The Fin-to-Finger Transition in one go
Traditional hypotheses
for the origin of tetrapods (= vertebrates with digits) focused on late Devonian Panderichthys and Elpistostege (Fig 1), two long-torso, wide-rib, small-lobefin taxa on the primitive side of the fin-to-finger transition. On the derived side were Latest Devonian Ichthyostega and Acanthostega (Fig 2), two short-torso, round-rib, large limb, multi-dactyl taxa. These two are also the oldest known tetrapods – but an early Middle Devonian tetrapod trackmaker (Fig 4) was much older.
That’s the first problem.
Second problem:
Most other basal tetrapods had only four fingers and five toes (Fig 6). So it was traditionally assumed that the second wave of Carboniferous tetrapods lost ancestral superfluous digits (Fig 2).
Figure 4. Acanthostega does not have much of a neck.
” data-large-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2017/06/acanthostega588.jpg?w=584″ src=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2017/06/acanthostega588.jpg?w=584″ alt=”Figure 4. Acanthostega does not have much of a neck.” class=”wp-image-27173″ srcset=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2017/06/acanthostega588.jpg?w=584 584w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2017/06/acanthostega588.jpg?w=135 135w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2017/06/acanthostega588.jpg?w=271 271w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2017/06/acanthostega588.jpg 588w” sizes=”(max-width: 584px) 100vw, 584px” />
Third problem:
Some of the most primitive Carboniferous tetrapods are long-torso, wide-body, small limb taxa, (Fig 1) resembling Panderichthys and Tiktaalik, except for the exchange of small lobefins for small limbs with small digits.
Fourth problem:
Late Devonian Tulerpeton (Fig 3) had five slender fingers and five toes with a near-reptile formula of non-ungual phalanges 1-2-3-4-4 on the manus and similarly for the pes.
Figure 5. Manus/Fin diagram from Cloutier et al. The phylogenetic order is wrong and Tulerpeton is not related (in the LRT). Greererpeton is substituted. 4 fingers is the primitive number.
” data-large-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2020/03/cloutier_hand_diagram588.gif?w=584″ src=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2020/03/cloutier_hand_diagram588.gif?w=584″ alt=”Figure 5. Manus/Fin diagram from Cloutier et al. The phylogenetic order is wrong and Tulerpeton is not related (in the LRT). Greererpeton is substituted. 4 fingers is the primitive number.” class=”wp-image-43452″ srcset=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2020/03/cloutier_hand_diagram588.gif?w=584 584w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2020/03/cloutier_hand_diagram588.gif?w=150 150w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2020/03/cloutier_hand_diagram588.gif?w=300 300w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2020/03/cloutier_hand_diagram588.gif 588w” sizes=”(max-width: 584px) 100vw, 584px” />
Here the traditional fin-to-finger transition hypothesis is revised
after a phylogenetic analysis that omits fewer taxa. Here basalmost tetrapods, like Pholidogaster and Colosteus (Fig 5), had a long but round (in cross-section) scaly torso with reduced, slender limbs. Here four fingers is the primitive number.
Here five fingers appear in basal Reptilomorpha.
Here Tulerpeton represents an as yet undiscovered Devonian radiation of terrestrial tetrapods, nesting as the proximal outgroup to the Reptilia. Here multi-dactyl taxa with robust limbs (Fig 2) are derived and these few taxa left no Carboniferous descendants.
Introduction
Two short definitions are followed here: Fish = pre-tetrapods have fins and lack digits. Conversely tetrapods have fingers or toes or both. This might seem obvious, but the widely-accepted definition of a stem tetrapod includes vertebrates with lobe fins and the widely-accepted definition of a crown tetrapod excludes many taxa with fingers. There are no known taxa in which one set of lobes/limbs had fins while the other set had digits. Finally, skull-only sister taxa of basal tetrapods are here considered tetrapods ‘by association’.
The focus of this blogpost
is restricted to the one and only fin-to-finger transition recovered in the large reptile tree (LRT, 2340 taxa). The large number of taxa in the LRT minimizes taxon exclusion, an ongoing problem in vertebrate paleontology.
The cladogram
The LRT is the online cladogram recovered after the inclusion of 2340 taxa ranging from Cambrian pre-fish to humans, birds, turtles and long lists of taxa in-between. This experiment began in 2011 with 240 taxa and grew ten-fold in 15 years. 268 pterosaurs and their proximal outgroups are tested separately with a different set of characters in the adjoining large pterosaur tree (LPT). Originally all taxa were on the same matrix, but when the number of taxa reached the software limit of 1500, the matrices had to be divided into overlapping subsets that continued to add taxa. 238 multi-state characters are scored in the LRT. The heuristic search option is used. Resolution is generally good throughout. Taxa colored red in the LRT have been tested, then deleted, generally because they represent only fragments of a skeleton.
