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Early Permian Vertebrates from the Cutler Formation of the ...

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C24 CONTRIBUTIONS TO PALEONTOLOGY<br />

depression on <strong>the</strong> anterior face <strong>of</strong> <strong>the</strong> conjoin.ed internal<br />

ridges <strong>of</strong> <strong>the</strong> pubes in Limnoscelops longifernur makes<br />

this face more nearly vertical than it is in Li1nnoscelis<br />

or Lim;nosceloides. In this, Lhnnoscelops is more like<br />

<strong>the</strong> captorhinids than are <strong>the</strong> o<strong>the</strong>r two genera.<br />

The external opening <strong>of</strong> <strong>the</strong> obturator foramen is<br />

similarly placed in Li1nnoscelops, Limnoscelis, and Limnosceloides.<br />

In Limnoscelis, as in Labidosaurus, <strong>the</strong><br />

internal opening <strong>of</strong> <strong>the</strong> foramen lies anterior to <strong>the</strong><br />

internal ridge <strong>of</strong> <strong>the</strong> pubis, but in Lirn/Jwrweloides, <strong>the</strong><br />

opening lies at <strong>the</strong> upper end <strong>of</strong> <strong>the</strong> ridge (Romer, 1952,<br />

p. 90). Limnoscelopslongifemu,r shows a somewhat intermediate<br />

condition in that <strong>the</strong> opening lies immediately<br />

anterior to a crest above <strong>the</strong> greatest depth <strong>of</strong><br />

<strong>the</strong> ridge, but <strong>the</strong> position <strong>of</strong> <strong>the</strong> opening is more like<br />

that in Lim;nosceloides than it is like that in Limnosceiis.<br />

The proximal fragment <strong>of</strong> an ulna is about fivesevenths<br />

<strong>the</strong> size <strong>of</strong> <strong>the</strong> corresponding part <strong>of</strong> Limnoscelis<br />

paludis figured by Williston (1912, fig. 27), but<br />

<strong>the</strong> two are morphologically indistinguishable.<br />

The hind led <strong>of</strong> Limnoscelops longifemur is represented<br />

in <strong>the</strong> type by only <strong>the</strong> proximal and distal ends<br />

<strong>of</strong> <strong>the</strong> left femur; <strong>the</strong>y are very similar to those <strong>of</strong><br />

Labidosaurus, but what is left <strong>of</strong> its stumps seems to<br />

indicate that <strong>the</strong> shaft was more slender and more like<br />

that <strong>of</strong> Oaptorhinus. The relative slenderness in Limnoscelops<br />

is a significant difference <strong>from</strong> Labidosaurus,<br />

a much smaller reptile by comparison <strong>of</strong> <strong>the</strong> proximal<br />

and distal ends. (See Case, 1911a, fig. 48.)<br />

The complete femur may have been at least 130 mm<br />

long <strong>from</strong> <strong>the</strong> most proximal point on <strong>the</strong> head to <strong>the</strong><br />

end <strong>of</strong> <strong>the</strong> posterior condyle, and at least 122 mm long<br />

<strong>from</strong> <strong>the</strong> most proximal point on <strong>the</strong> head to <strong>the</strong> center<br />

<strong>of</strong> <strong>the</strong> intercondylar notch. This calculated distance<br />

<strong>from</strong> head to intercondylar notch is about one and onefourth<br />

times <strong>the</strong> comparable length in Li'lnnosceloides<br />

dunkardensis (USNM 12166). The pelves are about<br />

<strong>the</strong> same size in <strong>the</strong>se two; <strong>the</strong>refore, Limnoscelops<br />

longifemur had proportionally much longer femora,<br />

and probably also had proportionally longer hind legs<br />

and slenderer shaft than Limtnosceloides dunkardensis:<br />

The stump <strong>of</strong> <strong>the</strong> shaft on <strong>the</strong> proximal fragment <strong>of</strong><br />

