A New Order of Fishlike Amphibia From the Pennsylvanian of Kansas Part 2

Although we are unable to separate, by sutures, the vomers from the palatines, the palatal surface of these bones and of the pterygoids is studded by numerous small teeth, as in Rhipidistia (Jarvik, 1954) and some of the early Amphibia (Romer, 1947). The stapes apparently reaches the quadrate, and could therefore serve in hyostylic suspension of the upper jaw.

The pectoral limb has an axial series of bones carrying hooklike f.l.a.n.g.es on their posterior edges. The other bones of the limb show little modification of form beyond the nearly flat, aquatic type seen in Rhipidistia. No distinct elbow or wrist joints are developed.

Characters of _Hesperoherpeton_ common to most primitive Amphibia, in contrast with Crossopterygii, are: Nares separated from edge of jaw; stapes having external process that may have met a tympanic membrane, thus giving the bone a sound-transmitting function. Apparently none of the opercular series was present.

There are two large palatal teeth, slightly labyrinthine in character, adjacent to each internal naris. The scapulocoracoid, as shown by Peabody (1958), is Anthracosaurian in structure, as are the long-stemmed clavicles. The limbs have digits rather than fin-lobes, although the digital number apparently is four and the number of bones in the ma.n.u.s is less than would be expected in a primitive amphibian.

The vertebrae are similar to those of Ichthyostegids, as described by Jarvik (1952), except that the pleurocentra are much larger.

In addition to this remarkable combination of crossopterygian and amphibian characters, _Hesperoherpeton_ is specialized in certain features of the skull. The orbits are much enlarged, probably in correlation with the diminutive size of the animal, and this has been accompanied by loss of several bones. The frontal and squamosal nearly meet each other, and both form part of the rim of the orbit. The bones of the posterior part of the dermal roof are greatly reduced, and there is none behind the squamosal except the projecting tabular; there is no indication of quadratojugal, jugal, intertemporal or postparietal. The foramen magnum is enormous. The external surfaces of the bones of the skull are nearly smooth.

Is it possible that the "primitive" and "specialized" features of this animal are actually larval? Are they not just the kind of characters that would be expected in an immature, aquatic embolomere of Pennsylvanian time? For several reasons we do not think this is the case. Except for the anterior part of the braincase, there is no indication that the skeleton was not well ossified. The postaxial processes on the humerus, ulna and ulnare could scarcely have been larval features only, since they are so clearly h.o.m.ologous with those in adult Rhipidistia; a larval limb should indeed be simple, but its simplicity is unlikely to involve paleotelic adult characters. The scapulocoracoid of our specimen is of practically the same shape and size as that in the only other known individual, the type; this would be probable if both were adults, but somewhat less likely if they were larvae of a much larger animal. The form of the stapes, tabular and otic notch suggest a functional tympanic membrane, which could not have occurred in a gill-breathing larva. On the other hand, an adult animal of pigmy size might be expected to have large orbits, large otic capsules and a large foramen magnum.

We conclude that _Hesperoherpeton_ lived and sought food in the weedy shallows at the margin of a pond or lagoon, and that for much of the time its head was partly out of water (Fig. 12). The animal could either steady itself or crawl around by means of the paddlelike limbs, but these probably could not be used in effective locomotion on land.

Like the Ichthyostegids, it probably swam by means of a fishlike tail.

[Ill.u.s.tration: FIG. 12. _Hesperoherpeton garnettense_ Peabody. Probable appearance in life. 0.5.]

TAXONOMY

Evidently _Hesperoherpeton_ is a small, lagoon-dwelling survivor of the Devonian forms that initiated the change from Crossopterygii to Amphibia (Jarvik, 1955). It shows, however, that this transition did not affect all structures at the same time, for some, as the braincase with its notochordal ca.n.a.l, the mandibular bones and axial limb bones, are unchanged from the condition normal for the Rhipidistia, but most other characters are of amphibian grade. To express these facts taxonomically requires that _Hesperoherpeton_ be removed from the family Cricotidae, suborder Embolomeri, order Anthracosauria, and placed in a new order and family of labyrinthodont Amphibia.

