The transition between terrestrial varanoid lizards and aquatic mosasaurs through the intermediate, semi-aquatic aigialosaurs is fully documented. Aigialosaurs are shown to possess a mosaic of mosasaurian (configuration of the skull, jaw and tail) and terrestrial varanoid characters (appendicular skeleton and trunk). / The taxonomic position of the Aigialosauridae within the superfamily Varanoidea is evaluated. Based on character states previously used to define the Varanoidea, neither the specific affinities of aigialosaurs nor the sister-group relationships of earlier members of the terrestrial varanoid assemblage can be securely established. For this reason, the specific character states involved have been reexamined and alternative hypotheses of relationship have been considered.

No description available.

[Truncated abstract] In ecomorphological or ecophysiological studies, variation in `design? traits (e.g. size, morphology and physiology) is thought to determine variation in ecologically-relevant performance traits, which in turn determines fitness in a particular habitat (Arnold 1983). Thus, natural selection is thought to act most directly on intermediate traits such as measures of locomotory performance. This thesis examined this process in the closely related group of Australian varanids lizards (Squamata: Varanidae). Phylogenetically, varanids are divided into three major clades. Size (mass and snout-to-vent length) is strongly correlated with these three clades. Two clades, (Gouldii and Komodoensis) are large, while the third clade (Odatria) has a smaller body size. Thus, there is considerable variation in size for various species. Size varied for species by three orders of magnitude. Size is also related to two ecological characteristics, foraging mode and habitat openness. Widely-foraging species were larger than sit-and-wait strategists, while species from open habitats were larger than species from semi-open or closed habitats. However, given the tight link between size and phylogeny we cannot separate adaptation of size to ecological traits from that of phylogenetic patterns. Of interest throughout this thesis was how variations in design (e.g. morphology and physiology) were related to ecological characteristics. Since body size also influences many of these morphological and physiological characteristics it is often necessary to remove the effects of size. Three design traits were examined in detail: body dimensions, vertebral number and metabolic rates. …Curiously, no performance variable linked differences in size-free body dimensions to retreat sites. This suggests that there is either a direct link between design and ecology (e.g. dorso-ventral compression), or some unmeasured performance variable related to retreat site. Given that most performance traits are thought to involve kinematic movements of the hindlimb limb and pelvis, and these were not best related to retreat site, then a direct link between design and ecology with respect to retreat site seems possible. In summary, this thesis provides evidence not only for links between design and ecology mediated by locomotory performance traits, but also direct links between design and ecology, for Australian varanid lizards.

Thesis (Ph.D.)--Australian National University, 2004.

Thesis (Ph.D.)--Ohio University, August, 2002. / Title from PDF t.p. Includes bibliographical references.

Thesis (MSc) -- Stellenbosch University, 2004. / ENGLISH ABSTRACT: Standard histological procedures revealed that there are two types of glands in the cloacal complex of the viviparous Cordylus cataphractus. These occur in the urodeal region of the complex (called urodeal glands), and in the posterior proctodeum region (called proctodeal glands). Urodeal glands are found in females only and differentiate at sexual maturity. Proctodeal glands occur in both sexes and the dorsal and ventral proctodeal glands are identical in structure. Seasonal activity in cloacal gland structure and secretory was studied. Urodeal glands show distinct seasonal variation in development and secretory activity, they become enlarged in vitellogenic females, remain active during pregnancy, with glandular activity peaking around parturition. Proctodeal glands, in contrast, show little or no seasonal variation in development or secretory activity in both sexes. Secretion quantity, however, is highly variable among individuals of the same sex, time of year, as well as reproductive stage, but evidently much less in females than in males. Although the possible functions of the cloacal glands are still unclear, two main functions are suggested: a) mechanical function such as lubrication or a substrate for genital products and b) pheromonal communication. I propose that urodeal glands are involved in some reproductive function rather than in social communication, because very little secretion is found during the mating season and peak glandular activity are evident around parturition. The opposite may be true for proctodeal glands, but further investigation is needed, particularly chemical analysis of secretion and behavioural manipulation, to determine the role of these glands in social communication. / AFRIKAANSE OPSOMMING: Standaard histologiese tegnieke dui aan dat daar twee tipes kliere voorkom in die kloakale kompleks van die lewendbarende akkedis, Cordylus cataphractus. Die een kliertipe word in die urodeale omgewing aangetref en staan bekend as die urodeaalkliere, terwyl die ander groep in die posterior area van die proktodeum voorkom en proktodeaalkliere heet. Urodeaalkliere word slegs in wyfies aangetref en ontwikkel eers na volwassenheid bereik is. Proktodeaalkliere kom in albei geslagte voor en dorsale en ventrale aspekte van hierdie kliere is identies aan mekaar. Die seisoenale aktiwiteit in struktuur en sekresie-aktiwiteit van hierdie kloakale kliere is ondersoek in mannetjies en wyfies wat gedurende verskillende maande van die jaar versamel is en dus in verskillende voorplantingstadia was. Urodeaalkliere het duidelike seisoenale variasie vertoon wat beide ontwikkeling van die klier en sekretoriese aktiwiteite betref. Die klier vergroot tydens vitellogenese, bly aktief tydens swangerskap en bereik maksimale aktiwiteitsvlakke rondom geboorte. In teenstelling hiermee het die proktodeaalkiere van beide geslagte weinig of self geen seisonale variasie in hul ontwikkeling of sekretoriese aktiwiteit getoon nie. Die hoeveelheid sekreet vrygestel was merkbaar minder in wyfies as in mannetjies, maar hoogs veranderlik tussen individue van dieselfde geslag of voortplantingstadium; of wat in dieselfde tydperk van die jaar versamel is. Alhoewel funksies van kloakale kliere steeds onduidelik bly, word twee hooffunksies voorgestel: a) ‘n meganiese funksie, byvoorbeeld as smeermiddel of as ‘n substraat vir geslagsprodukte en b) ‘n kommunikasie funksie, byvoorbeeld in die vrystelling van feromone. Ek stel voor dat die urodeaalkliere eerder by ‘n voortplantingsfunksie betrokke is, aangesien baie min sekreet gedurende die paarseisoen vervaardig word terwyl klieraktiwiteit rondom geboorte ‘n piek bereik. Die teenoorgestelde mag geld vir proktodeaalkliere, maar vereis verdere ondersoek. Veral ‘n chemiese analise van die sekresie en gedragsmanipulasie sal nuttig wees om ‘n moontlike rol van hierdie kliere in sosiale kommunikasie te bepaal.

