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The Burrowing Geckos of Southern Africa 5 1976.Pdf ANNALS OF THE TRANSVAAL MUSEUM ANNALE VAN DIE TRANSVAAL-MUSEUM- < VOL. 30 30 JUNE 1976 No. 6 THE BURROWING GECKOS OF SOUTHERN AFRICA, 5 (REPTILIA: GEKKONIDAE) By W.D. HAACKE Transvaal lvIllseum, Pretoria (With four Text-figures) ABSTRACT This study deals with the entirely terrestrial genera of southern African geckos and is published in five parts in this journal. In this part the phylogenetic and taxono­ mic affinities of these genera based on pupil shape and hand and foot structure are discussed. PHYLOGENETIC AND TAXONOMIC AFFINITIES In his classification of the Gekkonidae, Underwood (1954) placed six South African genera into the subfamily Diplodactylinae. These genera, i.e. Cho!ldrodac~ylus, Colopus, Palmatogecko, Rhoptropus, Rhoptropella and Ptmopus were supposed to share a peculiar variant of the straight vertical pupil which he called Rhoptropus-type. He notes that all of them occur in "desert or veldt" and appear to be adapted to the special conditions of South Africa. He further states that "Such a number of genera with several peculiar forms of feet in such a restricted area is somewhat surprising". At that time all except Rhoptropuswere considered to be monotypic genera. Since then two more species of Ptmopus and also the terrestrial, obviously related genus Kaokogecko have been described. In the present paper a special study has been made of the ground­ living, burrowing genera, which excludes Rhoptropus and Rhoptropella. Although it has been pointed out that the classification of the Gekkonidae according to the form of the digits (Boulenger, 1885) and the shape of the pupil (Underwood, 1954) is unsatisfactory (Stephenson, 1960; Kluge, 72 1964 and 1967) the possibilities of these characters as taxonomic indicators were reinvestigated in the genera in question and related forms. PUPIL SHAPE According to Underwood (1954) the six diplodactyline gekkonid genera all have the same type of pupil, which he referred to as £(boptroptls­ type. Pasteur (1960) found that the edge of the pupil of Sr:mrodacrylffs mauretanictfs and fascialtls, although straight-edged in life, became wavy after preservation. As the pupil always distorts in the same way after death, he assumes that this phenomenon corresponds to a histological peculiarity of the iris of these animals. Kluge (1964: 4) found that in live material the pupil shape varied slightly under different lighting conditions, but markedly so in preserved material. He thereby proved the unreliability of the pupil as a diagnostic character and no longer recognized Underwood's subfamilial groups. Amongst the genera under discussion Rhoptropella and Ptmoptls have a straight vertical pupil while the remaining ones, excluding Rhoptroptls} which is discussed below, have pupils with lobed margins which overlap to some extent, thereby closing down to four pinholes, a shape which Underwood (1954) referred to as Gekko-type. This shows that even by Underwood's standards this group of genera is not necessarily closely related and illustrates the danger of working with an inadequate series of specimens. Preserved specimens of Rhoptroptfs were also examined for pupil shape. The so-called Rhoptroptts-type is only present in R. afer. In R. bradfteldi and R. barnardi the anterior edge is nearly straight, while the posterior edge is lobed. R. bottltoni and R. tatmiosticttls have clearly Gekko-type pupils. The variation in the pupil shape of this genus thus confirms Kluge'S (1964) findiogs regarding the unreliability of this character as a diagnostic feature for sub-familial separation. PEDAL CHARACTERISTICS Boulenger (1885) based his classification of this group mainly on the external pedal characteristics, but Stephenson (1960) and Kluge (1967) pointed out the dangers involved in using the phalangeal formulae as an indication of relationship and phylogeny of the gekkonids. Nevertheless, phalangeal structure proved taxonomically important when splitting Crenodactyllfs from PI!Jllodacrylus and Ebenavia (Dixon & Kluge, 1964). Because of the unusual structure of the appendages of the terrestrial gekkonid genera of southern Africa a special examination of these appears to be warranted. 