Gene studies
Results recovered in the LRT do not match suprageneric results in genomic studies, which cannot test extinct taxa. The reason for this discrepancy is not yet known, but is widely acknowledged.
Results
In the LRT the following taxa are in the ancestral line toward tetrapods: Middle and late Devonian Eusthenopteron, Osteolepis, Glyptolepis, Gooloogongia, Middle Devonian Tinirau and Early Carboniferous Koilops (a skull-only taxon). In the LRT little Koilops gave rise to tetrapods on one branch and on the other branch, long-torso, low-body, small-fin taxa like Tiktaalik, Panderichthys and Elpistostege.
The following Carboniferous taxa are in the basalmost tetrapod clade: Aytonerpeton, Colosteus, Deltaherpeton and Pholidogaster.
The following Carboniferous taxa nested at the bases of the several radiations the followed the appearance of digits: Trypanognathus, giant early Permian Gaiasia and early Carboniferous Greererpeton. Long, low Trypanognathus converges with long, low Panderichthys (with the exception of digits replacing fins). Ichthyostega and Acanthostega arose from the Greerepton clade in the LRT.
Figure 1. The Early Carboniferous limbed osteolepid, Pholidogaster, compared to Middle Devonian Zalchemie tracks to scale.
” data-large-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2018/10/pholidogaster-zalchemie_tracks588.jpg?w=584″ src=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2018/10/pholidogaster-zalchemie_tracks588.jpg?w=584″ alt=”Figure 1. The Early Carboniferous limbed osteolepid, Pholidogaster, compared to Middle Devonian Zalchemie tracks to scale.” class=”wp-image-33734″ srcset=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2018/10/pholidogaster-zalchemie_tracks588.jpg?w=584 584w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2018/10/pholidogaster-zalchemie_tracks588.jpg?w=150 150w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2018/10/pholidogaster-zalchemie_tracks588.jpg?w=300 300w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2018/10/pholidogaster-zalchemie_tracks588.jpg 588w” sizes=”(max-width: 584px) 100vw, 584px” />
Tetrapod trackmakers
Early Middle Devonian multi-dactyl tetrapod trackways that can be matched to a multi-dactyl ichthyostegid have been described (Niedźwiedzki et al 2010). So the evolution of fingers from fins preceded the early Middle Devonian.
Before the transition from fins to fingers
Eusthenopteron had no metacarpals. Tiktaalik had three tiny metacarpals, perhaps four, but that lateral area is today taphonomically damaged. Digits are not present. Panderichthys had four larger individually-shaped = dissimilar metacarpals but still lacked phalanges = digits..
Figure 6. Colosteus relatives according to the LRT scaled to a common skull length. Their sizes actually vary quite a bit, as noted by the different scale bars. Only Pholidogaster and Colosteus are taxa in common with traditional colosteid lists.
” data-large-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2017/06/colosteids_recon2-5881.jpg?w=584″ src=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2017/06/colosteids_recon2-5881.jpg?w=584″ alt=”Figure 6. Colosteus relatives according to the LRT scaled to a common skull length. Their sizes actually vary quite a bit, as noted by the different scale bars. Only Pholidogaster and Colosteus are taxa in common with traditional colosteid lists.” class=”wp-image-27196″ srcset=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2017/06/colosteids_recon2-5881.jpg?w=584 584w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2017/06/colosteids_recon2-5881.jpg?w=143 143w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2017/06/colosteids_recon2-5881.jpg?w=287 287w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2017/06/colosteids_recon2-5881.jpg 588w” sizes=”(max-width: 584px) 100vw, 584px” />
During the transition from fins to fingers
Colosteus had four small, slender, cylindrical radiating metacarpals and slender digits of laterally increasing length. These tiny ?vestiges were unable to support the weight of the cylindrical body and smaller than ancestral lobes = limbs sans fins.
Aytonerpeton preserved robust pedal digits bones.
Pholidogaster preserved a five-metatarsal pes with mt4 the most robust and digit 4 likely the longest. Distal phalanges and unguals are taphonomically missing. The tarsus had two proximal elements (tibiale and fibular) and three distal elements with a simple hinge joint between them. The robust crus was about half the length of the longer, more slender femur. The fibula was more robust than the tibia. The manus was taphonomically absent. Here again the torso was roughly cylindrical in cross-section, as in Middle Devonian Tinirau.