Thesis (MSc)--University of Stellenbosch, 2000. / ENGLISH ABSTRACT: Cordylus giganteus is the largest lizard species of the family Cordylidae and is restricted in distribution to the highveld grasslands of the Free State, South Africa. Previous work on life history and physiological ecology suggested the need for further investigation into the social structure of C. giganteus, with the aim of improving knowledge on South African herpetofauna and contributing towards better conservation plans. Observations, recaptures, behavioural experiments and chemical analyses were made to investigate chemical communication, movements around and between burrows, spatial distribution and response to intruders. Sexual variation was found in both femoral gland proteins and lipids, while seasonal variation was found in lipids. Femoral gland proteins do not vary intra-individually. These results suggest alternate roles in communication for femoral gland proteins and lipids. Female C. giganteus showed a possible discriminatory ability between their own femoral gland secretion and that of other individuals. Burrow movements were characterised by very little time spent between burrows, high burrow fidelity and limited dispersal during seasons, other than movements associated with mating activity. Lizards remained in very close proximity to burrows. Male and female burrows were distributed in a clumped fashion, while male burrows were distributed randomly and female burrows were distributed randomly with respect to one another. There was a high degree of spatial association of male and female burrows. Male and female resident lizards responded aggressively to experimentally introduced conspecifics of the same sex, but exhibited no differential response towards neighbours or non-neighbours. Males and females exhibited similar levels of aggression towards intruders. The social system of C. giganteus can be defined as site defence, with very low frequencies of agonistic interactions. The social system of C. giganteus seems to be well suited to other aspects of its life history. / AFRIKAANSE OPSOMMING: Cordylus giganteus is die grootste akkedis spesie van die familie Cordylidae en is beperk in verspreiding tot die hoeveld graslande van die Vrystaat, Suid Afrika. Vorige werk oor die lewensgeskiedenis en fisiologiese ekologie het aangedui dat verdere inligting oor die sosiale struktuur van C. giganteus nodig is, met die doelom kennis oor Suid-Afrikaanse herpetofauna te verbeter, en om beter bewaringsstrategieë daar te stel. Observasies, hervangs, gedragseksperimente en chemise analyses was uitgevoer om aspekte oor kommunikasie, bewegings tussen en om gate, gat verspreiding en reaksie teenoor indringer akkedisse, te bestudeer. Femorale klier afskeidings het seksuele variasie in beide die proteiene en die lipiede getoon, terwyl net die lipide seisoenale variasie getoon het. Femoral klier proteiene het nie seisoenale variasie binne individue getoon nie. Hierdie resultate dui op alternatiewe funksies vir die femoral klier proteiene en lipiede. Wyfie C. giganteus het moontlik tussen hule eie femoral klier sekreet en die van ander wyfies onderskei. Baie min tyd tussen gate, hoë lojaliteit teenoor gate en beperkte bewegings weg van gate het gat bewegings gekarakteriseer. Net gedurende die paarseisoen was daar meer bewegings tussen gate. Andersins, het akkedisse baie naby hul eie gate gebly. Mannetjie en wyfie gate was naby mekaar versprei, terwyl mannetjies ewekansig teenoor mekaar versprei was, en wyfies ewekansig teenoor mekaar versprei was. Die verspreiding van mannetjies en wyfies het baie van die verspreiding van die ander geslag afgehang, sodat daar 'n groot assosiasie tussen hulle was. Akkedisse wat gate bewoon het, was baie aggressief teenoor indringer akkedisse, maar het geen verskil in reaksie teenoor naburige akkedisse of nie-naburige akkedisse getoon nie. Mannetjies en wyfies het dieselfde aggressiwiteit teenoor indringers getoon. Die sosiale struktuur van C. giganteus kan geklassifiseer word as skuilings-beskerming, maar met baie lae frekwensies van sosiale interaksies. Dit blyk dat die sosiale struktuur van C. giganteus baie goed pas by ander aspekte van sy lewensstyl.

