Thesis (MSc)--University of Stellenbosch, 2004. / ENGLISH ABSTRACT: Geographic variation in the presence/absence of generation glands in females of the cordylus-niger-oelofseni complex was described and correlated with climatic variables. Preserved and live specimens from 96 localities in the area south of 32°30' S and west of 19°45' E in the Western Cape, South Africa, were examined for the presence or absence of generation glands. Females from western coastal localities and from the Cape Fold Mountains generally lack generation glands, while females from inland lowland localities generally possess glands. A GIS analysis was performed to determine mean annual minimum temperature, mean annual maximum temperature, potential evaporation, mean annual precipitation, mean annual fog and mean annual cloud cover measured at 14:00 for each locality. Discriminant function and canonical analyses showed a highly significant correlation between the presence and absence of generation glands and five of the six climatic variables. It is suggested that the high incidence of fog and lower temperatures experienced along the west coast as a result of the cold Benguela Current, and the high incidence of orographic fog and cloud cover in the Cape Fold Mountains may affect energy allocation in females. In order to maintain their reproductive output, females may be forced to invest less energy in secondary sexual characters such as generation glands. Interspecific variation in the location, occurrence, number, and age of gland differentiation was investigated for cordylid lizards. For several species, data were obtained from the literature and for others data were obtained from material housed in the Ellerman Collection of the University of Stellenbosch. Cordylus cordylus and C. cataphractus were used for an evaluation of inter- and intrapopulation variation. Data for C. cordylus were obtained from five populations on the coastal lowlands of the Western Cape, while data on C. cataphractus come from the literature. The sex, snout-vent length, and number of generation glands were recorded for a representative sample of lizards at each locality. While in all species generation glands were located in the femoral region, Cordylus giganteus and C. microlepidotus had additional generation glands elsewhere on the body. It is suggested that additional generation glands in these species may be related to an increased need for chemical communication in a low-visibility environment in the case of C. giganteus, and in low population densities in the case of C. microlepidotus. Generation glands are present in all cordylid males, but females of some species may lack these glands. All cordylids display sexual dimorphism with regards to the presence and number of generation glands, with males having significantly more glands than females, indicating that these glands may play a more important role in males than in females. Low sexual dimorphism in at least two species suggests that generation glands may be equally important in both sexes of these species. In females, the presence of generation glands is closely regulated by climate. There is no indication of differences in generation gland profile between ground-dwelling and rock-dwelling species, except that in at least three ground-dwelling species, generation glands are already present at birth. In other species these glands differentiate only later in life. In the males of most species, generation glands start to differentiate prior to or at sexual maturity with a drastic increase in number when asymptotic body size is reached. The considerable inter- and intrapopulation variation in gland number in females, but not in males, underscores the hypothesis that the presence of generation glands in females is energy-linked and closely regulated by climate. An opposite geographic pattern of generation gland number in C. cataphractus, suggests that other factors than climate must be responsible for a difference in gland number and a difference in population density is suggested as explanation. To test for differences in aggression levels, females with and those without generation glands were staged in two experiments. In one experiment, females from different localities were staged against each other, while females from the same locality were paired in another experiment. No significant difference in aggression levels between females with and those without generation glands were observed in both experiments. It is suggested that generation glands do not exclusively function in territorial marking, but rather as a scent enhancer for femoral gland secretions. Inorder to elucidate the functional significance of generation glands, male response to female gland stimuli was investigated for 12 male C. cordylus. Males were presented with different female gland secretions (femoral, generation, and femoral and generation gland secretion combined) and water was used as a control. Total number of tongue-flicks and number of tongue-flicks directed at the secretion were counted. No significant difference was observed in the number of tongue-flicks between the different treatment groups. Males did, however, display leg-waving and head-rubbing behaviour. At present, the behaviour displayed by male C. cordylus in the presence of female gland stimuli, is unclear. The general absence of tongue-flicking of sandpaper labelled by female gland stimuli may be because of a flaw in the experimental design or because the experiment was conducted outside the breeding season. The display ofleg-waving and head-rubbing behaviour, however, suggests that cordylids do not exclusively rely on tongue-flicking for detection of conspecifics. The real function of generation glands thus still remains unresolved and further studies are needed to clarify this. / AFRIKAANSE OPSOMMING: Geografiese variasie in die aan-/afwesigheid van generasiekliere in wyfies van die cordylus-niger-oelofseni kompleks is beskryf en met klimaatsveranderlikes gekorreleer. Gepreserveerde en lewende materiaal, afkomstig van 96 lokaliteite in die area suid van 32°30' S en wes van 19°45'0 in die Wes-Kaap, Suid-Afrika, is vir die aan- ofafwesigheid van generasiekliere ondersoek. Generasiekliere is oor die algemeen afwesig in wyfies van westelike kuslokaliteite en die Kaapse Plooiberge, terwyl dit aanwesig is in wyfies van die binnelandse laagland lokaliteite. 