Dissertations / Theses on the topic 'Freshwater turtles'

Thesis (Ph. D.)--School of Biological Sciences, Faculty of Science, University of Sydney, 2005.
Title from title screen (viewed 13 January 2009) Submitted in fulfilment of the requirements for the degree of Doctor of Philosophy to the School of Biological Sciences, Faculty of Science. Includes bibliographical references. Also issued in print.

An unusual turtle fauna from Lightning Ridge, New South Wales (Albian) reveals that Australian turtles had a more extensive Mesozoic history than previously indicated. Reevaluation of several primitive groups provides novel information on turtle evolution in the southern hemisphere. Seven turtle taxa are identified at Lightning Ridge. Two are Testudines indet. and two indeterminate chelid groups are evinced by isolated elements. Three new taxa are assigned to the new family Spoochelyidae in the superfamily Meiolanoidea. Spoochelys ormondea n. fam., gen. et sp., Sunflashemys bartondracketti n. gen. et sp. and Opalania baagiwayamba n. gen. et sp. are predominantly land-living turtles with high-domed shells and short manus and pes. The sister-group relationship with the Meiolaniidae, supported by a suite of cranial and postcranial synapomorphies, increases the stratigraphic range of the horned turtles by around ~ 50 my. Primitive structures in Spoochelys (postparietal, supratemporal and interpterygoid vacuity), occur with derived features that are variably developed across Triassic and Jurassic turtles. Phylogenetic analysis precariously resolves the meiolanoids as sister group to a clade containing Palaeochersis and Proterochersis. Limited pleurodiran attributes suggest that meiolanoids may be pleurodiromorphs, closer to primitive pleurodires than to cryptodires. As basal ‘side-necked’ turtles, the Lightning Ridge meiolanoids permit first insights into cranial and postcranial progressions in pleurodiran stem taxa. Evidence of diverse meiolanoids in Early Cretaceous Australia and ancient radiations of meiolanoid-like turtles in southern Pangea, suggest that the horned turtles are a Triassic group and that the dichotomy between Pleurodira and Cryptodira occurred well before the Late Triassic. Early Cretaceous chelids at Lightning Ridge occur at higher palaeolatitude than in South America. The temporal range of Australian chelids is extended by ~ 50 my, demonstrating that chelids had a Jurassic history in Australia, with broad diversifications across the polar supercontinent. The palaeoecological setting of Lightning Ridge is comprehensively described for the first time. Diverse invertebrates and vertebrates include terrestrial, freshwater aquatic and rare marine forms that are anomalous at this near-polar palaeolatitude (~65-70oS). The anachronistic occurrence in Early Cretaceous Australia of distinctive radiations of ‘Triassic-type’ turtles, and other relic groups, implies prolonged intervals of biogeographic isolation in the eastern provinces of Pangea.

Doctor of Philosophy (PhD)
Over 85% of Australia’s population live in urban areas and many turtle populations occur on Australia’s east coast where urban development is particularly concentrated. In the state of NSW, over half of the freshwater coastal wetlands have been highly modified or completely destroyed, and urban freshwater creeks often have only a narrow strip of weedy bushland left along their banks. Even though habitat degradation may result in declines in density and distribution of turtle populations, there are few data on Australian freshwater turtles in urban areas. In addition to extreme habitat alteration, urban waterways are innundated with anthropogenic contaminants from sources including wet weather surface runoff and industrial and sewage discharges. Pollutants can impact all systems of the body with potentially severe effects on reproduction and survival that can result in deterioration of animal populations. Turtles are particularly susceptible to anthropogenic contaminants due to their intimate contact with the aquatic environment, an often high trophic level, their ability to accumulate toxins, and their longevity. For almost all contaminants, the degree of accumulation in and effect on reptile species is unknown. Sublethal effects in field situations are particularly poorly studied and have never been documented in pleurodiran turtles. As a pioneering work in Australian reptile ecotoxicology, this thesis takes a broad approach, but focuses primarily on immunotoxicity and reproductive toxicity – two areas that greatly impact the size and continuance of animal populations. The aim of the thesis is to provide baseline data on haematology, cellular immunology and tissue metal concentrations for freshwater turtles in Sydney – data which were lacking for all Australian turtle species prior to this study. After initial assessment of the distribution and density of freshwater turtles in Sydney, the study examines the potential for Sydney’s turtles as sentinel species for measuring the effects of