Alexander F. Flemming

Population fragmentation in the southern rock agama, Agama atra: more evidence for vicariance in Southern Africa

Mitochondrial DNA sequence data derived from two genes were used to infer phylogeographical relationships between 13 Agama atra populations. Three distinct geographical assemblages were found among the lizard populations. The first occurs in southern Namibia, the second is restricted to the western dry arid regions of South Africa, whereas the third is distributed throughout the more mesic southern and eastern parts of the subcontinent. Geographically structured differences among populations within Agama clades are probably the result of dispersal and historic isolations among populations. At the broader scale, there were marked congruences between the Agama genetic discontinuities and those described previously in other rock‐dwelling vertebrates such as Pronolagus rupestris and Pachydactylus rugosus. This suggests vicariance, probably as a response to natural climatic changes during the past three million years.

Extraordinary case of matrotrophy in the African skink Eumecia anchietae

The viviparous African skink, Eumecia anchietae, exhibits a matrotrophic fetal nutritional pattern. Until well after the limb bud stage, extravitelline nutritional provision is in the form of holocrine secretion originating from the stratified uterine epithelium of the uterine incubation chambers. Uterine secretions are absorbed by a specialized yolk sac ectoderm and chorioallantois through histotrophy. The yolk sac is not in close contact with the uterine lining from the limb bud stage onwards. The yolk sac ectoderm forms invaginations filled with uterine secretion and consists of a single layer of vacuolated hypertrophied cells bearing microvilli. The chorioallantois at the limb bud stage is extensive, well‐vascularized, and not intimately associated with the uterine epithelium. Where the uterus is folded, the chorioallantois may interdigitate loosely. Chorionic cells are low to high columnar, clearly vacuolated, and bear microvilli. The allantoic layer consists primarily of squamous cells exhibiting villous projections. By the time embryos have well‐defined digits, the specialized yolk sac ectoderm has regressed and the yolk sac lumen has been invaded by vitelline cells. The chorioallantois is very extensive and in areas greatly folded. Where it contacts the uterine epithelium, a proper chorioallantoic placenta is formed. Cell layers of the chorioallantois and uterine epithelium are thin and cuboidal to squamous in appearance. The chorioallantoic placenta is simple in structure, occurs throughout the incubation chamber, and is epitheliochorial in arrangement. It is unknown whether the placentome observed in other highly matrotrophic scincids is formed in late stage embryos of this species. J. Morphol. 247:264–287, 2001.

Morphology, development, and evolution of fetal membranes and placentation in squamate reptiles

Current studies on fetal membranes of reptiles are providing insight into three major historical transformations: evolution of the amniote egg, evolution of viviparity, and evolution of placentotrophy. Squamates (lizards and snakes) are ideal for such studies because their fetal membranes sustain embryos in oviparous species and contribute to placentas in viviparous species. Ultrastructure of the fetal membranes in oviparous corn snakes (Pituophis guttatus) shows that the chorioallantois is specialized for gas exchange and the omphalopleure, for water absorption. Transmission and scanning electron microscopic studies of viviparous thamnophine snakes (Thamnophis, Storeria) have revealed morphological specializations for gas exchange and absorption in the intra-uterine environment that represent modifications of features found in oviparous species. Thus, fetal membranes in oviparous species show morphological differentiation for distinct functions that have been recruited and enhanced under viviparous conditions. The ultimate in specialization of fetal membranes is found in viviparous skinks of South America (Mabuya) and Africa (Trachylepis, Eumecia), in which placentotrophy accounts for nearly all of the nutrients for development. Ongoing research on these lizards has revealed morphological specializations of the chorioallantoic placenta through which nutrient transfer is accomplished. In addition, African Trachylepis show an invasive form of implantation, in which uterine epithelium is replaced by invading chorionic cells. Ongoing analysis of these lizards shows how integration of multiple lines of evidence can provide insight into the evolution of developmental and reproductive specializations once thought to be confined to eutherian mammals.