'n GIS-analise is gedoen om die gemiddelde jaarlikse minimum temperatuur, gemiddelde jaarlikse maksimum temperatuur, potensiële verdamping, gemiddelde jaarlikse neerslag, gemiddelde jaarlikse mis en gemiddelde jaarlikse wolkbedekking, gemeet teen 14:00, vir elke lokaliteit te bepaal. Diskriminantefunksie en kanonieke analises toon 'n hoogs beduidende korrelasie tussen die aan- en afwesigheid van kliere en vyf van die ses klimaatsveranderlikes. Daar word voorgestel dat die hoë voorkoms van mis en lae temperature wat langs die weskus ervaar word as gevolg van die koue Benguela Seestroom, en die hoë voorkoms van orografiese mis en wolkbedekking in die Kaapse Plooiberge, die energie-toekenning in wyfies beïnvloed. Om hulle voortplantingsuitset te handhaaf, word wyfies moontlik gedwing om minder energie in sekondêre geslagskenmerke, soos generasiekliere, te belê. Interspesifieke variasie in die posisie, voorkoms, aantal en tyd van klierontwikkeling is vir verskeie gordelakkedisspesies ondersoek. Data vir verskeie spesies is vanuit die literatuur verkry, terwyl data vir ander spesies vanafmaterial in die Ellerman-versameling van die Universiteit van Stellenbosch verkry is. Cordylus cordylus en C. cataphractus is vir 'n evaluering van inter- en intrapopulasie variasie gebruik. Data vir C. cordylus is vanafvyfpopulasie wat op die kuslaaglande van die Wes-Kaap voorkom, ingesamel, terwyl data oor C. cataphractus vanuit die literatuur verkry is. Geslag, snoet-kloaak-lengte en die aantal generasiekliere is aangeteken vir 'n verteenwoordigende aantal individue van elke populasie. Terwyl generasiekliere in die femorale wyk van alle spesies aangetref word, het Cordylus giganteus en C. microlepidotus ook addisionele generasiekliere in ander dele van die liggaam. Daar is tot die gevolgtrekking gekom dat addisionele generasiekliere in hierdie spesies moontlik verwant is aan 'n groter behoefte aan chemiese kommunikasie in 'n lae sigbaarheidsomgewing in die geval van C. giganteus, en in 'n lae populasiedigtheid in die geval van C. microlepidotus. Generasiekliere word in alle gordelakkedis mannetjies aangetref, maar kan afwesig wees in wyfies van sommige spesies. Alle gordelakkedisse toon geslagsdimorfisme met betrekking tot die aanwesigheid en aantal generasiekliere, en mannetjies het beduidend meer kliere as wyfies, wat daarop dui dat hierdie kliere moontlik 'n meer belangrike rol by mannetjies as by wyfies speel. Lae geslagsdimorfisme inten minste twee spesies stel voor dat generasiekliere moontlik ewe belangrik in beide geslagte van hierdie spesies kan wees. In wyfies word die aanwesigheid van generasiekliere sterk deur klimaat gereguleer. Daar is geen aanduiding van verskille in generasieklierprofiel tussen grondbewonende en rotsbewonende spesies nie, behalwe dat generasiekliere reeds by geboorte teenwoordig is in ten minste drie grondbewonende spesies. In ander spesies ontwikkel hierdie kliere eers later. In mannetjies van die meeste spesies, begin generasiekliere voor oftydens geslagsrypheid ontwikkel, met 'n drastiese toename in aantal kliere wanneer asimptotiese liggaamsgrootte bereik word. Die aansienlike inter- en intrapopulasie variasie in aantal kliere in wyfies, maar nie in mannetjies nie, ondersteun die hipotese dat die teenwoordigheid van generasiekliere in wyfies aan energie gekoppel is en deur klimaat gereguleer word. 'n Teenoorgestelde geografiese patroon in die aantal generasiekliere in C. cataphractus, stel voor dat ander faktore as klimaat vir die verskil in aantal kliere verantwoordelik is en 'n verskil in populasiedigtheid word as verklaring voorgestel. Om vir verskille in aggressievlakke te toets, is wyfies met en wyfies sonder generasiekliere is teenoor mekaar getoets in twee eksperimente. In een eksperiment is wyfies van verskillende lokaliteite teenoor mekaar getoets, en in 'n ander eksperiment is wyfies van dieselfde lokaliteit teenoor mekaar getoets. Inbeide eksperimente is geen beduidende verskille is in die aggressievlakke van wyfies nie waargeneem nie. Daar word voorgestel dat generasiekliere nie uitsluitlik in gebiedsafbakening funksioneer nie, maar dat dit eerder as reukversterker vir die femorale kliere dien. Om die funksie van generasiekliere vas te stel, is die reaksie van 12 mannetjie C. cordylus op klierstimuli van wyfies getoets. Verskillende kliersekresies (femoraal, generasie, en femoraal en generasie gekombineerd) van wyfies is mannetjies voorgelê, en water is as 'n kontrole gebruik. Totale aantal tongskiete en aantal tongskiete wat na die sekresie gerig is, is getel. Geen beduidende verskil in die aantal tongskiete is tussen die verskillende groepe waargeneem nie. Mannetjies het egter beenwaai- en kopvryfgedrag getoon. Die rede vir die gedrag wat deur C. cordylus mannetjies in die teenwoordigheid van klierstimuli van wyfies vertoon word, is egter tans onduidelik. Die algemene afwesigheid van tongskiete na skuurpapier wat met klierstimuli van wyfies gemerk is, mag moontlik wees as gevolg van 'n fout in die eksperimentele ontwerp of omdat die eksperiment buite die paarseisoen uitgevoer is. Die vertoning van beenwaai- en kopvryfgedrag stel egter voor dat gordelakkedisse nie uitsluitlik op tongskiete staatmaak om lede van dieselfde spesie waar te neem nie. Die ware funksie van generasiekliere is dus steeds onopgelos en verdere studies word benodig om dit duidelikheid hieroor te kry.