Thesis (MSc)--University of Stellenbosch, 2001. / ENGLISH ABSTRACT: Cordy/ us cataphractus and C macropholis naturally occur in groups on a year round basis. I examined whether limited shelter availability might be responsible for this phenomenon. Individuals were provided with an excess of shelter sites and the observed pattern of shelter occupation was contrasted to that obtained for the solitary living species C polyzonus. Cordy/us cataphractus consistently occupied fewer shelters than its two congeners. Results suggest that grouping behaviour in this species is not the result of a limitation in refuge sites. In contrast, the pattern of shelter occupation by C macropholis did not differ significantly from that of C polyzonus, thus the aggregative behaviour observed in the former species may partially be attributed to limited shelter availability in its natural habitat. Conspecific recognition by means of pheromonal detection has been reported in many lizard families, but only for one member of Cordylidae, namely Cordy/us cordy/us. Cordylus cataphractus individuals were exposed to ceramic tiles that had been labeled by conspecific male and female substrate deposits. Labeling was achieved by using ceramic tiles as the bottom half of a shelter in a lizard's home cage for ten consecutive days. Washed tiles were used as controls. Labeled tiles did not elicit significantly more tongue-flicks than control tiles from test specimens in a series of trials. The apparent lack of conspecific recognition via pheromonal detection might be attributable to the group-living nature of this species, which primarily use visual cues for identification purposes. Tongue-flicking is generally utilized in C cataphractus in the detection of novel stimuli. Spatial dynamics of the group-living cordylid, C cataphractus, were investigated by studying patterns of intergroup movement. To date, literature has assumed that C cataphractus groups represent family units, implying low or delayed dispersal rates and high group fidelity. A mark-recapture experiment on six groups included three recaptures outside of and three recaptures within the mating season. High percentages of male, female and juvenile lizards left their groups, but might be an artifact of microhabitat disturbance. High proportions of males, females and juveniles entering the groups, both outside of mld within the mating season, clearly implied a high rate of intergroup movement. Activity and foraging patterns were studied in group-living C. cataphractus lizards by conducting direct field observations between September 1998 and October 1999. The influence of general weather patterns as well as the effect of group size on these patterns were investigated by observing a small-sized, medium-sized and large group. Each group was observed for a minimum of three sunny days per season, with scans conducted every 30 minutes between the hours of 08:00 and 20:00. There was a high rate of activity during the mating season in spring (September). Activity declined sharply mld foraging ceased at the onset of warm, dry conditions (February). Virtually no activity occurred just prior to the first winter rains (July), after which the lizards emerged to forage and replenish energy stores before onset of the next mating season. Selective inactivity might be employed to reduce energy expenditure in this group-living lizard, where intraspecific competition is stringent. Groups of different sizes display different thresholds at which it becomes energetically viable for group members to emerge, as well as different patterns of spatial use around their respective home crevices. Members of larger groups perched further from their crevices, ran further to catch prey mld had less potential prey capture events per individual. Lizards in the larger groups also displayed sporadic lengthy foraging excursions. Such movements might explain the evolution of tail-biting behaviour and heavy armour, which are defensive strategies associated with open areas. / AFRIKAANSE OPSOMMING: Cordylus cataphractus en C. macropholis word dwarsdeur die jaar in groepe in hul natuurlike habitat aangetref. Daar is nagevors of 'n beperking in aantal beskikbare skuilplekke vir hierdie verskynsel verantwoordelik is. Individue is van 'n oormaat skuilings voorsien en