pollution on haematology, cellular immunity, and parasite loads; and considers the relationships between urban metal pollution and reproductive variables. The relative suitability of non-lethally sampled tissues (blood, carapace, egg) for use in biomonitoring is also assessed. Three species of Australian freshwater turtles were found in the Sydney region, with Chelodina longicollis occurring naturally in the area, and populations of Emydura macquarii and Elseya latisternum likely to have originated from translocated individuals. The North American turtle Trachemys scripta elegans was not encountered during this study despite concerns that it was establishing in the Sydney area. Chelodina longicollis populations were widespread, although poor recruitmment was indicated by low capture rates and comparatively low percentage of juveniles at some sites. Not so widespread, Emydura macquarii was present in much larger numbers than C. longicollis and with a high juvenile component in some areas of southeastern Sydney. I provide information on erythrocyte and leucocyte parameters in C. longicollis over a range of sites, pollution conditions, and seasons. In C. longicollis, numbers of lymphocytes, heterophils and eosinophils varied over sites, but not due to pollution from sewage treatment plant outfalls. There was significant temporal variation in erythrocyte, lymphocyte, eosinophil, heterophil, and basophil number, the heterophil:lymphocyte ratio, and haematocrit, but not consistently among sites. Future studies should ensure simultaneous sampling across sites for comparative purposes. Similarly, turtle populations downstream of sewage treatment plant outfalls showed no consistent difference in number, body condition, blood haemogregarine load, or leech (haemogregarine vector) load from upstream populations. Leech (Helobdella papillornata, with some Placobdella sp.) load and haemogregarine numbers increase dramatically once C. longicollis reach a carapace of 110 mm. The number of leeches on turtles varied across season, year, and site. Turtles with large numbers of leeches had reduced haematocrit, but the presence of leeches had no other correlations with haematological parameters. Haemogregarine numbers did not change across season or year, and were not correlated with haematological variables. The hypothesis that pollutants lead to an increase in normal blood protozoa due to reduced immunity thus was not supported. The concentration of metals in C. longicollis and E. macquarii carapace and in lagoon sediments varied significantly over four urban and four national park sites, but not based on this split. Pollution in periurban areas, such as illegal dumping of toxic wastes and atmospheric deposition of pollutants, means that each site must be classified separately as to degree of metal pollution. There was little or no affect of species, size, sex, or gravidity on metal concentrations in the carapace of adult turtles. Emydura macquarii had higher concentrations of blood Fe than C. longicollis from a different site, but this is possibly due to an increase in haemoglobin resulting from the site’s low aquatic oxygen concentration rather than any increased environmental exposure. Chelid turtles in Sydney do not show much promise as a biomonitoring tool. Carapace analysis is largely discounted as a potential tool for metal biomonitoring due to poor correlations between potentially toxic metals in non-lethally samplable tissues (carapace, claw) and internal organs (liver, kidney) or bone (femur). However, carapace metal concentrations still potentially reflect long-term metal presence or different dietary exposures as evidenced by the significant variation in concentrations over sites. A rare correlation was found for concentrations of aquatic Pb and carapace Pb, and a correlation was also found for concentrations of blood Pb and carapace Pb in E. macquarii. Thus any potential for tissue biomonitoring seems to lie with this highly ecotoxicologically relevant metal. Although two other ecotoxicologically relevent metals, Cu and Se, were significantly higher in egg contents of C. longicollis compared to E. macquarii, these elements are also essential and a lack of baseline values means it is not known if this simply reflects natural taxonomic variation. Ni, a metal of toxicological concern in sea turtles, was not present in egg contents, and only variably present in eggshell. The absence of Pb from eggs, despite its presence in many maternal tissues, suggests that selective metal uptake into eggs may be protective of toxic elements, rather than eggs serving as a maternal method of toxic metal elimination as has been previously suggested. The paucity of toxic metal detection in eggs renders them unlikely tissues for biomonitoring. The maternal tissue or tissues or environmental source from which egg metals originate remains obscure, although a significant negative effect of maternal carapace concentrations of Ca and Mg on eggshell thickness in E. macquarii indicates that