Abstract There are a remarkable variety of sex determination systems among different animal taxa. In most animals, sex is determined chromosomally. Although in an increasing number of animals sex determination has been found to be influenced primarily by the environment. Species with genotypic sex determination (GSD) have their sex determined at the time of fertilization, by genetic factors alone and those with environmental sex determination (ESD) have their sex determined by environmental factors that act after fertilization. Temperature-dependent Sex Determination (TSD), whereby the sex of the developing embryos depends on the temperature at which they develop is widespread in oviparous reptiles and occurs in all crocodilians, marine turtles and tuatara examined to date and is common in many freshwater turtles and lizards. SECTION ONE Temperature-dependent sex determination (TSD) was never expected to occur in viviparous reptiles, as thermoregulation by pregnant females would result in relatively stable gestation temperatures. Temperature-dependent sex determination and viviparity goes against all the basic assumptions that TSD occurs in oviparous reptiles where temperatures within a nest vary widely. However, skewed sex ratios as a result of incubation temperature indicated the possibility of TSD in the viviparous lizard Eulamprus tympanum. In my first experiments I show the first recorded case of a viviparous reptile with TSD. The developing embryos of the viviparous skink E. tympanum are subject to TSD, with gestation temperature having a highly significant effect on sex and warmer temperatures giving rise to male offspring (Chapter 1). Sex is fully determined at the time of birth and can be differentiated histologically into testes or ovaries (Chapter 2). The morphology and histological characteristics of the gonads of neonatal E. tympanum resulting from the treatment temperatures described in chapter 1 illustrate that sex in E. tympanum is easily distinguished at the time of birth and corresponds with the presence or absence of hemipenes. Males are histologically characterised by an elongated gonad consisting of seminiferous tubules with either no cortical epithelium or, if present at all, in a very thin band. If they are present, M�llerian ducts, showing signs of degeneration, are attached to the kidney by a shortened mesosalpinx. Females are histologically characterised by an irregularly shaped gonad consisting of a thick cortical epithelium that occasionally contains oocytes. The M�llerian ducts are obvious structures attached to the kidney by a fibrous mesosalpinx. The presence or absence of hemipenes is a reliable technique for determining sex in newborn E. tympanum. Sex determination is easiest to perform on neonates within the first few days of birth as hemipenes become increasingly difficult to evert as neonates age, however, with practice they are easily identified without full eversion. SECTION TWO The thermal biology of E. tympanum in the field is restricted by both the thermal properties of their habitat (Chapter 3) and behavioural modifications when faced with a predation threat (Chapter 4). The available temperatures in the field suggest that TSD is biologically relevant in the species and not just a laboratory artefact; E. tympanum can attain mean selected temperatures achieved in the laboratory but the proportion of time at the temperature is restricted. Females actively thermoregulate in the field, although they are restricted in their efficiency of thermoregulation by environmental constraints, for example, microhabitat structure, weather conditions, predator avoidance and social ranking. The highly territorial nature and high densities of E. tympanum present in Kanangra Boyd National Park potentially force less dominant individuals into less favourable habitats that are significantly cooler. An important point is that gravid females in more favourable habitats in the period encompassing the middle third of development (the assumed sex determining period) are selecting higher temperatures, with lower variance and have greater thermoregulatory efficiency than during the rest of pregnancy, therefore, thermoregulating more precisely during this thermosensitive period (Chapter 3). Chemosensory cues provide important information on the risk of predation. Hence, chemoreception is a common mechanism used by many species to detect the presence of, and subsequently respond to, a potential predator. The perceived risk of predation may force retreat to sub-optimal conditions, forcing a trade-off between the risk of predation and the ability to acquire resources. The basking regime maintained by gravid female E. tympanum, can directly alter sex ratios of offspring produced through temperature-dependent sex determination (Chapter 1). The avoidance of predator scents may restrict basking ability and in turn alter the sex of offspring produced. I measured responsiveness to chemical cues using tongue flicks as an indicator of chemical discrimination in females of different reproductive condition. I then measured activity and basking behaviour of gravid and non-gravid females in experimental enclosures in the presence of various chemical stimuli to determine if basking opportunity is compromised by the presence of a predator scent. Females respond differently depending upon reproductive condition, with gravid females responding most significantly to a predator scent. Activity, basking frequency, and time spent in the open (basking duration) are significantly reduced in gravid females in the presence of a