Invasive implantation and intimate placental associations in a placentotrophic African lizard, Trachylepis ivensi (Scincidae)

In the viviparous lizard Trachylepis ivensi (Scincidae) of central Africa, reproducing females ovulate tiny ∼1 mm eggs and supply the nutrients for development by placental means. Histological study shows that this species has evolved an extraordinary placental pattern long thought to be confined to mammals, in which fetal tissues invade the uterine lining to contact maternal blood vessels. The vestigial shell membrane disappears very early in development, allowing the egg to absorb uterine secretions. The yolk is enveloped precocially by the trilaminar yolk sac and no isolated yolk mass or yolk cleft develops. Early placentas are formed from the chorion and choriovitelline membranes during the neurula through pharyngula stages. During implantation, cells of the chorionic ectoderm penetrate between uterine epithelial cells. The penetrating tissue undergoes hypertrophy and hyperplasia, giving rise to sheets of epithelial tissue that invade beneath the uterine epithelium, stripping it away. As a result, fetal epithelium entirely replaces the uterine epithelium, and lies in direct contact with maternal capillaries and connective tissue. Placentation is endotheliochorial and fundamentally different from that of all other viviparous reptiles known. Further, the pattern of fetal membrane development (with successive loss and re‐establishment of an extensive choriovitelline membrane) is unique among vertebrates. T. ivensi represents a new extreme in placental specializations of reptiles, and is the most striking case of convergence on the developmental features of viviparous mammals known. J. Morphol. 2011.

ACTIVE GENERATION GLANDS PRESENT IN NEONATES OF SOME CORDYLID LIZARDS : A CASE STUDY OF CORDYLUS MACROPHOLIS (SAURIA: CORDYLIDAE)

Generation glands are holocrine epidermal glands occurring on the ventral aspect of the thigh of cordylid lizards. In most species these glands seemingly start to differentiate with the onset of sexual maturity, but macroscopic signs of generation gland activity were noted in neonates of the large‐scaled girdled lizard, Cordylus macropholis. The glands of neonatal, subadult, and adult individuals were examined microscopically using standard histological techniques. The glands of the five neonatal specimens examined, including both males and females, all had the same basic structure and displayed two layers of mature glandular material. In subadult and adult specimens, the number of layers varied from seven to nine. The structure of the generation glands of C. macropholis is similar to that of the few other Cordylus species that have been described to date. They are of the protruding kind with multiple mature glandular generations. Juveniles of an additional 12 cordylid species have been examined for the presence of active generation glands. Active glands were found to be present in neonates of C. tasmani and C. tropidosternum, both of which are, like C. macropholis, terrestrial species. In C. cordylus and C. coeruleopunctatus, active generation glands are absent in neonates, but differentiate soon after birth. In other cordylid species, generation glands apparently differentiate only with the onset of sexual maturity. J. Morphol. 235:177–182, 1998.

Foraging mode of serpentiform, grass-living cordylid lizards: A case study of Cordylus anguina

Foraging mode of the Cape grass lizard, Cordylus anguina, was determined by using three criteria: 1) the number of movements per minute (MPM) and the percentage of time spent moving (PTM) during periods of activity; 2) prey chemical discrimination ability; 3) stomach contents. To determine MPM and PTM, observations were made from an observation tower using a semi-natural outdoor enclosure, and through a one-way glass panel using a semi-natural indoor enclosure. Each lizard (n = 10) was observed for a 10 minute period during peak activity and the times it was moving and the times it was stationary were recorded. Data obtained in the indoor- and outdoor enclosures did not differ significantly and were pooled. MPM (0.37 ± 0.21 S.D.) and PTM (1.92 % ± 0.93 S.D.) values recorded for C. anguina fall within the range given as characteristic for sit-and-wait foragers. Nine C. anguina were habituated in glass terraria to accept mealworms offered to them. When all lizards accepted food without hesitation, they were tested for their ability to discriminate among three different odours presented to them in a randomized block design: prey odours consisting of mealworm surface odours, distilled water as an odourless control stimulus, and cologne as a pungency control. The number of tongue-flicks directed at the cotton applicator containing the stimulus odour during 60 seconds was recorded. No statistically significant differences were found among the responses to the three treatment odours. An analysis of the stomach contents of 21 C. anguina revealed a diet of mostly diurnally active arthropods.