there may be mobilisation of Ca and Mg from the carapace for eggshell formation. The only metal whose eggshell concentration correlated with eggshell thickness was Mg, indicating that ecotoxic metals previously associated with eggshell thinning are not problematic in the Sydney chelids. As with North American turtles living at polluted sites, none of the chelid hatchlings were found to have any overt abnormalities. Hatching success was poor and hatching mass low for eggs of both C. longicollis and E. macquarii, although results from natural nests are required to determine whether or not this was an outcome of hormonally-induced oviposition and artificial incubation. It is difficult to interpret metal concentrations found in the soft tissues, calcified tissues, and eggs of chelonians due to the paucity of comparative data, and much more research is required on tissue metal concentrations before patterns will emerge. This especially applies to pleurodires for which no previous information is available. From comparisons with the limited data available for other freshwater turtles, marine turtles, and other aquatic reptiles, it does not appear that Sydney’s turtle populations have unusually high metal concentrations in tissues. Exclusion of toxic metals such as Pb from the egg may also be protective to the developing embryo. An ability to live in polluted habitats, while limiting the accumulation of toxic contaminants, may be one key to their persistence in urban waterways from which other freshwater fauna have disappeared. Reproductive impacts such as low embryo survival and small hatchling weights require more rigorous examination, but may have less effect on these animals which have such naturally high egg and hatchling mortality. Although it was generally hard to demonstrate biochemical, physiological or population impacts of contaminants, C. longicollis from a site with severe sewage pollution did display unusual alterations in a number of haematological variables, body condition, and carapace bone structure. Despite this, the population was large and had a comparatively high ratio of juveniles. Additionally, the adverse haematological alterations appeared reversible. Thus, successful populations in Sydney probably are more dependent on basic ecological needs being met, than on low levels of environmental contaminants. The ongoing persistence of chelid populations in Sydney is likely to be dependent to some extent on their opportunistic diets, which generally make animals less vulnerable to habitat modification and the reduction in prey item diversity following pollution (Mason 1996, Allanson & Georges 1999), with a further benefit possibly bestowed at some sites on E. macquarii by its omnivory.

The major question addressed by this project was to determine if the long-necked, freshwater turtle Chelodina oblonga, vocalise underwater and whether their vocal activity could be related to behavioural or ecological aspects of their lives. These turtles often live in wetlands where visibility is restricted due to habitat complexity or light limitation caused by factors such as tannin-staining, or turbidity. For many aquatic animals, sound is a useful means of communication over distances beyond their visual acuity. This thesis gives the first detailed account of the underwater vocal repertoire of C. oblonga. In total, over 230 days were spent in the field and more than 500 hours of tape recordings were made for this research. Initially, a number of recordings took place in three wetlands known to support turtle populations: Blue Gum Lake; Glen Brook Dam; and Lake Leschenaultia in Perth, Western Australia; in order to determine the nature of the freshwater sound field and place turtle vocalisations into the context in which they were vocalising. The wetlands differed in terms of degree of enrichment, substrate material, water depth and habitat complexity. Recordings were made over a four-week period in the last month of summer and the first week of autumn (Feb-Mar 2003). Invertebrate sweeps were also taken over a two-week period at each recording site to determine if invertebrate distributions were related to patterns of sonic activity. To determine the influence of wind on ambient noise; recordings were undertaken on winter mornings (June-August, 2003) at Blue Gum Lake and Glen Brook Dam at locations north, south, west and east for four different wind speeds – Beaufort Wind Scale (BWS) 0,1,2 & 3. There were seven distinctive calls recognised in the recordings. The frequency bandwidth most utilised by organisms was between 3 kHz up to around 14 kHz, with the exception of the ‘bird-like song’; which extended from 500 Hz up to around 10 kHz. Blue Gum Lake contained a more diverse and abundant assemblage of invertebrates than Lake Leschenaultia and Glen Brook Dam. Correspondingly, a greater diversity of calls was recorded at Blue Gum Lake, as well as the presence of chorus activity, which was not heard at the two less-enriched sites. The periods of greatest diversity