Appendicular Skeleton in Amphisbaenians (Reptilia: Squamata)
The appendicular skeletal elements in 32 species from all 23 extant genera of amphisbaenians were surveyed via the preparation and study of radiographs, dry skeletons, and cleared-and-stained specimens. Here I report on the presence or absence of those elements, describe their morphology, and make comparisons to several other limbless or limb-reduced squamate taxa that have previously been suggested as close relatives to amphisbaenians. Variability found here in the appendicular skeleton provides a new source of character data for ongoing phylogenetic studies of the group. Comparisons of these elements to those found in potential squamate sister groups indicate that some amphisbaenians share plesiomorphic similarities with limbed squamates such as gymnophthalmid lizards, whereas others share only absence or reduction features with other limbless or limb-reduced squamates, such as dibamid lizards and snakes. This pattern is considered potentially inconsistent with previously proposed hypotheses of amphisbaenian sister groups within squamates. For the pectoral girdle, every stage of reduction—including complete loss—occurs among lizards, amphisbaenians, and snakes. However, the pelvic girdle is never completely lost in lizards or amphisbaenians regardless of the degree of limb reduction or loss; all these taxa retain at least the ilium, and this is also true of most basal snakes. These pelvic girdle elements are variously modified among these taxa, a fact that, together with their widespread persistence, suggests both different patterns of reduction among higher taxa and possible new functional roles beyond the primitive one of supporting the hind limb.Abstract

(A) Most amphisbaenians are completely limbless, as in this species, Amphisbaena alba (photo credit: R. McDiarmid). (B) Bipes biporus, one of the three species of amphisbaenians (all in the genus Bipes) that exhibits limbs (forelimbs only)

Pectoral girdles and forelimbs in amphisbaenians. (A) Ventral view of Blanus cinereus. (B) Ventral and right lateral views of Bipes biporus. Ventral views of (C) Agamodon anguliceps, (D) Diplometopon zarudnyi, (E) Pachycalamus brevis (redrawn from Gans, 1960), and (F) Trogonophis wiegmanni

Radiographs of (A) Bipes biporus and (B) Corucia zebrata, showing relative positions of pectoral girdle and forelimbs

Pelvic girdles and hind limbs in amphisbaenians. Each set represents a ventral view on the left and a right lateral view on the right. The paired bony element occurring in all taxa other than Bipes biporus and Blanus cinereus represents the ilium. (A) Amphisbaena alba, (B) Anops kingi, (C) Aulura anomala, (D) Baikia africana, (E) Blanus cinereus, (F) Cadea blanoides, (G) Cynisca leucura, (H) Dalophia ellenbergeri, (I) Leposternon microcephalum, (J) Mesobaena huebneri, (K) Monopeltis capensis, (L) Zygaspis quadrifrons, (M) Bipes biporus, (N) Rhineura floridana, (O) Agamodon anguliceps, (P) Diplometopon zarudnyi, and (Q) Trogonophis wiegmanni

Pectoral girdles (when present) and pelvic girdles in some non-amphisbaenian squamates. Each set represents a ventral view of the pectoral girdle on the left (if a pectoral girdle is present) and a ventral view of the pelvic girdle on the right. (A) Acontias meleagris, (B) Anniella pulchra (no pectoral girdle present), (C) Dibamus novaeguineae, (D) Bachia trisanale