1. Ptenopus garrulus (TM 32705) MANUS: Phalangeal formula 2,3,4,5,3 (Fig. 1c). A small sesamoid occurs dorsal to the radiale while a large, flat sesamoid covers most of the palmar surface. The first and fifth metacarpals are shorter and thicker than the remaining ones. A prominent protuberance occurs near the proximal epiphysis of metacarpal 1. All the phalangeal articulation surfaces are rather flat and the phalanges are tightly joined indicating that only a 73 minimum of lateral movement is possible. Proximally the terminal, claw­ bearing phalanges extend over the distal condyle of the penultimate phalanges and a sesamoid occurs just behind this extension. Ventrally a sesamoid occurs below the metacarpal-phalangeal joint as well as below all the interphalangeal joints except the terminal ones of fingers two to five. Ventrally the proximal articuution surfaces of all except the terminal phalanges extend backwards. The flexor muscles, in which the above­ mentioned sesamoids are embedded, are attached to these extensions. The terminal claw-bearing phalanges have a midventral swelling to which the contractor tendon is attached (cf. Mahendra, 1941: 296). PES: Phalangeal formula 2,3,4,5,4 (Fig. 1a and b). Four small sesamoid bones, of which three lie helow the tibio-fibulare and the fourth below the cuboid, occur on the palmar surface of the tarsus. Metatarsals one to four are long and thin while five is only half as long as four. The fifth metatarsal is dub-shaped and much less hooked than in the other genera under discussion. The shape and articulations of the tarsal phalanges are similar to the situation described for those of the hand. However, the ventral, backward extension of the proximal articulation surfaces is less well developed. Large sesamoids occur ventral to the metatarsal phalan­ geal joints, while smaller to minute ones occur under the first joint of toes 2 and 5, first and second of toe 3 and first to third of toe 4. Dorsally sesamoids occur above the metatarsal-phalangeal joints as well as above the first interphalangeal joint of toe 1, the two terminal joints of toes 2 and 3 and the three terminal ones of toes 4 and 5. The claws of Ptmopus are not retractile and the fifth toe is not opposable. Furthermore the phalanges are capable of only limited lateral movement, and the digits are not widely spread but lie nearly parallel to each other. In later discussions this type of structure will be referred to as a straight­ fingered or straight-toed pattern. 2. Ptenopus carpi (TM 32332) MANUS: Phalangeal formula 2,3,4,5,3. In general similar to that de­ scribed for garrulus but with slightly thicker metacarpals, and very large sesamoids on the ventral side of the fingers, except the terminal joint of finger 1 where none is present. On the dorsal side the sesamoids are very poorly developed and in some cases are hardly discernible. PES: Phalangeal formula 2,3,4,5,4. General structure very similar to that described for garrulus but with only two sesamoids on the palmar surface below the tibio-fibulare. The sesamoids on the ventral side are much better developed than those of garrulus but those on the dorsal side are much less well developed and only a single, minute sesamoid occurs just proximally to the terminal joint on all five toes. 3. Ptenopus kochi (TM 31656) MANUS: Phalangeal formula 2,3,4,5,3. Basically identical to the struc­ ture described for garrulus but with less well developed sesamoids. No sesamoids were noticed dorsal to the terminal joint nor ventral to the third joint of digit 4. PES: Phalangeal formula 2,3,4,5,4. Similar in structure to that de­ scribed for garrulus but differing on the following points: Only two sesa- 74 moids occur ventral to the tibio-fibulare while two minute sesamoids oc­ cur next to the small first tarsal. A minute sesamoid occurs near the proxi­ mal epiphysis on the femoral side of metatarsals 2 to 4. In the toes sesa­ moid bones occur in the identical positions to those of germ/us except in toe 5 where the one ventral to the first interphalangeal joint is absent. 4. Palmatogecko rangei (TM 22830 female, TM 33199 male) MANUS: Phalangeal formula 3,3,4,5,3 (Fig. 2f). No sesam~ids occur on the ventral side of the carpus but a minute one occurs dorsolateral to the first carpal. The male specimen has an extra one dorsolateral to the distal epiphysis of the ulna. The metacarpals are spread to the extreme so that the first and fifth fingers are pointing in opposite directions. Of the meta­ carpals, which are elongate and normal, the second is the longest while the fifth is the shortest. Cartilaginous, slightly ossified, club-shaped structures occur lateral to the distal end of the metatarsals and the first phalanx of finger 4. These supports of the soft interdigital membrane have of the first phalanx of fingers 1, 2 and 3, as well as of the second phalanx of finger 4, have lateral, cartilaginous, partly ossified rod-like extensions, similar to those occurring in the toes of Kaokogecko. The terminal two phalanges of fingers 1, 2 and 5 and the terminal three phalanges of fingers 3 and 4 are bent upwards in similar fashion to those in geckos with vertically retractile claws and adhesive subdigital lamellae. No sesa­ moids were seen in the manus. The cartilaginous, rod-like reinforcements of the interdigital web are better developed in the female specimen than in the male. This may be due to age. PES: Phalangeal formula 3,3,4,5,4 (Fig. 2e). A single sesamoid occurs ventral to the first tarsaL Metatarsals 1 to 4 are thin and elongate while the fifth is so short and hooked that it no longer resembles a metatarsal but appears to be a true tarsal element.
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