and abundance of macroinvertebrates was synonymous with the increased sonic activity at dusk and midnight with noise levels greatest at dusk in particular, and to a lesser extent at midnight. There was no difference in ambient noise at Blue Gum Lake or Glen Brook Dam at wind speeds of Beaufort Wind Scale 0, 1 and 2. Turtles from three populations were recorded in artificial environments: consisting of round, plastic, above-ground ponds (1.8m dia. x 0.65m depth), which were set up to recreate small wetlands. Recordings occurred from September to October, 2003 and from February to December, 2004 as well as January, 2005. Seven hatchling and five juvenile turtles (CL <10cm) were also recorded in order to ascertain whether very young turtles vocalised. Hatchlings were recorded in a glass aquarium (35.5cm length x 20cm width x 22.0cm depth) and juveniles were placed into a below-ground outdoor pond (1m length x 0.5m width x 0.4m depth). Recordings occurred from as early as 4.30am (dawn recordings) to as late as 1.30am (evening recordings). The recordings revealed that turtles utilise an underwater acoustic communication system (calling at the water’s surface was also noted but these were not recorded or a part of this research) involving a repertoire of both complex and percussive sounds with short, medium and potentially long-range propagation characteristics. Complex structures included harmonically related elements (richly or sparsely) and different rates of frequency modulation. Frequency use extended beyond the in-air auditory sensitivity known for a single species of turtle studied from the family Chelidae; with calls ranging from around 100 Hz in some of the percussive displays, to as high as 3.5 kHz in some complex calls, with ‘clicks’ extending beyond the 20 kHz upper limit of the recording system. However, most of C. oblonga’s vocalisations had dominant frequencies below 1 kHz. Turtles were intermittent callers with an extensive vocal repertoire of seventeen (17) vocal categories - highly suggestive of complex social organisation. Vocalisations included: a) clacks; b) clicks; c) squawks; d) hoots; e) short chirps; f) high short chirps; g) medium chirps; h) long chirps; i) high calls; j) cries or wails; k) cat whines; l) grunts; m) growls; n) blow bursts; o) staccatos; p) a wild howl; and q) drum rolling. Also, two sustained ‘pulse-bouts’ were recorded during the breeding months, hypothesised to function as acoustic advertisement displays – possibly ‘calling songs’. Hatchling turtles were not heard to vocalise within the audible range. Only a single complex vocalisation was heard produced by the juvenile turtles, with a number of percussive calls. Preliminary playback trials were conducted under free-field conditions and within an artificial environment, which consisted of a below ground rectangular tank (2.4m length x 0.8m width x 0.6m deep). A number of turtle calls recorded in the artificial ponds were selected for playback. A UW 30 speaker was used for broadcast of calls. The free-field playbacks occurred at Mabel Talbot Lake and Blue Gum Lake during the months of April and May, 2005. Playback using 14 seconds of an artificially constructed sequence from the sustained ‘pulse-bout’ occurred in the artificial channels. This sequence consisted of some of the first phase pulses followed by a section of the ‘vibrato’. The preliminary free-field playback trials indicated that turtles had some interest in the calls being played by responding with an ‘alert posture’. Turtles were shown to remain in the alert posture for a significantly longer time than when no sound was played or when white noise was played. The extensive repertoire and initial responses to the free-field playbacks indicated that sound has some biological importance for C. oblonga, although results of playbacks under artificial conditions were inconclusive.

The major question addressed by this project was to determine if the long-necked, freshwater turtle Chelodina oblonga, vocalise underwater and whether their vocal activity could be related to behavioural or ecological aspects of their lives. These turtles often live in wetlands where visibility is restricted due to habitat complexity or light limitation caused by factors such as tannin-staining, or turbidity. For many aquatic animals, sound is a useful means of communication over distances beyond their visual acuity. This thesis gives the first detailed account of the underwater vocal repertoire of C. oblonga. In total, over 230 days were spent in the field and more than 500 hours of tape recordings were made for this research. Initially, a number of recordings took place in three wetlands known to support turtle populations: Blue Gum Lake; Glen Brook Dam; and Lake Leschenaultia in Perth, Western Australia; in order to determine the nature of the freshwater sound field and place turtle vocalisations into the context in which they were vocalising. The wetlands differed in terms of degree of enrichment, substrate material, water depth and