, as discovered earlier here in the LRT. Let me know if there was an earlier discovery for this hypothesis of interrelationships and I will promote that citation.

Occasionally
a salamander will have six fingers. We’ll look at that strange case soon.

Figure 5. Utegenia diagram showing five fingers on each hand. This is the most primitive taxon in our lineage to have all five.

Figure 4. Utegenia diagram showing five fingers on each hand. This is the most primitive taxon in our lineage to have all five.

In the course of this study
I learned that the Tiktaalik clade and Panderichthys needed to switch places on the LRT. This has been updated in most cases (Fig.1).

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

Figure 5. Tulerpeton manus and pes in situ, reconstructed by Lebdev and Coates 1995 and newly reconstructed here.

Evidently Boisvert et al. were using
an outdated tree topology and did not recognize the problem that arose between the digits presented by Tiktaalik and those presented by Panderichthys (Fig. 2). Of course that puts Tiktaalik cousins, Koilops and Spathicephalus in the lobefin grade (Fig. 1), lacking fingers. Both currently lack post-cranial data, but were originally thought to be tetrapods.


References
Boisvert CA, Mark-Kurik E and Ahlberg PE 2008. The pectoral fin of Panderichthys and the origin of digits. Nature 456:636–638.