The integument is the body organ interfacing between the internal and external environments of a lizard. This study explores aspects of its structure, texture, and coloration, relating them to survival strategies of horned lizards. The dermal chromatophore structure of Phrynosoma modestum is described. Color change depends on two cell types, melanophores and iridophores. This cellular arrangement may be typical of lizards utilizing color change for thermoregulation. Color pigment cells (xanthophores and erythrophores) function in pattern formation and background color matching. Experiments on the regulation of color change revealed that (1) -MSH is the prime skin darkening agent, (2) - and -adrenergic receptors also play a role in color change, and (3) temperature is a factor in in vitro and in vivo darkening and lightening responses. Apparently the primary role of color change is thermoregulation. Intraepidermal mechanoreceptors on dorsal body, limb, and head scales were studied, their ultrastructure is described, and their role in defense and survival is explored. The use of the integument for "rain-harvesting" of drinking water by P. cornutum is described, including stereotyped behavior, stereoscopic SEM examination of interscalar channels, experiments on capillary water flow, and an evaluation of micro-ornamentation. Comparisons are made with the putative use of the integument for the collection of rain, fog, or dew for drinking by Moloch and Phrynocephalus. A stereotypic behavior that utilizes positioning of occipital horns and dorsal scale armament is described and hypothesized to be an ophidian antipredator defense. Attacks by Onychomys torridus on P. cornutum and P. modestum were studied to further evaluate the role of occipital horns, dorsal scalation, and dermal collagen as antipredator defenses. Attack behaviors of the grasshopper mice and defense behaviors of the lizards are detailed. The use of color pattern, integumental structures, and mimetic behavior by P. modestum in avoidance of predator detection, through "stone-mimicry," is hypothesized and supported. Aspects of social communication in four species were studied. The importance of olfaction for intraspecific communication and lack of visual color signals is related to the need for chromatic crypticity in order to avoid detection by predators.