predator stimulus. Under laboratory conditions, gravid females modify their behaviour and forego the opportunity to bask when there is a perceived predation risk (Chapter 4). SECTION THREE As female viviparous reptiles can regulate the temperature of the embryo by maternal temperature selection (Chapter 1), the occurrence of TSD in E. tympanum opens the possibility for females to select the sex of offspring. Reproducing females may benefit by facultatively adjusting their investment into sons over daughters or vice versa, in response to population wide shifts in adult sex ratios. Female E. tympanum, can manipulate the sex of their offspring in response to sex imbalances in the population using temperature-dependent sex determination (Chapter 5). When adult males are scarce, females produce male-biased litters and when adult males are common, females produce female-biased litters. The cues used by a female to assess the adult population are not known, but presumably depends upon the female�s experience throughout the breeding season and is the subject of further investigation (Chapter 6). The maternal manipulation of offspring sex ratio in E. tympanum suggests a selective advantage of temperature-dependent sex determination. Any facultative sex ratio response needs to recognise the scarcity of one sex in order to overproduce that sex in the next generation; offspring sex ratio will vary inversely with adult sex ratio. Maternal sex allocation in E. tympanum is linked with population (or adult) sex ratio (Chapter 5), and one of the mechanisms by which females recognise an imbalance may be linked to visual recognition of males (Chapter 6). Females maintained throughout pregnancy without any male stimulus produce entirely male offspring (Chapter 5). In contrast females exposed to male stimulus produce both sexes (Chapter 5). Females respond differently to varying degrees of male stimulus and visual recognition of males in a population may be more important than chemoreception. In the absence of visual cues, females produce more male offspring, even when chemosensory cues are present (Chapter 6). The study system presented here offers many advantages over oviparous species with TSD, due to E. tympanum being relatively short lived and fast maturing. Thus, the fitness consequences over multiple generations as a result of gestation can be investigated. Viviparity allows maternal control of embryonic temperature during gestation and a means of maternal sex allocation. Until now the maternal side of TSD and sex allocation has been where the mother deposits her eggs and the allocation of sex steroid hormones at oviposition, both of which have been difficult to study. The work presented and the study system itself should inspire great interest in TSD and viviparous reptiles.

Abstract There are a remarkable variety of sex determination systems among different animal taxa. In most animals, sex is determined chromosomally. Although in an increasing number of animals sex determination has been found to be influenced primarily by the environment. Species with genotypic sex determination (GSD) have their sex determined at the time of fertilization, by genetic factors alone and those with environmental sex determination (ESD) have their sex determined by environmental factors that act after fertilization. Temperature-dependent Sex Determination (TSD), whereby the sex of the developing embryos depends on the temperature at which they develop is widespread in oviparous reptiles and occurs in all crocodilians, marine turtles and tuatara examined to date and is common in many freshwater turtles and lizards. SECTION ONE Temperature-dependent sex determination (TSD) was never expected to occur in viviparous reptiles, as thermoregulation by pregnant females would result in relatively stable gestation temperatures. Temperature-dependent sex determination and viviparity goes against all the basic assumptions that TSD occurs in oviparous reptiles where temperatures within a nest vary widely. However, skewed sex ratios as a result of incubation temperature indicated the possibility of TSD in the viviparous lizard Eulamprus tympanum. In my first experiments I show the first recorded case of a viviparous reptile with TSD. The developing embryos of the viviparous skink E. tympanum are subject to TSD, with gestation temperature having a highly significant effect on sex and warmer temperatures giving rise to male offspring (Chapter 1). Sex is fully determined at the time of birth and can be differentiated histologically into testes or ovaries (Chapter 2). The morphology and histological characteristics of the gonads of neonatal E. tympanum resulting from the treatment temperatures described in chapter 1 illustrate that sex in E. tympanum is easily distinguished at the time of birth and corresponds with the presence or absence of hemipenes. Males are histologically characterised by an elongated gonad consisting of seminiferous tubules with either no cortical epithelium or, if present at all, in a very thin band. If they are present, M�llerian ducts, showing signs of degeneration, are attached to the kidney by a shortened mesosalpinx. Females are histologically characterised by an irregularly shaped gonad consisting of a thick cortical epithelium that occasionally contains oocytes. The M�llerian ducts are obvious structures attached to the kidney by a fibrous mesosalpinx. The presence or absence of hemipenes is a reliable technique for determining sex in newborn E. tympanum. Sex determination is easiest to perform on neonates within the first few days of