Intergroup-movement in a group-living lizard, Cordylus cataphractus, from South Africa : short communication

Extracted from text ... Short communication Intergroup-movement in a group-living lizard, Cordylus cataphractus, from South Africa LOUISE VISAGIE1, P. LE FRAS N. MOUTON2, AND ALEXANDER F. FLEMMING1 1, 2Department of Zoology, University of Stellenbosch, Private Bag X1, Matieland 7602, South Africa, 2Corresponding author:pnm@.sun.ac.za Key words.- Cordylidae, Armadillo Lizard, group stability, mark-recapture. 75 African Journal of Herpetology, 2002 51(1): 75-80. The armadillo lizard, Cordylus cataphractus, is a rock-dwelling cordylid restricted to the arid, western region of South Africa (Mouton 1988). It naturally occurs in groups (Peers 1930; Mouton et al. 1999) and laboratory experiments have shown that its grouping behaviour is the result of mutual ..

Aseasonal reproduction and high fecundity in the Cape grass lizard, Cordylus anguinus , in a fire-prone habitat

We investigated life history characters of the Cape grass lizard, Cordylus anguinus , and relate them to survival in the fire-prone habitat in which it occurs. Unlike in other cordylids, reproductive activity was found to be asynchronous among females, with vitellogenic and gravid females encountered virtually throughout the year. Aseasonal breeding will circumvent reduction or complete loss of reproductive effort for any given year due to fire. Female grass-lizards attain significantly larger body sizes than males. Clutch size ranged from three to seventeen and was positively correlated with snout-vent length. Maximum clutch size is more than three times that recorded for any other cordylid. We suggest that high fecundity will allow quick recruitment after a fire. To determine the reproductive cycle exhibited by males, testicular volume and seminiferous tubule diameter were measured, and spermatogenic activity assessed qualitatively, using Lichts classification system. Our data indicate that C. anguinus has a post-nuptial spermatogenic cycle. The species differs from other cordylids having a post-nuptial cycle, in that spermatogenesis already commences in spring.

Geographic variation in sexual size dimorphism in the rock agama, Agama atra (Sauria: Agamidae)

Significant sexual dimorphism in overall size occurs in the rock agama, Agama atra (Sauria; Agamidae), with males growing larger than females. Geographic variation in the degree of sexual size dimorphism also exists, males growing significantly larger than females in Namaqualand and Namibia compared to populations in other areas. Sexual differences in scaling of head, limb and tail dimensions were mainly the result of differential asymptotic sizes reached by the sexes. Head size was also influenced by a faster increase in head dimensions with increasing snout to vent length in males compared to females, probably as a result of sexual selection. In females, scaling of limb and tail dimensions was decreased compared to males, possibly a result of differential energy allocation to reproduction.

Generation gland morphology in cordylid lizards: An evolutionary perspective

To elucidate the functional significance of the three distinct types of generation glands that have been identified among cordylid lizards, we mapped gland type to the terminal taxa in the most recent phylogenetic tree for the Cordylidae. We used the phylogenetic programme Mesquite and applied the principle of parsimony to infer character states for the ancestral nodes in the tree. For those species where information on gland type was not available from the literature, we conducted a histological investigation of generation gland morphology, using standard histological techniques. We included two species of the sister family Gerrhosauridae in the analysis to serve as outgroups. In both Gerrhosaurus typicus and G. flavigularis, scales immediately anterior to the femoral pores displayed glandular activity, but differed from generation glands of cordylids in the absence of mature glandular generations. Among the cordylids investigated, we identified a fourth type of generation gland in Pseudocordylus subviridis, P. spinosus, and in the two Hemicordylus species, one where the glands consistently comprise of two mature glandular generations. In H. capensis, both single‐ and two‐layer type glands are present. Our reconstruction of ancestral character states suggests a minimum of six transformations from one gland type to another during the evolutionary history of the family. The reconstruction furthermore suggests that the single‐layer type gland reappeared at least once (in Hemicordylus) in the Cordylinae after having been lost. The reconstruction also unequivocally shows that the pit‐like multiple‐layer type gland evolved directly from the single‐layer type and not from the protruding multiple‐layer type. The two‐layer type gland appears to be an intermediary condition between the multiple‐layer and single‐layer types. The evolutionary transformation of generation gland type appears to be linked to changes in lifestyle and associated changes in degree of territoriality and the need for chemical communication. J. Morphol. 275:456–464, 2014.