The behavioral repertoire of the Puerto Rican anoles, Anolis cooki, A. cristatellus and A. monensis was studied in the laboratory. The display of A. cooki and A. monensis revealed great similarities; consisting of one head bob pattern highly stereotyped in its temporal sequencing and total duration, sometimes accompanied by non-stereotyped dewlap movements. A. cristatellus display behavior consisted of two distinct head bob patterns (Type A and Type B). The A display is strongly stereotyped and used in the assertion and courtship contexts, while B display is less stereotyped and appeared only in the challenge context. Similar display modifiers used during aggressive interactions were found in A. cooki, A. cristatellus, A. monensis and two other Puerto Rican anoles, A. evermanni and A. gundlachi. In all these species the modifiers were progressively added during prolonged aggressive encounters and seemed to indicate the increasing arousal of the combatants. A grading scale was designed to measure the level of aggression in conspecific and congeneric interactions. It was based on the different behaviors that the lizards used during their agonistic encounters. The scale assigned a numerical value to each behavior according to their order of appearance in the sequence of behaviors performed in the encounter. Thus, a numerical value was derived for the level of aggression exhibited in each interaction, and that was called the "aggression index" (AI). The AI of A. cooki and A. cristatellus conspecific interactions was significantly higher than the AI of their congeneric interactions. The high AI of the cooki-cristatellus interactions was not explained by Murray's (1971) "mistaken identity argument", but as a function of their wide overlap in resource requirements. / Master of Science

Thesis (MSc (Botany and Zoology))--University of Stellenbosch, 2006. / The role of the southern harvester termite, Microhodotermes viator, and several climatic parameters in the distribution of the group-living lizard, Cordylus cataphractus, was investigated. Microhodotermes viator is considered the most important prey item of C. cataphractus and termitophagy as the causative agent in the evolution of group-living in this species. One would therefore expect a high degree of correspondence in the ranges of C. cataphractus and M. viator. As climate will also play a role in the distribution of any species, various climatic variables were investigated to determine their influence on the distribution of C. cataphractus. Species distributions were visualized using the minimum polygon technique and the degree of overlap was determined using standard geographic information systems (GIS) techniques. A total of 53 C. cataphractus localities were investigated for the presence of termites. The climatic limits of the geographical distribution of C. cataphractus were investigated by means of three models, namely Classification Trees, General Discriminant Analysis and Logistic Regression. The range of C. cataphractus was completely included within the range of M. viator Microhodotermes viator was included in the diet of C. cataphractus at 73 % of the localities sampled within the lizard’s range. The current geographical range of C. cataphractus is mainly correlated with two climatic factors, namely the low summer rainfall and high monthly solar radiation. The restricting role of both these factors can be directly linked to the group-living nature of C. cataphractus. If termitophagy were the overarching cause of group-living in C. cataphractus, then one would expect a close relationship between termite density and lizard density and termite density and lizard group size. I investigated these relationships at both a local and regional scale. For the local scale study, 25 quadrats of 25 × 25 m were plotted at a selected site, and for the regional scale study, ten 35 × 35 m quadrats at sites throughout the lizard’s range were used. In each quadrat, a range of variables were recorded, the most important of which were lizard density, lizard group sizes, termite foraging port density, distance to nearest termite foraging ports, vegetation height and vegetation cover. I found that the density of termite foraging ports determines C. cataphractus density. Vegetation height and cover affects crevice selection by C. cataphractus groups, probably because an unobstructed view is necessary to locate termite activity at foraging ports. I also investigated possible differences in the use of termites by different sized groups of C. cataphractus during different times of the year. Faecal samples, collected once a month at Eland’s Bay from small, medium and large groups from January 2005 to December 2005, were analysed for the presence of termite head material. I found that large groups fed on termites to a greater extent than small groups during certain times of the year and there was a general tendency for this phenomenon throughout the year. The results collected in this study indicate that the southern harvester termite, M. viator, plays a central role in the ecology of the group-living lizard, C. cataphractus.

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.

Determining the past, present and future effects of alien species on native biodiversity is a globally recognized problem. The Wall Lizard, Podarcis muralis, was introduced to Vancouver Island from Europe in 1970. To assess potential competitive interactions with the native Northern Alligator Lizard (Elgaria coerulea), I investigated several aspects of P. muralis ecology. D i h i o n along manmade corridors (e.g. roads and powerlines) and human-facilitated jump dispersal are contributing to range expansion; three separate populations currently exist. Reproductive output of P. muralis was not affected by amount eaten while gravid, but feeding level and incubation temperature affected offspring phenotypes. In comparative tests of locomotor performance, P. muralis was faster than Z. coerulea, but only at moderate temperatures. I found no effect of P. muralis dour on E. coeruela behaviour, but E. coeruela avoided cover objects that housed P. muralis. These two species use similar habitats, but were found on different substrates and mesoslopes. Based on the results of these experiments there is a potential for comvetition between these two lizard species.