die waargenome patroon van skuilplekbesetting is gekontrasteer met dié verkry vir '11 enkelwonende species, naamlik C. polyzonus. Cordylus cataphractus het voortdurend minder skuilplekke beset as sy kongeneriese species. Groepsgedrag in C. cataphractus is dus nie die resultaat van 'n tekort aan beskikbare skuilplekke nie. In teenstelling hiermee het die patroon van skuilplekbesetting vertoon deur C. macropholis nie noemenswaardig verskil van dié van C polyzonus nie. Dus mag die groepsgedrag wat gewoonlik in C. macropholis waargeneem word 'n uitvloeisel wees van 'n beperking op die beskikbaarheid van skuilplekke in sy natuurlike habitat. Herkenning van speciesgenote deur waarneming van ferornone is vir verskeie akkedisfamilies gerapporteer, maar slegs vir een lid van Cordylidae, naamlik C. cordylus. Cordylus cataphractus individue is blootgestel aan keramiekteëls wat gemerk is met substraat-neerleggings van manlike en vroulike speciesgenote. Sodanige merking is uitgevoer deur die keramiekteëls vir tien dae lank te gebruik as die onderste helfte van akkedisse se skuilplekke in hul tuishokke. Gewaste teëls is as kontroles gebruile Gemerkte teëls het nie statisties beduidend meer tongskiete van die akkedisse uitgelok as kontrole teëls in 'n reeks toetse nie. Die klaarblyklike afwesigheid van speciesgenootherkenning deur middel van feromoon waameming kan moontlik toegeskryf word aan die groeplewende gedrag van C. cataphractus, wat primêr visuele informasie vir identifikasie gebruik. Cordylus cataphractus gebruik tongskiete oor die algemeen vir die waarneming vannuwe omgewingsstimuli. Die ruimtelike dinamika van 'n groeplewende gordelakkedis, C. cataphractus, is ondersoek deur die patroon van intergroepbeweging na te vors en groepsgetrouheid te meet. Tot op datum is daar in die literatuur aanvaar dat C cataphractus groepe familie-eenhede verteenwoordig, wat lae of vertraagde verpreidingstempo' sasook' n hoë mate van groepsgetrouheid impliseer. 'n Merk-hervang eksperiment is op ses groepe uitgevoer, insluitend drie hervang episodes buite en drie binne die paarseisoen. 'n Hoë persentasie manlike, vroulike en onvolwasse akkedisse het hul groepe verlaat, maar dit mag bloot 'n uitvloeisel van versteuring van die mikrohabitat wees. 'n Hoë proporsie mannetjies, wyfies en onvolwassenes het egter by die groepe aangesluit, beide buite en binne die paarseisoen, wat onteenseglik op 'n hoë mate van intergroep beweging dui. Daar kom derhalwe veel vryer bewegingspatrone in hierdie species voor as voorheen vermoed en dit is hoogs onwaarskynlik dat groepe familie-eenhede verteenwoordig. Aktiwiteits- en voedingspatrone van die groeplewende akkedis, C cataphractus, is bestudeer in 'n reeks veldobservasies tussen September 1998 en Oktober 1999. Afgesien van die invloed van algemene weerspatrone. is die effek van groepgrootte op hierdie patrone ondersoek deur 'n klein, medium en groot groep te observeer. Elke groep is waargeneem oor . 'n minimum periode van drie dae per seisoen met opnames elke 30 minute, tussen 08:00 ten 20:00. 'n Hoë aktiwiteitsvlak is gedurende lente tydens die paarseisoen gehandhaaf (September). Met die aanvang van warm en droë kondisies (Februarie) het aktiwiteitsvlakke skerp begin daal en voedingsgedrag is gestaak. Feitlik geen aktiwiteit was waarneembaar kort voor die eerste winterreëns (Julie) nie, waamá akkedisse intensiefbegin voed het om energiestore op te bou voor die aanvang van paartyd. Selektiewe onaktiwiteit verminder moontlik energieverbruik in hierdie groeplewende akkedis, waar intraspesifieke kompetisie streng is. Verskillende groepgroottes beskik oor verskillende perke waar dit vir die individu energeties voordelig sou wees om uit die skeur te kom, asook 'n variasie in ruimtelike gebruik om die skeur. Individue van die twee groter groepe het nader aan hul rotsskeur gesit, het verder van die skeur af prooi vang en het beduidend minder potensiële voedingskanse gehad. Akkedisse in die groter groepe het voorts sporadiese lang voedingsekskursies getoon. Hierdie bewegings mag die evolusie van stertbytgedrag en 'n swaar pantser, verdedigingstrategieë wat met oop areas geassosieer word, verduidelik.

published_or_final_version / Zoology / Doctoral / Doctor of Philosophy

published_or_final_version / Anatomy / Doctoral / Doctor of Philosophy