After the transition from fins to fingers
Trypanognathus had four ‘hourglass’ shaped metacarpals and four long proximal phalanges tipped by four unguals. Digit 3 was the largest. Phalanx 3.1 was the largest. Unguals 3 and 4 were the largest. The flattened wide body and small limbs were convergent with similar traits on members of the Tiktaalk clade.
Greererpeton (Fig 6) had four ‘hourglass’ shaped metacarpals (Fig 6) and four long proximal phalanges. The medial two were tipped by sharp-ish unguals. Digits 3 and 4 had an extra row of phalanges tipped by sharp-ish unguals with the formula 1, 1, 2, 2 (not counting the unguals). Digit 3 was barely the largest. Phalanx 3.1 was the largest. The pes had five digits tipped with sharpish unguals with the following non-ungual number: 1, 1, 2, 3, 3. The body was again flat and wide. The limbs were small and unable to support the weight of the torso. Greerepeton descendants include Ichthyostega (Fig 7) and Acanthostega in the LRT.
Figure 6. Not sure which is more correct, but this Ichthyostega data shows little to no wiggle room for the larger skull, more for the smaller skull.
” data-large-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2017/06/ichthyostega588.jpg?w=584″ src=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2017/06/ichthyostega588.jpg?w=584″ alt=”Figure 6. Not sure which is more correct, but this Ichthyostega data shows little to no wiggle room for the larger skull, more for the smaller skull.” class=”wp-image-27176″ srcset=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2017/06/ichthyostega588.jpg?w=584 584w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2017/06/ichthyostega588.jpg?w=150 150w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2017/06/ichthyostega588.jpg?w=297 297w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2017/06/ichthyostega588.jpg 588w” sizes=”(max-width: 584px) 100vw, 584px” />
Multi-dactyl taxa
Ichthyostega (Fig 7) does not preserve a manus. However, pedal digit 1 appears in triplicate, all narrower and shorter than the lateral four robust metatarsals and toes with the following non-ungual number: 2(x3), 2, 3, 3, 2. The pelvis, femur and radius were robust. The scapula and humerus were also robust. The derived torso was short and cylindrical, distinct from the the long, low, small-finned pre-tetrapods.
The manus of Acanthostega preserves three short straight digits (again in triplicate) where digit 1 is otherwise located in other taxa. Digit 2 is aligned with the axis of the antebrachium. Fingers 3–6 radiate laterally with digit 4 the longest. So digits 1, 2 and 6-8 appear to be novel in appearance after 4-fingered ancestral taxa. The pes had a vestige digit 1, a short digit 2, digits3-5 equally long, digit 6 shorter and digit 7 shorter still. Digit 8 was a vestige no longer than mt7. These traits are not found in Carboniferous taxa, indicating that Ichthyostega and Acanthostega were derived, not primitive, as they nest in the LRT some distance from the origin of digits.
The earliest and most derived Reptilomorph
Late DevonianTulerpeton nested at the base of the Reptilia = Amniota in the LRT. Viséan Silvanerpeton is the last common ancestor of all extant mammals, reptiles and birds. ‘Amniota’ is a junior synonym in the LRT. The right hand of Tulerpeton had only five fingers. The purported sixth finger was the emerging, but otherwise buried tip of the left fourth finger.
Speculation on the advantage of digits over lobe fins
Tiny digits tipping small, slender limbs in Colosteus and Pholidogaster (Fig 5) ancestors apparently deep in the Early Devonian must have provided some niche advantage not afforded to their lobe-fin ancestors and sister taxa. Primitive tiny limbs and relatively weak girdles would have been less able to lift the relatively large torso off the substrate, so the terrestrial option seems to be untenable – that is unless one considers the present activity of finless moray eels as they hunt for crabs on rocky shorelines. YouTube video here.
Compared to lobe fins, digit tips might have providedindividual points of sensory input, perhaps for navigation along shallow water bottoms cluttered with detritus. Digits can wrap around objects, preventing the rest of the body from drifting in weak currents. Thus digits could have been anchors, not paddles. Digits can cling to or lodge against the substrate when traversing through shallow waters. Perhaps limbs were useful during exaggerated sinusoidal motions of the torso that provided the mail thrust, as in moray eels.
References
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Source: https://pterosaurheresies.wordpress.com/2026/05/28/the-fin-to-finger-transition-in-one-go/
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