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, 2004. / ENGLISH ABSTRACT: There is overwhelming evidence that the Armadillo Lizard, Cordylus cataphractus, forms permanent aggregations, and that termites are possibly the most important component of the diet of this species. In addition, the spinose morphology and defensive tail-biting behaviour displayed by this lizard species strongly imply that individuals move away from the crevice, where they are more vulnerable to predation. Therefore the aim of this part of the study was to investigate whether C. cataphractus harvest termites at the termite foraging ports and to discuss the likely ecological implications of termitophagy for this species. A quadrate at the Graafwater study site, including several crevices housing C. cataphractus groups, was measured out. All the foraging ports of the subterranean harvester termite (Microhodotermes viator) present in the quadrate, were located and their positions recorded in respect to the distance from the nearest crevice housing lizards. The presence of C. cataphractus tracks at the foraging ports was used to verify whether individuals visited specific termite foraging ports. Tracks were found at foraging ports located at an average distance of 6.1 m, but were also located at foraging ports up to 20 m from the nearest crevice. The results strongly support the hypothesis that C. cataphractus individuals move away from the crevice to feed on termites. Termitophagy and the heavy armature of C. cataphractus appear to be key factors in the group-living behaviour of this species. Termitophagy allows individuals to live in groups as competition for food at the home crevice will be limited. Theheavy armature of C. cataphractus renders it clumsy and slow-moving, compared to other cordylids. Group-living will, however, eliminate juvenile dispersal and mate-searching excursions, activities which, in the light of the heavy morphology of the species, may result in high predation impact. The global objective of the study was to investigate social structure and space-use in Cordylus cataphractus. For the correct interpretation of data on the latter, it was necessary to first establish an ethogram for the study animal. Members of a multi-male group located at the Graafwater site, including all age-sex classes, were observed in the field by telescope, for a total of 102 hours. All individuals were observed to predominantly display maintenance behavioural act systems, however, agonistic act systems were also observed. Aggression levels were determined for the respective agonistic behavioural acts observed. Males were highly aggressive and displayed the most aggression (67.42 %). Females and sub-adults displayed aggression for 34.97 % and 37.75 % of the total observation time, respectively. Juveniles did not display aggression at all. Mating behaviour was also observed and described. An ethogram was thus established for the first time for C. cataphractus, and was compared to the ethograms available for a few other cordylids. The spatial distribution of individuals within a population has an important impact on the social structure that ensues. Living in a group imposes several pressures on the individual and competition for mates is one. The main aim of the study was to determine the space-use of all theindividuals included in a multi-male Cordylus cataphractus group (n = 55), to elaborate on the mating system. Specific focus was on the adult males and females of the multi-male group, because, ultimately, the manner in which individuals, particularly the adult males and females, utilise space, determines the mating system. The mating system is an important aspect of social structure. A two-dimensional grid pattern was delineated at a free-living, multi-male group’s crevice, situated at the Graafwater study site, to be able to plot the position of each individual during observations. All marked individuals, that were visible, were scanned through a telescope and their respective positions were recorded at 30-minute intervals. Arcview 3.2 Geographical Information Software was used to create a computerised model replicate of the two-dimensional grid pattern at the crevice, and to graphically depict the recorded positions for each individual. The space-use of males overlapped greatly with the space-use of females (66.12 ± 31.81 %), but a 0 % overlap in space-use was observed among adult males. The space-use of sub-adults and juveniles overlapped highly among one another, as well as with the space-use of adult males and females. The results showed conclusively that adult male C. cataphractus are territorial, defending specific sites at the crevice. The territory of each male included in the multi-male group incorporated at least one female. The space-use of several females incorporated the space-use of two or more males. Males were observed to mate with one or more females, and one female was observed to mate with two different males on more than one occasion. Thus the evidence strongly suggests that C. cataphractus males display typical territorial