habitat complexity. Recordings were made over a four-week period in the last month of summer and the first week of autumn (Feb-Mar 2003). Invertebrate sweeps were also taken over a two-week period at each recording site to determine if invertebrate distributions were related to patterns of sonic activity. To determine the influence of wind on ambient noise; recordings were undertaken on winter mornings (June-August, 2003) at Blue Gum Lake and Glen Brook Dam at locations north, south, west and east for four different wind speeds - Beaufort Wind Scale (BWS) 0,1,2 and 3. There were seven distinctive calls recognised in the recordings. The frequency bandwidth most utilised by organisms was between 3 kHz up to around 14 kHz, with the exception of the 'bird-like song'; which extended from 500 Hz up to around 10 kHz. Blue Gum Lake contained a more diverse and abundant assemblage of invertebrates than Lake Leschenaultia and Glen Brook Dam. Correspondingly, a greater diversity of calls was recorded at Blue Gum Lake, as well as the presence of chorus activity, which was not heard at the two less-enriched sites. The periods of greatest diversity and abundance of macroinvertebrates was synonymous with the increased sonic activity at dusk and midnight with noise levels greatest at dusk in particular, and to a lesser extent at midnight. There was no difference in ambient noise at Blue Gum Lake or Glen Brook Dam at wind speeds of Beaufort Wind Scale 0, 1 and 2. Turtles from three populations were recorded in artificial environments: consisting of round, plastic, above-ground ponds (1.8m dia. x 0.65m depth), which were set up to recreate small wetlands. Recordings occurred from September to October, 2003 and from February to December, 2004 as well as January, 2005. Seven hatchling and five juvenile turtles (CL < 10cm) were also recorded in order to ascertain whether very young turtles vocalised. Hatchlings were recorded in a glass aquarium (35.5cm length x 20cm width x 22.0cm depth) and juveniles were placed into a below-ground outdoor pond (1m length x 0.5m width x 0.4m depth). Recordings occurred from as early as 4.30am (dawn recordings) to as late as 1.30am (evening recordings). The recordings revealed that turtles utilise an underwater acoustic communication system (calling at the water's surface was also noted but these were not recorded or a part of this research) involving a repertoire of both complex and percussive sounds with short, medium and potentially long-range propagation characteristics. Complex structures included harmonically related elements (richly or sparsely) and different rates of frequency modulation. Frequency use extended beyond the in-air auditory sensitivity known for a single species of turtle studied from the family Chelidae; with calls ranging from around 100 Hz in some of the percussive displays, to as high as 3.5 kHz in some complex calls, with 'clicks' extending beyond the 20 kHz upper limit of the recording system. However, most of C. oblonga's vocalisations had dominant frequencies below 1 kHz. Turtles were intermittent callers with an extensive vocal repertoire of seventeen (17) vocal categories - highly suggestive of complex social organisation. Vocalisations included: a) clacks; b) clicks; c) squawks; d) hoots; e) short chirps; f) high short chirps; g) medium chirps; h) long chirps; i) high calls; j) cries or wails; k) cat whines; l) grunts; m) growls; n) blow bursts; o) staccatos; p) a wild howl; and q) drum rolling. Also, two sustained 'pulse-bouts' were recorded during the breeding months, hypothesised to function as acoustic advertisement displays - possibly 'calling songs'. Hatchling turtles were not heard to vocalise within the audible range. Only a single complex vocalisation was heard produced by the juvenile turtles, with a number of percussive calls. Preliminary playback trials were conducted under free-field conditions and within an artificial environment, which consisted of a below ground rectangular tank (2.4m length x 0.8m width x 0.6m deep). A number of turtle calls recorded in the artificial ponds were selected for playback. A UW 30 speaker was used for broadcast of calls. The free-field playbacks occurred at Mabel Talbot Lake and Blue Gum Lake during the months of April and May, 2005. Playback using 14 seconds of an artificially constructed sequence from the sustained 'pulse-bout' occurred in the artificial channels. This sequence consisted of some of the first phase pulses followed by a section of the 'vibrato'. The preliminary free-field playback trials indicated that turtles had some interest in the calls being played by responding with an 'alert posture'. Turtles were shown to remain in the alert posture for a significantly longer time than when no sound was played or when white noise was played. The extensive repertoire and initial responses to the free-field playbacks indicated that sound has some biological importance for C. oblonga, although results of playbacks under artificial conditions were inconclusive.