Cnemidophorus costatus barrancorum was studied in the short-tree forest of southeastern Sonora, Mexico from November 1985 to December 1988. Adult females varied in snout-vent length from 71 to 101 mm, while males reached a maximum of 121mm. Ontogenetic change in color-pattern was pronounced, however sexual dimorphism in color-pattern was slight. The mean number of dorsal scales around mid-body was 99.4. Mating occurred in late July. Females laid two clutches of eggs per season, with clutch size varying from 2-8 (X = 4.2). The maximum shelled egg dimensions were 10.5mm x 15.6mm. Hatchlings appeared in early fall, and were surface-active throughout November. Yearlings emerged from hibernation in February-March. Adults were active from mid-spring into September. C. c. barrancorum had a unimodal daily activity pattern, with activity beginning when air temperature exceeded 30°C. The mean body temperature of surface-active animals was 40.0°C.

Baited pitfall traps were used to sample the lizard fauna at Birdlings Flat on Kaitorete Spit, Canterbury. Four species had been recorded previously from this area; Leiolopisma maccanni Patterson and Daugherty, Leiolopisma nigriplantare polychroma Patterson and Daugherty, Leiolopisma lineoocellatum (Dumeril and Dumeril) and Hoplodactylus maculatus (Gray). Three of these species (L. maccanni, L. n. polychroma and H. maculatus) were captured during the course of the study. The aim of this present study was to examine the nature of the ecological relationship among these three species at Birdlings Flat. Capture data indicated that L. maccanni was almost entirely confined to the dunelands while L. n. polychroma was associated exclusively with shrublands on old dune ridges behind the sand dunes. H. maculatus' distribution encompassed both of these major habitats. Separation on the basis of habitat was thought to be the most important niche variable for these two diurnal skinks. There was some temporal separation in activity of these two species, with L. maccanni active earlier in the day than L. n. polychroma. However, there was a high degree of overlap in the activity periods of these two species. Temporal differentiation between the nocturnal gecko H. maculatus and the two diurnal skinks is thought to be an important means by which these species coexist. The most common prey items consumed by all three species were Diptera, Araneae, C. propinqua seeds, Hemiptera, unidentified arthropod eggs, Coleoptera and Hymenoptera. Dietary differences between the two skink species were apparent although these differences appeared to be related to the preferred habitats of the respective species. Density estimates for L. maccanni varied between 1050/ha and 1850/ha while L. n. polychroma density varied between 200/ha and 400/ha. The density of H. maculatus was not calculated but appeared to be intermediate to the density of the two skink species. The apparent disappearance of L. lineoocellatum from an area where they were once relatively common is cause for concern. There is no obvious reason for this decline although it may be related to the combined impacts of predation, collection and habitat disturbance.

Habitat loss is one of the leading causes of species decline across all taxa and conservation practices require information on population trends. The Mescalero Sands ecosystem, New Mexico, USA, is experiencing landscape changes associated with oil and gas development. The dune-dwelling lizard community contains a habitat specialist, Sceloporus arenicolus, that is of particular interest because it has a very limited geographic distribution that is entirely subject to oil and gas development. Distance sampling is widely used to estimate population densities of many vertebrate taxa however assumptions can be difficult to satisfy with certain species or in certain habitats. Researchers must investigate the likelihood that assumptions can be satisfied before implementing any population sampling method. I had two objectives. First to investigate the precision of population densities of dune-dwelling lizards estimated via distance sampling that was coupled with double-observer surveys. Second to compare abundances of dune-dwelling lizards among sites that varied in oil and gas development. I conducted distance line transects and compared those density estimates to densities obtained from total removal plots. I quantified the amount of oil and gas development, habitat quantity and quality and correlated those to lizard abundances to investigate the effects of oil and gas development on lizard populations. I found large differences in density estimates from distance sampling and total removal plots that resulted from violation of distance sampling assumptions. Although distance sampling is a low cost method, it does not produce reliable density estimates for dune-dwelling lizards and is not an appropriate sampling method in this system. I did not find oil and gas development effects on the habitat quantity, quality or on the abundances of lizards. Lizard abundances were most strongly correlated to habitat quantity. Lizard abundances may be influenced by complex interactions between oil and gas development and habitat quantity and quality but controlling for those interactions was beyond the scope of my study. Before and after experiments and long-term studies at multiple sites would be required to more fully address the effects of oil and gas development on lizard populations in the Mescalero Sands.

Habitat loss is one of the leading causes of species decline across all taxa and conservation practices require information on population trends. The Mescalero Sands ecosystem, New Mexico, USA, is experiencing landscape changes associated with oil and gas development. The dune-dwelling lizard community contains a habitat specialist, Sceloporus arenicolus, that is of particular interest because it has a very limited geographic distribution that is entirely subject to oil and gas development. Distance sampling is widely used to estimate population densities of many vertebrate taxa however assumptions can be difficult to satisfy with certain species or in certain habitats. Researchers must investigate the likelihood that assumptions can be satisfied before implementing any population sampling method. I had two objectives. First to investigate the precision of population densities of dune-dwelling lizards estimated via distance sampling that was coupled with double-observer surveys. Second to compare abundances of dune-dwelling lizards among sites that varied in oil and gas development. I conducted distance line transects and compared those density estimates to densities obtained from total removal plots. I quantified the amount of oil and gas development, habitat quantity and quality and correlated those to lizard abundances to investigate the effects of oil and gas development on lizard populations. I found large differences in density estimates from distance sampling and total removal plots that resulted from violation of distance sampling assumptions. Although distance sampling is a low cost method, it does not produce reliable density estimates for dune-dwelling lizards and is not an appropriate sampling method in this system. I did not find oil and gas development effects on the habitat quantity, quality or on the abundances of lizards. Lizard abundances were most strongly correlated to habitat quantity. Lizard abundances may be influenced by complex interactions between oil and gas development and habitat quantity and quality but controlling for those interactions was beyond the scope of my study. Before and after experiments and long-term studies at multiple sites would be required to more fully address the effects of oil and gas development on lizard populations in the Mescalero Sands.

Thesis (Ph. D.)--University of Queensland, 2003. / Includes bibliographical references.

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