In ectothermic vertebrates, body segmentation is often linked to final body length. This thesis focuses on the realtionship between final body length, ecological (temperature) and genetic (sex) factors and the count of body vertebrae in the Paroedura geckos, closely focusing on the Paroedura picta species. Individuals of this species were exposed to three different constant rearing temperatures, where they achieved different final sizes. We tested the correlation between the number of presacral vertebrae and final body size and the effect of rearing temperature and sex on the vertebrae number. The data point to high genetic and enviromental canalisation of the vertebrae count. Large conservation in the number of presacral vertebae were found even across the genus Paroedura regardless of the considerable evolutionary changes in body size. Powered by TCPDF (www.tcpdf.org)

Orientador: Carlos Alberto Hussni / Coorientador: Carlos Roberto Teixeira / Banca: Marcos Jun Watanabe / Banca: Rogério Loesch Zacariotti / Resumo: As lesões cutâneas em répteis possuem diferentes causas e decorrem em cicatrizes e disecdise, predispondo a dermatites futuras, indesejáveis, podendo determinar a retirada do animal da exposição e levar ao abandono destes quando criados domesticamente. Com o objetivo de comparar diferentes produtos tópicos e procedimentos na busca da cicatrização da pele, foram utilizados doze lagartos adultos, machos, da espécie Tupinambis merianae. Em cada animal foram produzidas quatro lesões circulares na pele do dorso, dispostas em quadrilátero. Compostos três grupos de quatro animais, o Grupo 1 compreende o comparativo entre solução fisiológica e policresuleno; no Grupo 2, tintura de iodo a 2% e o não tratamento, considerado como controle; o Grupo 3 clorexidine e o enxerto autólogo de pele. As lesões do lado direito destinaram-se à análise macroscópica diária, com planimetria a cada sete dias e observação do tempo total de cicatrização. Das lesões esquerdas, foram coletadas amostras para biópsias e conseqüentes exames histopatológicos, a cada sete dias. Os lagartos apresentaram processo cicatricial semelhante aos mamíferos, notoriamente mais lento, entretanto. O enxerto não se mostrou vantajoso neste experimento, sofrendo necrose nos quatro animais. O policresuleno destacou-se por impermeabilizar a ferida em 4 dias, enquanto que os demais grupos obtiveram o mesmo efeito entre 8 e 12 dias. As lesões se mostraram completamente epitelizadas entre 40 e 53 dias, não havendo diferença entre os grupos quanto ao tempo total de cicatrização, retração tecidual e aparência da cicatriz / Abstract: Skin lesions in reptiles are caused by different factors and result in scarring and dysecdysis, predisposing to dermatitis. It may let the animal out of exhibitions or lead to the abandonment when created as pets. In order to compare different topical products and procedures for skin repair, twelve adult male tegu lizards (Tupinambis merianae) were used. Four circular square arranged skin lesions were created at the dorsum of each animal. Animals were divided in three groups (four animals/group) and each group was treated with two medications. In Group 1, animals were treated with saline and policresulen, in Group 2, with tincture of iodine and untreated (control group), and in Group 3, chlorhexidine and autologous skin graft. The lesions on the right side were macroscopicaly evaluated daily and the planimetry was realized every seven days. The left lesions were sampled every seven days for histopathological examination. The lizards showed healing process similar to mammals; however, it was notably slower. In this experiment, the skin graft showed no advantage, undergoing necrosis in all four animals. Policresulen closed the wounds in four days, while other groups had the same effect between 8 and 12 days. The lesions were completely epithelialized between 40 and 53 days, with no difference between groups regarding the total time of healing, tissue retraction and appearance of the scar / Mestre

Animals trade-off limited resources among competing demands. Trade-offs are difficult to quantify because it is challenging to measure investment into disparate physiological systems using a common scale. Additionally, biologists desire methods to more precisely measure energy status in wild animals. I used stable isotopes to help solve both of these problems. I examined natural spatial and temporal variation in stable isotope signatures of wild lizards and found significant variation. In the lab, I was able to demonstrate the utility of nitrogen stable isotope ratios of uric acid pellets for measuring nutritional stress. By tracing labeled amino acids through the bodies of gravid female lizards, I demonstrated that vitellogenesis and wound healing compete for amino acids and quantified the direction and magnitude of the trade-offs. I showed that reproductive-immune trade-offs vary based on reproductive stage and energy availability, have effects on metabolism and immune function, and are influenced by hormonal mechanisms. My findings shed light on the interconnectedness of stable isotope endpoints and key physiological systems in animals. I showed that isotopic signatures of physiological stress can be reflected at a large scale in natural populations, and I made novel measurements of the size and direction of trade-offs, which were formerly limited to physiological and performance outcomes.