Order PLESIOPODA

(_plesios_, Gr., near, almost; _podos_, Gr., foot)

Labyrinthodontia having limbs provided with digits, but retaining posterior f.l.a.n.g.es on axial bones as in Rhipidistia, without joint-structure at elbow and wrist essential for terrestrial locomotion; neurocranium having separate otico-occipital section, large notochordal ca.n.a.l, no occipital condyle, as in Rhipidistia; nares separate from rim of mouth; pectoral girdle anthracosaurian; vertebrae having U-shaped intercentrum and paired, but large, pleurocentra.

Probably a.s.sociated with the characters of the order, as given above, are the connection of pectoral girdle with skull, and the presence of a tympanic membrane, the stapes functioning in both sound-transmission and palatoquadrate suspension.

Family HESPEROHERPETONIDAE

Orbits and foramen magnum unusually large in correlation with reduced size of animal; squamosal forming posterior margin of orbit; circ.u.morbital series absent (except for pos...o...b..tal); sensory pits on squamosal and frontal.

Characters defining the family are evidently the more specialized cranial features, which probably evolved during Mississippian and early Pennsylvanian times.

The definition of the genus and species may be left to rest upon Peabody's (1958) original description and the present account, until the discovery of other members of the family gives reason for making further distinctions.

SUMMARY

_Hesperoherpeton garnettense_ Peabody (1958), based on a scapulocoracoid and part of a vertebra, was originally placed in the order Anthracosauria, suborder Embolomeri, family Cricotidae. A new skeleton from the type locality near Garnett, Kansas (Rock Lake shale, Stanton formation, Upper Pennsylvanian), shows that the animal has the following rhipidistian characters: Large notochordal ca.n.a.l below foramen magnum, otico-occipital block separate from ethmosphenoid, postaxial processes on three axial bones of forelimb, pectoral girdle (probably) articulated with tabular. Nevertheless, _Hesperoherpeton_ has short digits, an anthracosaurian type of pectoral girdle, an otic rather than spiracular notch, nostrils separate from the mouth, and vertebrae in which the intercentrum is U-shaped and the pleurocentra large but paired. The stapes reaches the quadrate.

_Hesperoherpeton_ is placed in a new order, PLESIOPODA, on the basis of the characters stated above, and a new family, HESPEROHERPETONIDAE.

Specialized characters of the family include: Reduction of circ.u.morbital bones, bringing the squamosal to the edge of the orbit, loss of certain bones of the temporal region, and relative enlargement of the orbits and foramen magnum, in correlation with the diminutive size of the animal. The structural characters of _Hesperoherpeton_ suggest to us that it lived in the shallow, weedy margins of lagoons, rested with its head partly out of water, and normally did not walk on land.

LITERATURE CITED

EATON, T. H., JR.

1951. Origin of tetrapod limbs. Amer. Midl. Nat., 46: 245-251.

JARVIK, E.

1952. On the fish-like tail in the ichthyostegid stegocephalians.

Meddel. om Grnland, 114: 1-90.

1954. On the visceral skeleton in _Eusthenopteron_ with a discussion of the parasphenoid and palatoquadrate in fishes. Kgl. Svenska Vetenskapsakad. Handl., 5: 1-104.

1955. The oldest tetrapods and their forerunners. Sci. Monthly, 80: 141-154.

MOORE, R. C., FRYE, J. C., and JEWETT, J. M.

1944. Tabular description of outcropping rocks in Kansas. Kansas State Geol. Surv. Bull., 52: 137-212.

PEABODY, F. E.

1952. _Petrolacosaurus kansensis_ Lane, a Pennsylvanian reptile from Kansas. Univ. Kansas Paleont. Contrib., Vertebrata, Art. 1: 1-41.

1958. An embolomerous amphibian in the Garnett fauna (Pennsylvanian) of Kansas. Jour. Paleont., 32: 571-573.

ROMER, A. S.

1937. The braincase of the Carboniferous crossopterygian _Megalichthys nitidus_. Mus. Comp. Zool. Bull., 82: 1-73.

1947. Review of the Labyrinthodontia. Mus. Comp. Zool. Bull., 99: 1-368.

1957. The appendicular skeleton of the Permian embolomerous amphibian _Archeria_. Univ. Michigan Contrib. Mus. Paleont., 13: 103-159.

WATSON, D. M. S.

1926. The evolution and origin of the Amphibia. Phil. Trans. Roy.

Soc. London, (B) 214: 189-257.

_Transmitted January 13, 1960._

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