The major question addressed by this project was to determine if the long-necked, freshwater turtle Chelodina oblonga, vocalise underwater and whether their vocal activity could be related to behavioural or ecological aspects of their lives. These turtles often live in wetlands where visibility is restricted due to habitat complexity or light limitation caused by factors such as tannin-staining, or turbidity. For many aquatic animals, sound is a useful means of communication over distances beyond their visual acuity. This thesis gives the first detailed account of the underwater vocal repertoire of C. oblonga. In total, over 230 days were spent in the field and more than 500 hours of tape recordings were made for this research. Initially, a number of recordings took place in three wetlands known to support turtle populations: Blue Gum Lake; Glen Brook Dam; and Lake Leschenaultia in Perth, Western Australia; in order to determine the nature of the freshwater sound field and place turtle vocalisations into the context in which they were vocalising. The wetlands differed in terms of degree of enrichment, substrate material, water depth and habitat complexity. Recordings were made over a four-week period in the last month of summer and the first week of autumn (Feb-Mar 2003). Invertebrate sweeps were also taken over a two-week period at each recording site to determine if invertebrate distributions were related to patterns of sonic activity. To determine the influence of wind on ambient noise; recordings were undertaken on winter mornings (June-August, 2003) at Blue Gum Lake and Glen Brook Dam at locations north, south, west and east for four different wind speeds - Beaufort Wind Scale (BWS) 0,1,2 and 3. There were seven distinctive calls recognised in the recordings. The frequency bandwidth most utilised by organisms was between 3 kHz up to around 14 kHz, with the exception of the 'bird-like song'; which extended from 500 Hz up to around 10 kHz. Blue Gum Lake contained a more diverse and abundant assemblage of invertebrates than Lake Leschenaultia and Glen Brook Dam. Correspondingly, a greater diversity of calls was recorded at Blue Gum Lake, as well as the presence of chorus activity, which was not heard at the two less-enriched sites. The periods of greatest diversity and abundance of macroinvertebrates was synonymous with the increased sonic activity at dusk and midnight with noise levels greatest at dusk in particular, and to a lesser extent at midnight. There was no difference in ambient noise at Blue Gum Lake or Glen Brook Dam at wind speeds of Beaufort Wind Scale 0, 1 and 2. Turtles from three populations were recorded in artificial environments: consisting of round, plastic, above-ground ponds (1.8m dia. x 0.65m depth), which were set up to recreate small wetlands. Recordings occurred from September to October, 2003 and from February to December, 2004 as well as January, 2005. Seven hatchling and five juvenile turtles (CL < 10cm) were also recorded in order to ascertain whether very young turtles vocalised. Hatchlings were recorded in a glass aquarium (35.5cm length x 20cm width x 22.0cm depth) and juveniles were placed into a below-ground outdoor pond (1m length x 0.5m width x 0.4m depth). Recordings occurred from as early as 4.30am (dawn recordings) to as late as 1.30am (evening recordings). The recordings revealed that turtles utilise an underwater acoustic communication system (calling at the water's surface was also noted but these were not recorded or a part of this research) involving a repertoire of both complex and percussive sounds with short, medium and potentially long-range propagation characteristics. Complex structures included harmonically related elements (richly or sparsely) and different rates of frequency modulation. Frequency use extended beyond the in-air auditory sensitivity known for a single species of turtle studied from the family Chelidae; with calls ranging from around 100 Hz in some of the percussive displays, to as high as 3.5 kHz in some complex calls, with 'clicks' extending beyond the 20 kHz upper limit of the recording system. However, most of C. oblonga's vocalisations had dominant frequencies below 1 kHz. Turtles were intermittent callers with an extensive vocal repertoire of seventeen (17) vocal categories - highly suggestive of complex social organisation. Vocalisations included: a) clacks; b) clicks; c) squawks; d) hoots; e) short chirps; f) high short chirps; g) medium chirps; h) long chirps; i) high calls; j) cries or wails; k) cat whines; l) grunts; m) growls; n) blow bursts; o) staccatos; p) a wild howl; and q) drum rolling. Also, two sustained 'pulse-bouts' were recorded during the breeding months, hypothesised to function as acoustic advertisement displays - possibly 'calling songs'. Hatchling turtles were not heard to vocalise within the audible range. Only a single complex vocalisation was heard produced by the juvenile turtles, with a number of percussive calls. Preliminary playback trials were conducted under free-field conditions and within an artificial environment, which consisted of a below ground rectangular tank (2.4m length x 0.8m width x 0.6m deep). A number of turtle calls recorded in the artificial ponds were selected for playback. A UW 30 speaker was used for broadcast of calls. The free-field playbacks occurred at Mabel Talbot Lake and Blue Gum Lake during the months of April and May, 2005. Playback using 14 seconds of an artificially constructed sequence from the sustained 'pulse-bout' occurred in the artificial channels. This sequence consisted of some of the first phase pulses followed by a section of the 'vibrato'. The preliminary free-field playback trials indicated that turtles had some interest in the calls being played by responding with an 'alert posture'. Turtles were shown to remain in the alert posture for a significantly longer time than when no sound was played or when white noise was played. The extensive repertoire and initial responses to the free-field playbacks indicated that sound has some biological importance for C. oblonga, although results of playbacks under artificial conditions were inconclusive.