birth as hemipenes become increasingly difficult to evert as neonates age, however, with practice they are easily identified without full eversion. SECTION TWO The thermal biology of E. tympanum in the field is restricted by both the thermal properties of their habitat (Chapter 3) and behavioural modifications when faced with a predation threat (Chapter 4). The available temperatures in the field suggest that TSD is biologically relevant in the species and not just a laboratory artefact; E. tympanum can attain mean selected temperatures achieved in the laboratory but the proportion of time at the temperature is restricted. Females actively thermoregulate in the field, although they are restricted in their efficiency of thermoregulation by environmental constraints, for example, microhabitat structure, weather conditions, predator avoidance and social ranking. The highly territorial nature and high densities of E. tympanum present in Kanangra Boyd National Park potentially force less dominant individuals into less favourable habitats that are significantly cooler. An important point is that gravid females in more favourable habitats in the period encompassing the middle third of development (the assumed sex determining period) are selecting higher temperatures, with lower variance and have greater thermoregulatory efficiency than during the rest of pregnancy, therefore, thermoregulating more precisely during this thermosensitive period (Chapter 3). Chemosensory cues provide important information on the risk of predation. Hence, chemoreception is a common mechanism used by many species to detect the presence of, and subsequently respond to, a potential predator. The perceived risk of predation may force retreat to sub-optimal conditions, forcing a trade-off between the risk of predation and the ability to acquire resources. The basking regime maintained by gravid female E. tympanum, can directly alter sex ratios of offspring produced through temperature-dependent sex determination (Chapter 1). The avoidance of predator scents may restrict basking ability and in turn alter the sex of offspring produced. I measured responsiveness to chemical cues using tongue flicks as an indicator of chemical discrimination in females of different reproductive condition. I then measured activity and basking behaviour of gravid and non-gravid females in experimental enclosures in the presence of various chemical stimuli to determine if basking opportunity is compromised by the presence of a predator scent. Females respond differently depending upon reproductive condition, with gravid females responding most significantly to a predator scent. Activity, basking frequency, and time spent in the open (basking duration) are significantly reduced in gravid females in the presence of a predator stimulus. Under laboratory conditions, gravid females modify their behaviour and forego the opportunity to bask when there is a perceived predation risk (Chapter 4). SECTION THREE As female viviparous reptiles can regulate the temperature of the embryo by maternal temperature selection (Chapter 1), the occurrence of TSD in E. tympanum opens the possibility for females to select the sex of offspring. Reproducing females may benefit by facultatively adjusting their investment into sons over daughters or vice versa, in response to population wide shifts in adult sex ratios. Female E. tympanum, can manipulate the sex of their offspring in response to sex imbalances in the population using temperature-dependent sex determination (Chapter 5). When adult males are scarce, females produce male-biased litters and when adult males are common, females produce female-biased litters. The cues used by a female to assess the adult population are not known, but presumably depends upon the female�s experience throughout the breeding season and is the subject of further investigation (Chapter 6). The maternal manipulation of offspring sex ratio in E. tympanum suggests a selective advantage of temperature-dependent sex determination. Any facultative sex ratio response needs to recognise the scarcity of one sex in order to overproduce that sex in the next generation; offspring sex ratio will vary inversely with adult sex ratio. Maternal sex allocation in E. tympanum is linked with population (or adult) sex ratio (Chapter 5), and one of the mechanisms by which females recognise an imbalance may be linked to visual recognition of males (Chapter 6). Females maintained throughout pregnancy without any male stimulus produce entirely male offspring (Chapter 5). In contrast females exposed to male stimulus produce both sexes (Chapter 5). Females respond differently to varying degrees of male stimulus and visual recognition of males in a population may be more important than chemoreception. In the absence of visual cues, females produce more male offspring, even when chemosensory cues are present (Chapter 6). The study system presented here offers many advantages over oviparous species with TSD, due to E. tympanum being relatively short lived and fast maturing. Thus, the fitness consequences over multiple generations as a result of gestation can be investigated. Viviparity allows maternal control of embryonic temperature during gestation and a means of maternal sex allocation. Until now the maternal side of TSD and sex allocation has been where the mother deposits her eggs and the allocation of sex steroid hormones at oviposition, both of which have been difficult to study. The work presented and the study system itself should inspire great interest in TSD and viviparous reptiles.