polygyny whereas females may be promiscuous. / AFRIKAANSE OPSOMMING: Daar is oorweldigende bewyse dat die pantsergordelakkedis, Cordylus cataphractus, permanent in groepe bly en dat termiete waarskynlik die belangrikste komponent in die dieet van hierdie spesie is. Die stekelrige morfologie en die stert-byt beskermingsgedrag is verder sterk aanduidings dat hierdie akkedisse tyd weg van hul klipskeure spandeer waar hulle meer gevoelig vir predasie is. Die doelwit van die eerste deel van die studie was om te bepaal of C. cataphractus termiete by die voedingspoorte van die termietneste oes en verder ook om die moontlike ekologiese implikasies wat termitofagie vir die spesie inhou, te bespreek. ‘n Kwadraat was by die Graafwater studielokaliteit uitgemeet om verskeie akkedisgroepe in te sluit. Al die voedingspoorte van die suidelike grasdraer termiet (Microhodotermes viator) binne die kwadraat is gevind en die afstande na die naaste klipskeur wat akkedisse huisves, is vir elke voedingspoort bepaal. Die teenwoordigheid van C. cataphractus spore in die sand by die voedingspoorte was gebruik om besoeke aan spesifieke voedingspoorte te bevestig. Spore is by voedingspoorte gemiddeld 6.1 m vanaf die naaste akkedisgroep gevind, maar tot so ver as 20 m vanaf die naaste akkedisgroep. Die resultate verleen sterk ondersteuning vir die hipotese dat C. cataphractus individue op termiete weg van hul klipskeur af voed. Termitofagie en die swaar pantser van C. cataphractus is waarskynlik sleutelfaktore in die groeplewe-gedrag van die spesie. Termitofagie laat groeplewe toe omdat kompetisie vir voedsel by die klipskeur beperk sal wees. Die swaar pantser van C. cataphractus het tot gevolg dat die spesie lomp en stadig is in vergelyking met ander gordelakkedisse. Groeplewe sal egter natale spreiding en maatsoek-ekskursies, aktiwiteite wat in die lig van die lompheid van die spesie swaar predasie impak tot gevolg kan hê, uitskakel. Die oorhoofse doelwit van die studie was om die sosiale struktuur en spesifiek ruimtebenutting by C. cataphractus te ondersoek. Vir korrekte interpretasie van data oor lg. was dit egter eers nodig om ‘n etogram vir die studiedier saam te stel. Lede van ‘n multi-mannetjie groep, wat alle ouderdomsgrootte klasse ingesluit het, is by die Graafwater studielokaliteit m.b.v. ‘n teleskoop vir ‘n totaal van 102 uur dopgehou. Al die individue in die groep het oorwegend instandhoudingshandelingsisteme vertoon, alhoewel agonistiese handelingsisteme ook vertoon is. Aggressievlakke is vir die verskeie agonistiese gedragshandelings bepaal. Mannetjies was hoogs aggressief en het die meeste aggressie vertoon (67.42 %). Wyfies en subvolwassenes het aggressie vir 34.97 % en 37.75 % van die totale observasietyd respektiewelik getoon. Jong individue het geen aggressie getoon nie. Paringsgedrag was ook waargeneem en beskryf. ‘n Etogram is dus vir die eerste keer vir C. cataphractus opgestel en is ook vergelyk met etogramme van ander gordelakkedisse. Die ruimtelike verspreiding van individue binne ‘n populasie het ‘n belangrike invloed op die sosiale struktuur wat daaruit voortvloei. Om in ‘n groep te lewe, plaas verskeie drukke op die individue binne die groep en kompetisie vir maats is een hiervan. The hoofdoelwit van die studie was om die ruimtebenutting van al die individue binne ‘n multi-mannetjie groep van C. cataphractus (n = 55) te bepaal en om die paringsisteem in so ‘n groep te ondersoek. Die spesifieke fokus was op die volwasse mannetjies en wyfies, omdat die paringsisteem deur die wyse waarop volwasse individue ruimte benut, bepaal word. Die studie is by die Graafwater studielokaliteit gedoen. Die oppervlak buite ‘n klipskeur wat deur ‘n multi-mannetjie groep bewoon word is opgedeel in sektore om die kartering van die posisies van die individue tydens observasietye moontlik te maak. Al die individue is vooraf met verfkodes langs die kant van die kop gemerk en op gegewe observasiedae is individuele posisies elke 30 min aangeteken. Arcview 3.2 Geographical Information Software was gebruik om ‘n tweedimensionele, gerekenariseerde replikaat van die klipskeur en onmiddelike omgewing te skep en om die waargenome posisies van individue grafies voor te stel. Die ruimtebenutting van mannetjies het grootliks met dié van wyfies oorvleuel, terwyl zero oorvleueling tussen mannetjies waargeneem is. Ruimtebenutting van subvolwasse en jong individue het ook grootliks onderling oorvleuel en ook met dié van volwasse mannetjies en wyfies. Die resultate toon duidelik aan dat volwasse mannetjies territoriaal is en dat spesifieke standplase rondom die klipskeur verdedig word. Die territorium van elke volwasse mannetjie het of ‘n gedeelte of die hele ruimtebenutting van ten minste een volwasse wyfie ingesluit. Ruimbenutting van verskeie wyfies het met dié van twee of selfs meer mannetjies oorvleuel. Mannetjies is waargeneem om met meer as een wyfie te paar en ten minste een wyfie het met meer as een mannetjie gepaar. Die resultate dui dus sterk daarop dat C. cataphractus mannetjies tipies terrotoriale poligenie vertoon terwyl wyfies weer promisku is.