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 homologous 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.
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 canal, 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)