Among vertebrates, oviparous animals lay eggs with a calcified eggshell and eggs are laid in an external environment, while viviparous animals retain eggs in oviduct until they give birth to live young. Viviparity has evolved in the lineage of snakes and lizards (squamates) over 100 times, more than all other vertebrate groups. Embryos of oviparous squamates obtain calcium from both yolk and eggshell while their viviparous counterparts lack a calcified eggshell and must obtain their calcium solely from yolk, or from yolk plus placental transfer. During embryonic development, squamates rely on calcium to build the skeleton before hatching. The extent of skeletal ossification at hatching or birth varies considerably among vertebrates. This study aims to determine if skeletal development/ossification varies in association with reduced eggshell calcium in embryos of viviparous squamate species. We studied the amount of ossification and overall size of embryos and hatchlings from an oviparous and a viviparous population of the reproductively bimodal lizard Zootoca vivipara. Previous studies suggested that limb development is delayed, and that overall size is reduced in viviparous squamates. We tested the hypothesis that viviparous embryos and hatchlings are more skeletally immature and smaller in size than oviparous embryos and hatchlings in squamates. To achieve this, specimens from both populations, spanning multiple stages of embryonic development including hatchlings, were cleared & stained to reveal skeletal cartilage and bone. Lengths of total body, humerus, femur, skull and Meckel’s cartilage were measured from photographs of cleared & stained specimens taken with a Cannon EOS 70D camera on a Motic, Leica MZ9 Compound Microscope, with a measurement reference in each picture. Photos were calibrated to the measurement reference and total length measurements were obtained using iSolutionLite® software. In addition to total measurements, the lengths of ossified portions on the humerus and femur were also measured. Preliminary results revealed that total length of the skull and body are reduced in embryos and hatchlings of viviparous specimens. Total length of the limbs and of Meckel’s cartilage are not significantly different between populations. However, it appears that the amount of ossification in the limbs is reduced in oviparous specimens. These preliminary findings do not support our hypothesis and indicate that reduction in eggshell calcium in embryos of viviparous populations does not negatively impact ossification during development but does influence overall size.

L'anthropisation croissante et les changements globaux entrainent de nombreuses perturbations dans l'environnement conduisant à la dégradation, et même à la destruction d'habitat. Le modèle allostatique proposé par McEwen et Wingfield en 2003 permet de mieux appréhender les relations entre physiologie et réponses démographique dans le cadre de la physiologie de la conservation. A travers des approches expérimentales en milieux controlé et semi-naturel nous avons testé l'effet de d'un stress chronique sur les changements des traits d'histoires de vie et les paramétres physiologique chez le lézard vivipare (Zootoca vivipara). Nous discutons de la caractérisation de la charge allostatique et des effets à court terme et long terme d'un stress chronique et l'utilisation de la charge allostatique comme indicateur de la dégradation de l'habitat. Ce travail pourra être utilisé afin de mieux comprendre et prédire la dynamique des populations naturelles soumises aux changements globaux. / Increasing anthropogenic and global changes are causing many disturbances in the environment leading to degradation and even destruction of habitat. The allostatic model proposed by McEwen and Wingfield in 2003 provides a better understanding of the relationships between physiology and demographic responses within the framework of conservation physiology. Experimental approaches in controlled and semi-natural environments have tested the effect of chronic stress on changes in life histories and physiological parameters in the viviparous lizard (Zootoca vivipara). We discuss the characterization of allostatic charge and the short-term and long-term effects of chronic stress and the use of allostatic charge as an indicator of habitat degradation. This work can be used to better understand and predict the dynamics of natural populations subjected to global changes.