Limnoscelops longifernrur is about 17 mm thick ( measured<br />

in <strong>the</strong> plane <strong>of</strong> <strong>the</strong> head <strong>of</strong> <strong>the</strong> femur) as compared<br />

to about 25 mm in Limnosceloides dunkardensis<br />

(calculated <strong>from</strong> Romer, 1952, fig. 12). The proximal<br />

and distal ends <strong>of</strong> <strong>the</strong> femur in Limnosceloides are not<br />

so wide in comparison with <strong>the</strong> shaft as in Limnoscelis<br />

whose femur is short, stoutly built, and in general comparable<br />

to <strong>the</strong> femur in Seymouria. (See Romer, 1952,<br />

fig.12; 1956, figs.171A., B, 0, F.)<br />

The trochanteric crest is set <strong>of</strong>f <strong>from</strong> <strong>the</strong> head by a<br />

distinct notch as in Labidosaurus (Romer, 1956, fig.<br />

171G), but in contrast to Lhnnoscelis, makes almost<br />

a right angle with <strong>the</strong> head, and does not flare widely<br />

at <strong>the</strong> front. The trochanteric crest in Labidosaurus<br />

and L,imnoscelis is directed anteriorly as well as ventrally,<br />

so that it makes an obtuse ventral angle with <strong>the</strong><br />

head; only <strong>the</strong> base <strong>of</strong> <strong>the</strong> crest is preserved in Limnosceloides<br />

dunkardensis (USNM 12166). There is a<br />

prominent excavation on <strong>the</strong> posterior side <strong>of</strong> <strong>the</strong> posterior<br />

condyle as in Labidosaurus and Lim;noscelis, but<br />

in contrast to Limnosceloides.<br />

To sum up <strong>the</strong> significant morphologic features <strong>of</strong><br />

Limnoscelops longifemur: The vertebrae are typically<br />

limnoscelid, <strong>the</strong> preserved parts <strong>of</strong> <strong>the</strong> pelvis are much<br />

like those in Lirrunosceloides except for <strong>the</strong> thicker symphysis,<br />

and for <strong>the</strong> almost vertical, captorhinomorphlike<br />

face <strong>of</strong> <strong>the</strong> conjoined internal ridges <strong>of</strong> <strong>the</strong> pubes.<br />

Romer (1956) recognizes three families <strong>of</strong> captorhinmnorph<br />

cotylosaurs: <strong>the</strong> Limnoscelidae, <strong>the</strong> Romeriidae,<br />

and <strong>the</strong> Captorhinidae. The consensus seems to be<br />

that vertebral structure is more conservative than <strong>the</strong><br />

structure <strong>of</strong> <strong>the</strong> appendages, as illustrated by <strong>the</strong> currently<br />

accepted classification <strong>of</strong> labyrinthodont amphibians;<br />

thus, <strong>the</strong> classification <strong>of</strong> Limnoscelops must depend<br />

more on its vertebrae than on <strong>the</strong> o<strong>the</strong>r materials<br />

available to us. We <strong>the</strong>refore tentatively assign Limnoscelops<br />

to <strong>the</strong> family Limnoscelidae. Romer's assignment<br />

<strong>of</strong> Limnosceloides to this same family was also<br />

only tentative. Limnoscelis, known only <strong>from</strong> <strong>the</strong> <strong>Cutler</strong><br />

(El Cobre Canyon) <strong>of</strong> New Mexico, has heret<strong>of</strong>ore<br />

been <strong>the</strong> only unquestioned representative <strong>of</strong> this family.<br />

Limnoscelops, for <strong>the</strong> time being, is best thought<br />

<strong>of</strong> as a limnoscelid advanced in <strong>the</strong> direction <strong>of</strong> <strong>the</strong><br />

captorhinids.<br />

The romeriids and captorhinids are closely related,<br />

<strong>the</strong> captorhinids having apparently been derived <strong>from</strong><br />

<strong>the</strong> romeriids. The postcranial skeleton <strong>of</strong> <strong>the</strong> romeriids<br />

is, unfortunately, still unknown. Protorothyris and<br />

Romeria <strong>of</strong> <strong>the</strong> romeriids are known <strong>from</strong> <strong>the</strong> lower<br />

part <strong>of</strong> <strong>the</strong> Wichita Group in Texas, M elanothyris is<br />

known <strong>from</strong> <strong>the</strong> lower part <strong>of</strong> <strong>the</strong> Dunkard Group in<br />

<strong>the</strong> eastern United States, and Oephalerpeton may be<br />

a romeriid <strong>from</strong> <strong>the</strong> Allegheny <strong>Formation</strong> <strong>of</strong> Illinois.<br />

The Wichita Group has yielded only a few primitive<br />

captorhinids, but <strong>the</strong> advanced Oaptorhinus and Labidosaurus<br />

are common in <strong>the</strong> over-lying Clear Fork<br />

Group.<br />

The known limnoscelids could not have been <strong>the</strong> actual<br />

ancestors <strong>of</strong> <strong>the</strong> romeriids or captorhinids if we<br />

consider <strong>the</strong>ir respective ages. The phylogenetic divergence<br />

must <strong>the</strong>refore have occurred no later than<br />

Middle Pennsylvanian time.

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