The life history theory studies the characteristics combinations that maximize the organisms reproduction and survival in the environment where they live. The species use the habitat in a diverse way and this use depends on the environmental conditions in the available habitats. Phrynops tuberosus, a freshwater turtle found in the north-northeastern regions of Brazil, is the animal-model chosen for the life history traits and habitat use studies. This work aims understand how this species reproduce, maintain and use the habitat in Caatinga. The study generated biological and ecological knowledge of the species and allowed the testing of hypotheses and predictions of life history traits, sexual dimorphism and habitat use. Selection of clutching sites, males smaller than females and preference of larger animals for environments most exposed were found in the populations studied. The results facilitate understanding of reproduction and survival of turtles in a semiarid environment.
A teoria de histÃria de vida estuda as combinaÃÃes de caracterÃsticas que maximizam a reproduÃÃo e a sobrevivÃncia dos organismos no ambiente onde vivem. As espÃcies utilizam o ambiente de modo diverso, e esse uso depende das condiÃÃes existentes nos hÃbitats disponÃveis. Phrynops tuberosus, cÃgado encontrado nas regiÃes norte-nordeste do Brasil, Ã o animal-modelo escolhido para o estudo dos traÃos de histÃria de vida e de uso de hÃbitat. Esse trabalho objetiva entender como a espÃcie se reproduz, organiza-se e utiliza o hÃbitat na caatinga. O estudo gerou conhecimentos biolÃgicos e ecolÃgicos da espÃcie e permitiu o teste de hipÃteses e previsÃes sobre traÃos de histÃria de vida, dimorfismo sexual e uso de hÃbitat. SeleÃÃo de locais de desova, machos menores que fÃmeas e preferÃncia de animais maiores por ambientes mais expostos foram encontradas nas populaÃÃes estudadas. Os resultados encontrados facilitam a compreensÃo da reproduÃÃo e da sobrevivÃncia dos quelÃnios em um ambiente do semiÃrido.

An ecological study was undertaken on four populations of Krefft�s River Turtle Emydura krefftii inhabiting the Townsville Area of Tropical North Queensland. Two sites were located in the Ross River, which runs through the urban areas of Townsville, and two sites were in rural areas at Alligator Creek and Stuart Creek (known as the Townsville Creeks). Earlier studies of the populations in Ross River had determined that the turtles existed at an exceptionally high density, that is, they were superabundant, and so the Townsville Creek sites were chosen as low abundance sites for comparison. The first aim of this study was to determine if there had been any demographic consequences caused by the abundance of turtle populations of the Ross River. Secondly, the project aimed to determine if the impoundments in the Ross River had affected the freshwater turtle fauna. Specifically this study aimed to determine if there were any difference between the growth, size at maturity, sexual dimorphism, size distribution, and diet of Emydura krefftii inhabiting two very different populations. A mark-recapture program estimated the turtle population sizes at between 490 and 5350 turtles per hectare. Most populations exhibited a predominant female sex-bias over the sampling period. Growth rates were rapid in juveniles but slowed once sexual maturity was attained; in males, growth basically stopped at maturity, but in females, growth continued post-maturity, although at a slower rate. Sexual maturity was at 6-7 years of age for males, which corresponded to a carapace length of 150-160 mm, and 8-10 years of age for females, which corresponded to a carapace length of 185-240 mm. The turtles were omnivorous, although in the Ross River they ate more submerged vegetation (by percent amount and occurrence) than those of the Townsville Creeks. Turtles in Townsville Creeks ingested more windfall fruit and terrestrial insects.