This study focused on the interactions between small skinks and their major predators/competitors (birds) and prey (arthropods) in core and edge areas of small, long established remnant Cumberland Plain woodlands of Richmond, New South Wales, Australia. Eight study sites were selected (4 edge and 4 core) within the peri-urban environment to compare the abundance and diversity of small skinks, birds and arthropods. Of the three taxa, only arthropods maintained a similar abundance and diversity between edge and core areas. Birds and skinks were either in lower numbers in the perimeter areas (skinks), or were interior/perimeter specialists (birds). Arthropod diversity and abundance of preferred skink prey, was also examined. It was concluded that the distribution of arthropod prey was similar between core and edge areas, and therefore, was not considered to have an influence on small skinks’ ability to inhabit edges. However, large, aggressive/noisy birds (including skink and small bird predators) dominated edge areas. Those birds encountered in the edge foraged in a variety of niches, such as on the ground or they swooped from vantage points. This, coupled with other anti competitor behaviour (e.g. noise, aggression, flocking), placed these birds in proximity to skinks in the edge areas. However, as most of these birds were also predators of skinks, it was concluded that the evidence supported a predator/prey relationship in the perimeter area, rather than a competitive one. This was supported by predation rates on skinks, using decoys. Birds preyed on small skink in greater numbers in the perimeter of woodland remnants than in their interior. This predation pressure was sustained throughout the year. It was concluded that predation and/or displacement of skinks and small birds resulted in lower numbers being observed in edge compared to core areas. This study demonstrated that old, small remnants not continually exposed to major disturbances (e.g. logging, agricultural practices) can re-establish stability in terms of environmental conditions. However, a stable environment or a single taxonomic group does not necessarily predict the response individual taxa will have to edge habitats and that this can alter the interactions between dependent groups such as prey/predator or competitors. . / Doctor of Philosophy (PhD)

Approximately two thirds of the world’s land is directly supporting human population contributing to an accumulation of disturbed habitats. This dissertation investigates the impact of human mediated habitat disturbance, in introduced Eucalyptus plantations, on community diversity and population divergence using Brazilian Cerrado lizards as a model. Data was collected along a gradient from undisturbed cerrado to disturbed Eucalyptus plantations. Community diversity differences and indicator species were identified. Furthermore, the role of phenotypic divergences were determined based on populations able to persist in disturbed habitats. Dispersal, food availability (body condition), competition and predation (caudal autotomy) were tested as potential mechanisms driving phenotypic divergences. Additionally, I investigated phylogenetic community structure differences between habitats to test for a phylogenetic signal to disturbance. The evidence showed community diversity indices were significantly lower in Eucalyptus plantations with a decrease along the cerrado– Eucalyptus gradient. Furthermore, 29 % of the Cerrado species suffered local extinctions in the disturbed habitat and of these 80 % are endemic species. One indicator species was identified for the disturbed habitat and seven species were identified for the undisturbed habitat. Species able to persist in both habitats demonstrated morphological trait divergences. These species showed short dispersal distances with only two individuals dispersing between habitats indicating a mechanism driving the observed phenotypic divergences. Another mechanism is body condition, which was higher in the disturbed habitats, reflecting increased food availability possibly due to the decreased abundances. Caudal autotomy showed no difference between the habitats indicating that competition and predation are not driving phenotypic divergences. Phylogenetic community structure demonstrated a phylogenetic signal to disturbance. The undisturbed habitat consists of communities with more closely related species compared to the disturbed habitat: indicating evolutionary forces such as habitat filtering as the stronger process structuring these communities. Whereas, disturbed communities are structured by ecological forces such as competition. This research provides information for the preservation and maintenance of the Cerrado biodiversity and has an even broader impact since habitat change caused by human activities touches a plethora of ecosystems. / text

Bill Brooks is worried about the future of his organization, perhaps even more than he is about the future of horned lizards. Brooks is the president and a founding member of the Horned Lizard Conservation Society, a nonprofit with the goal of protecting all species of horned lizard. The HLCS was founded in Austin in 1991, after its members realized that populations of Texas horned lizards, Phrynosoma cornutum, had crashed at some point in the preceding decades; they wanted to both figure out what had happened to the lizards and how to bring them back. Cornutum is an iconic species and the state reptile, and almost all Texans over the age of 40 can remember seeing them in the hundreds as children. Flash-forward a quarter of a century, and a lot of these founding members are either old and grey or deceased. HLCS membership is down, and Brooks thinks that if there aren’t any of his kind left (Texans that grew up with “horny toads”), there’ll be no one left to care. Herpetologists are working on returning healthy horned lizard populations to the wild, and both reintroduction and relocation programs are showing signs of success. But can they work, and will it be too late for Brooks’ organization and a Texas identity dependent upon horned lizards? / text