Submitted by (edna.saturno@ufrpe.br) on 2017-02-17T13:31:07Z No. of bitstreams: 1 Mirna Regina dos Santos Tambourgi.pdf: 973009 bytes, checksum: e1b23c1d9979ea2656cba5d134038e44 (MD5) / Made available in DSpace on 2017-02-17T13:31:07Z (GMT). No. of bitstreams: 1 Mirna Regina dos Santos Tambourgi.pdf: 973009 bytes, checksum: e1b23c1d9979ea2656cba5d134038e44 (MD5) Previous issue date: 2010-02-17 / In the period from December 2003 to December 2009, the reproductive tract were collected from 201 individuals of Carcharhinus longimanus, caught by of commercial tuna boats in the fleet, who were pelagic longline fisheries and hand line in the area located between latitudes 06 º 45'N and 23 ° 36'S and longitudes 018 ° 44'W and 053 º 13'W. Among the 201 individuals captured, 98 were females (74.5 – 227 cm TL) and 103 were males (72 – 242 cm TL) with a sexual proportion males to females of 1:1. Females were categorized as immature (N = 78; 74.5 – 170 cm TL), matures (N = 13; 165 – 223 cm TL), pré-ovulatory (N = 2; 181 – 187 cm CT) and pregnant (N = 5; 169 – 227 cm TL). The uterine fecundity ranged from 1 to 10 embryos. The cycle of pregnancy is annual with birth occurring at the end / beginning of the year. The size at first maturity of females was estimated at 170 cm TL. Of the 103 males captured, 75 were immature (72 to 160 cm TL), 8 maturing (170 - 196) and 20 were mature (160 - 242 cm TL). The size of sexual maturity was between 160 and 170 cm. / No período de dezembro de 2003 a dezembro de 2009, foram coletados os aparelhos reprodutores de 201 espécimes de Carcharhinus longimanus, capturados por meio de barcos da frota comercial atuneira, que realizaram pescarias com espinhel pelágico e linha de mão, na área localizada entre as latitudes 06º45’N e 23º36’S e as longitudes de 018º44’W e 053º13’W. Deste total, 98 eram fêmeas (74,5 a 170 cm CT), maduras (N= 13; 165 a 223 cm CT), pré-ovulatórias (N= 2; 181 e 187 cm CT) e prenhes (N= 5; 169 a 227 cm CT). A fecundidade ovariana variou de 1 a 10 folículos vitelogênicos e a uterina, de 1 a 10 embriões. O ciclo de gestação é anual com nascimento ocorrendo ao final/início do ano. O tamanho de primeira maturação das fêmeas desta espécie é de 170 cm CT. Dos 103 machos capturados, 75 eram imaturos (72 – 160 cm CT), 8 em maturação (170 - 196) e 20 eram maduros (160 – 242 cm CT). O tamanho de primeira maturação sexual entre 160 e 170 cm.

Submitted by Cláudia Bueno (claudiamoura18@gmail.com) on 2016-02-11T12:23:04Z No. of bitstreams: 2 Dissertação - Michelle Furquim Leão - 2007.pdf: 19618077 bytes, checksum: 185360e935aab77ea3553e7a7d1c4093 (MD5) license_rdf: 23148 bytes, checksum: 9da0b6dfac957114c6a7714714b86306 (MD5) / Approved for entry into archive by Luciana Ferreira (lucgeral@gmail.com) on 2016-02-12T10:57:42Z (GMT) No. of bitstreams: 2 Dissertação - Michelle Furquim Leão - 2007.pdf: 19618077 bytes, checksum: 185360e935aab77ea3553e7a7d1c4093 (MD5) license_rdf: 23148 bytes, checksum: 9da0b6dfac957114c6a7714714b86306 (MD5) / Made available in DSpace on 2016-02-12T10:57:42Z (GMT). No. of bitstreams: 2 Dissertação - Michelle Furquim Leão - 2007.pdf: 19618077 bytes, checksum: 185360e935aab77ea3553e7a7d1c4093 (MD5) license_rdf: 23148 bytes, checksum: 9da0b6dfac957114c6a7714714b86306 (MD5) Previous issue date: 2007-08-20 / Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - CAPES / The sharp toxicity of the herbicide original Roundup (Glifosato), one of the more acquaintances and maybe one of the more used dessecantes in direct planting in the farmings in the area Centro Oeste and in every country now, it was investigated through the effects detected in the fish Poecilia viviparous... / A toxicidade aguda do herbicida Roundup original (Glifosato), um dos mais conhecidos e talvez um dos mais utilizados dessecantes em plantio direto nas lavouras na região Centro Oeste e em todo país atualmente, foi investigada através dos efeitos detectados no peixe Poecilia vivípara...