Cette étude a pour objectifs d’actualiser les données sur la répartition et la variation génétique des deux espèces de tortues d’eau douce Mauremys leprosa et Emys orbicularis en Tunisie et d’évaluer leur vulnérabilité. Elle a également pour but de déterminer la diversité et d’évaluer la spécificité des hémogrégarines (Protozoa: Apicomplexa: Haemogregarinidae) en se basant sur les analyses morphologiques et moléculaires. L’Emyde lépreuse n’est actuellement pas menacée en Tunisie contrairement à la Cistude d’Europe qui est en situation critique. Les analyses morphologiques et moléculaires des parasites sanguins ont permis de montrer la présence de co-infection chez un même individu de tortue. L’aspect moléculaire a concerné la caractérisation et l’identification des espèces appartenant au même genre Haemogregarina à travers l’utilisation des marqueurs mitochondriaux et nucléaires. Ainsi on a pu comparer le degré d’informations génétiques des deux marqueurs et de montrer que le gène mitochondrial COI permettait d’identifier plus de variabilité que celle trouvée avec le gène 18S ARNr. En outre, la faible spécificité de ces parasites sanguins envers les espèces hôtes définitives et intermédiaires, s’est révélée un facteur qui augmente le risque de transmission des hémogregarines d’une espèce hôte exotique vers une espèce hôte indigène
The objective of this study is to update data on distribution and genetic variation of t two species of freshwater turtles Mauremys leprosa and Emys orbicularis in Tunisia and to assess their vulnerability. It also aims to determine the diversity and to evaluate the specificity of the hemogregarins (Protozoa: Apicomplexa: Haemogregarinidae) based on morphological and molecular analyzes. The Mediterranean pond turtle is currently not threatened in Tunisia unlike the European pond turtle which is in critical situation. Morphological and molecular analyzes of blood parasites have reveald the presence of co-infection in the same turtle specimen. The molecular aspect concerned the characterization and identification of species belonging to the same genus Haemogregarina through the use of mitochondrial and nuclear markers. Thus we were able to compare the level of genetic information of the two markers and to demonstrat that the mitochondrial COI genes identified more variability than those found with the 18S rRNA gene. In addition, the low specificity of these blood parasites towards the final and intermediate host species has been shown to be a factor which increases the risk of transmission of hemogregarins from an exotic host species to a native host species

An ecological study of Emydura macquarii macquarii in the south-east region of Australia was conducted between October 1995 and March 1998. E. m. macquarii is an abundant and widespread species of short-necked turtle that is highly variable in morphology and related life history attributes. No study in Australia had previously looked at geographic variation in biological traits in freshwater turtles, hence the level of variation in E. m. macquarii had been poorly documented. The principal aims of this study were to investigate the plasticity of life history traits across populations of E. m. macquarii and to speculate on possible causes. A more intensive study was also conducted on a rare and suspected declining population of E. m. macquarii in the Nepean River to determine whether relevant management and conservation measures; were required. The study involved comparing various life history attributes between five populations of E. m. macquarii (Brisbane River, Macleay River, Hunter River, Nepean River and Murray River). The populations were specifically chosen to account for the range of variation in body size within this subspecies. Body size (maximum size, size at maturity, growth rates), population structures (sex ratios, age and size structures), reproductive traits (clutch mass, clutch size, egg size, egg content, etc.) and other attributes were collected for each population. Patterns of life history traits, both within and among populations, were explored so that causes of variation could be sought. Geographic variation in Body Size and other Related Life History Traits Body size in E. m. macquarii differed markedly between populations. Females ranged in maximum sizes (carapace length) of 180 mm in the Macleay River to over 300 mm in the Murray River. E. m. macquarii was sexually dimorphic across all populations with females larger than males in all cases. Maximum body size was positively related to the size at which a turtle matures. The size at maturity in turn was positively related to juvenile growth rates. Age was a more important factor for males in terms of timing of maturity whereas in females it was body size. Morphological variation was not only great between populations, but also within populations. Maximum body size was unrelated to latitude; hence it was inferred that habitat productivity had the most important influence on geographic variation in body size. Population structures also differed between populations. Sex ratios did not differ in the Brisbane, Macleay and Murray Rivers. However, a male bias was present in the Nepean River population and a female bias in the Hunter River. Juveniles were scarce in the Brisbane and Macleay Rivers but numerous in the Nepean and Hunter Rivers. Geographic Variation in Reproduction There was large variation in reproductive traits across populations of E. m. macquarii. Nesting season began as early as mid-September in the Brisbane River and as late as December in the Hunter River, and continued until early January. Populations in the Hunter and Murray Rivers are likely to produce only one clutch per season while populations from the Macleay and Nepean Rivers can produce two, and on some occasions, three clutches annually. The majority of females would appear to reproduce every year. Clutch mass, clutch size, and egg size varied greatly both within and among populations. A large proportion of variation in reproductive traits was due to the effects of body size. E. m. macquarii from large-bodied populations such as in the Brisbane and Murray Rivers produced bigger eggs than small-bodied populations. Within a population, clutch mass, clutch size, and egg size were all correlated with body size, except the Nepean River. The variability of egg size was smaller in large-bodied populations where egg size was more constant. Not all variation in reproductive traits was due to body size. Some of this variation was due to annual differences within a population. Reproductive traits within a population are relatively plastic, most likely a result of changing environmental conditions. Another source is the trade-off between egg size and clutch size. A negative relationship was found between egg size and clutch size (except the Brisbane River). Reproductive variation was also influenced by latitudinal effects. Turtles at lower latitudes produces more clutches, relatively smaller clutch sizes, clutch mass and larger eggs than populations at higher latitudes. