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Ontogenetic bone growth in the caecilian skull

Monday, February 27, 2017 14:41
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(Before It's News)

Back to an old subject…
Earlier we looked at the skull of Dermophis, an extant caecilian from Mexico (Fig. 1) based on Digimorph.org images. There were comments from anamniote experts criticizing my labeling of the bones, suggesting I had a ‘magic fusion detector.’ I was encouraged to check out Wake and Hanken 1982, which documents the growth of the Dermophis skull (Fig. 2).

Figure 1. Dermophis, the extant Mexican caecilian, with bones, even if fused to one another, identified. The quadratojugal and squamosal are absent. Black and white image from Digimorph.org. Coloring the bones makes them so much easier to read and understand. ” data-medium-file=”https://pterosaurheresies.files.wordpress.com/2017/02/dermophis_5881.jpg?w=584&h=1022?w=171″ data-large-file=”https://pterosaurheresies.files.wordpress.com/2017/02/dermophis_5881.jpg?w=584&h=1022?w=584″ class=”size-full wp-image-26113″ src=”https://pterosaurheresies.files.wordpress.com/2017/02/dermophis_5881.jpg?w=584&h=1022″ alt=”Figure 1. Dermophis, the extant Mexican caecilian, with bones, even if fused to one another, identified. The quadratojugal and squamosal are absent. Black and white image from Digimorph.org. Coloring the bones makes them so much easier to read and understand.” width=”584″ height=”1022″ srcset=”https://pterosaurheresies.files.wordpress.com/2017/02/dermophis_5881.jpg?w=584&h=1022 584w, https://pterosaurheresies.files.wordpress.com/2017/02/dermophis_5881.jpg?w=86&h=150 86w, https://pterosaurheresies.files.wordpress.com/2017/02/dermophis_5881.jpg?w=171&h=300 171w, https://pterosaurheresies.files.wordpress.com/2017/02/dermophis_5881.jpg 588w” sizes=”(max-width: 584px) 100vw, 584px” />

Figure 1. Dermophis, the extant Mexican caecilian, with bones, even if fused to one another, identified. The quadratojugal and squamosal are absent. Coloring the bones makes them so much easier to read and understand. Skull from Digimorph.org and used with permission.

Wake and Hanken discuss
some of the earlier hypotheses regarding the origin of the skull bones in caecilians. “The belief of Marcus et al, (’35) that the well-developed skull of caecilians is a retained primitive feature has been challenged by many authors, however, all of whom interpret the stegokrotaphy of the caecilian skull as being secondarily derived from a reduced skull typical of other Recent amphibians.”

Unfortunately for Wake and Hanken,
the publication of Eocaecilia (Jenkins and Walsh 1993; Eaerly Jurassic, 190 mya) came eleven years later. That settled the issue.

Figure 1. Dermophis skull elements according to Wake and Hanken 1982. ” data-medium-file=”https://pterosaurheresies.files.wordpress.com/2017/02/dermophis-skull-growth.gif?w=584&h=529?w=300″ data-large-file=”https://pterosaurheresies.files.wordpress.com/2017/02/dermophis-skull-growth.gif?w=584&h=529?w=584″ class=”wp-image-26245 size-full” src=”https://pterosaurheresies.files.wordpress.com/2017/02/dermophis-skull-growth.gif?w=584&h=529″ alt=”Figure 1. Dermophis skull elements according to Wake and Hanken 1982. ” width=”584″ height=”529″ />

Figure 2. Dermophis skull elements according to Wake and Hanken 1982. Two of the larger growth series specimens  are shown here,  Red = pterygoid/quadrate. Also shown are the source of the fused bones based on phylogenetic relationship to Acherontiscus. Note the green ellipse = supratemporal, as in Eocaecilia.

Eocaecilia retains
the supratemporal and postfrontal, two bones thought by Wake and Hanken to have been absent in recent amphibians including caecilians. However, the elliptical supratemporal and the strip-like postfrontal both become temporarily visible in the 6.85 cm immature skull and then become fused to what Wake and Hanken label the squamosal. Their squamosal encircles the tiny orbit. Squamosals usually do not do that on their own, as everyone familiar with tetrapods knows. It doesn’t even contact the squamosal in Eocaecilia.

Figure 1. Eocaecilia skull with original and new bone identifications based on comparisons to sister taxa listed here. Like Brachydectes, the jaw joint has moved forward, beneath the jugal now fused to the quadratojugal creating a long retroarticular process, otherwise rare in amphibians. Also rare is the fusion of the squamosal with the postorbital. ” data-medium-file=”https://pterosaurheresies.files.wordpress.com/2017/02/eocaecilia_skull5882.jpg?w=584&h=363?w=300″ data-large-file=”https://pterosaurheresies.files.wordpress.com/2017/02/eocaecilia_skull5882.jpg?w=584&h=363?w=584″ class=”size-full wp-image-26115″ src=”https://pterosaurheresies.files.wordpress.com/2017/02/eocaecilia_skull5882.jpg?w=584&h=363″ alt=”Figure 1. Eocaecilia skull with original and new bone identifications based on comparisons to sister taxa listed here. Like Brachydectes, the jaw joint has moved forward, beneath the jugal now fused to the quadratojugal creating a long retroarticular process, otherwise rare in amphibians. Also rare is the fusion of the squamosal with the postorbital.” width=”584″ height=”363″ srcset=”https://pterosaurheresies.files.wordpress.com/2017/02/eocaecilia_skull5882.jpg?w=584&h=363 584w, https://pterosaurheresies.files.wordpress.com/2017/02/eocaecilia_skull5882.jpg?w=150&h=93 150w, https://pterosaurheresies.files.wordpress.com/2017/02/eocaecilia_skull5882.jpg?w=300&h=186 300w, https://pterosaurheresies.files.wordpress.com/2017/02/eocaecilia_skull5882.jpg 588w” sizes=”(max-width: 584px) 100vw, 584px” />

Figure 3. Eocaecilia skull with original and new bone identifications based on comparisons to sister taxa listed here. Like Brachydectes, the jaw joint has moved forward, beneath the jugal now fused to the quadratojugal creating a long retroarticular process, otherwise rare in amphibians. Also rare is the fusion of the squamosal with the postorbital.

Wake and Hanken reported:
“Our analysis of skull development in Dermophis has several implications for this controversy. First, as presented above, we did not observe several of the embryonic ossification centers whose supposed presence has been used to ally caecilians and early amphibians, particularly the microsaurs.” Again, they did not have the blueprint of Eocaecilia to work with, as we do now. They did not mention the microsaur, Acherontiscus (Carroll 1969; Namurian, Carboniferous; Fig. 4), in their paper. This taxon phylogenetically and chronologically precedes caecilians in the large reptile tree (LRT). Microbrachis is also related, but has a shorter torso and longer legs than Acherontiscus and Eocaecilia.

Figure 4. Acherotisicus has large cheek bones (squamosal, quadratojugal) that appear to fuse in Eocaecilia and Dermophis. ” data-medium-file=”https://pterosaurheresies.files.wordpress.com/2017/02/acherontiscus_588.jpg?w=584&h=696?w=252″ data-large-file=”https://pterosaurheresies.files.wordpress.com/2017/02/acherontiscus_588.jpg?w=584&h=696?w=584″ class=”size-full wp-image-26210″ src=”https://pterosaurheresies.files.wordpress.com/2017/02/acherontiscus_588.jpg?w=584&h=696″ alt=”Figure 4. Acherotisicus has large cheek bones (squamosal, quadratojugal) that appear to fuse in Eocaecilia and Dermophis.” width=”584″ height=”696″ srcset=”https://pterosaurheresies.files.wordpress.com/2017/02/acherontiscus_588.jpg?w=584&h=696 584w, https://pterosaurheresies.files.wordpress.com/2017/02/acherontiscus_588.jpg?w=126&h=150 126w, https://pterosaurheresies.files.wordpress.com/2017/02/acherontiscus_588.jpg?w=252&h=300 252w, https://pterosaurheresies.files.wordpress.com/2017/02/acherontiscus_588.jpg 588w” sizes=”(max-width: 584px) 100vw, 584px” />

Figure 4. Acherotisicus has large cheek bones (squamosal, quadratojugal) that appear to fuse in Eocaecilia and Dermophis.

Earlier I used the term bone ‘buds’
to represent small ossification centers from which the adult skull bone would eventually develop. This term caught some flak, but as you can see (Fig. 2) the adult skull bones do indeed develop from smaller ‘buds’.

Wake and Hanken concluded:
“We heartily concur with the idea of a long and separate evolutionary history for caecilians, independent of frogs and salamanders, as has been expressed by Carroll and Currie (’75). However, the resemblances between the cranial morphology of caecilians and that of their purported ancestors, the microsaurs, are only superficial, and many significant differences remain. Further, there are real differences in the postcranial elements, which were not within the purview of Carroll and Currie’s study. Based on our observations of skull development in Dermophis mexicanus, we believe that there is now little evidence for the hypothesis of primary derivation of the caecilian skull from any known early amphibian group.”

So Wake and Hanken gave up —
but this was before the advent of widespread computer-aided phylogenetic analysis, Now, like flak itself, you don’t have to actually hit a target. You can get really close and still knock it down. So ‘superficial’ resemblances, if nothing else in the gamut of included taxa comes closer, become homologies. That’s what happens in the LRT.

Based on what Wake and Hanken 1982 wrote,
skull buds are not apparent. Based on what Wake and Hanken 1982 traced, skull buds for all pertinent bones are indeed present.

And caecilians are cemented down
as living microsaurs close to Eocaecilia, Acherontiscus and Microbrachis based on morphology, phylogeny and ontogeny.

References
Jenkins FA, Walsh DM and Carroll RL 2007. Anatomy of Eocaecilia micropodia, a limbed caecilian of the Early Jurassic. Bulletin of the Museum of Comparative Zoology 158(6): 285-366.
Jenkins FA and Walsh M 1993. 
An Early Jurassic caecilian with limbs. Nature 365: 246–250.
Marcus H, Stimmelmayr E and Porsch G 1935. Beitrage zur Kenntnis der Gymnophionen. XXV. Die Ossifikation des Hypogeophisschddels. Morphol. Jahrb. 76;375-420.
Wake MH and Hanken J 1982. Development of the Skull of Dermophis mexicanus (Amphibia: Gymnophiona), With Comments on Skull Kinesis and Amphibian Relationships. Journal of Morphology 173:203-222.



Source: https://pterosaurheresies.wordpress.com/2017/02/28/ontogenetic-bone-growth-in-the-caecilian-skull/

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