Förlust av biologisk mångfald är den planetära gräns som överskrids mest, och medför bland de största hoten mot biosfärens integritet. Reptiler är funktionella delar i ekosystem, men habitatförlust, förändrad markanvändning och klimatförändringar orsakar globala populationsminskningar. Ekologisk övervakning möjliggör förståelse för arters tillstånd, och utveckling av bevarandeåtgärder. Denna förekomststudie syftar till att undersöka lokal reptilfauna, som inventerades med artificial cover objects (ACO) i två naturområden på Billingen, ett av de västsvenska platåbergen, i Skövde kommun 2021. Data från miljöbeskrivning och inventering samt över väderförhållanden samlades in, och samband mellan observationer och miljöfaktorer testades med statistiskt t-test för positiva skillnader mellan datagrupper. Skogsödla (Zootoca vivipara), kopparödla (Anguis fragilis), snok (Natrix natrix) och huggorm (Vipera berus),alla allmänna reptilarter i regionen, observerades i undersökningsområdet. Nio observationer gjordes under 20 regnfria dagar i juni och juli, med temperaturer över 18°C. Biotopkvaliteterna brynmiljöer och nektarresurser, solexponering och de öppna biotoperna beteshage och elledningsgata, förekom i signifikant högre utsträckning där observationer gjordes, än på platser utan observationer. Markfuktighet, mänsklig närvaro, naturvärden och områdesskydd uppvisade ingen signifikans. Resultaten överensstämmer med andra studier om reptilarternas habitat, och kan ses gälla på lokal nivå trots ett relativt litet dataunderlag. Effektiva reptilbevarande åtgärder i närområdet kan innebära upprätthållande och skapande av ytor med hög solinstrålning och strukturell komplexitet, fortsatt men inte utökat lågintensivt naturbete, samt ökad halt avträdslagsblandning och brynmiljöer i produktionsskog. Kunskapen kring reptilers ekologi, samtövervakningen av populationer, behöver utökas. Det rekommenderas vidare studier om reptiler, deras habitat och bevarande av biologisk mångfald på lokala nivåer. / Biodiversity loss is the most exceeded planetary boundary, and entails some of the largest threats against biosphere integrity. Reptiles are functional parts of ecosystems, but habitat loss and changes in land use and climate are causing global population declines. Ecological monitoring enables understanding of species’ conditions, and development of conservation measures. This occurrence study aims to investigate local reptile fauna, surveyed using artificial cover objects (ACOs), in two areas on Billingen, one of Western Sweden’s table mountains, in Skövde municipality 2021. Environmental, survey and weather data were collected, and relations between observations and environmental factors were tested with statistical t-test for positive differences between data groups. Zootoca vivipara, Anguis fragilis, Natrix natrix and Vipera berus, all regionally common reptile species, were observed in the investigated area. Nine observationswere made during 20 rain-free days in June and July, with temperatures above 18°C. Solarexposure, biotope qualities Forest edges and Nectar resources, and the open biotopes Pasture and Powerline corridor, occurred in significantly higher extent where observations were made, than on sites without observations. Soil moisture, Human presence, Natural values and Environmental protection did not exhibit any significance. The results correspond to other studies on the reptilespecies’ habitat, and may apply on a local scale despite a relatively small data amount. Effective conservation measures for local reptiles can imply maintaining and creating spaces with high solar radiation and structural complexity, continued but not extended low-intensity grazing, and increasing tree species mixture and edge zones in production forests. Knowledge on reptile ecology and population monitoring needs to increase. Further studies on reptiles and biodiversity conservation on local scales are recommended.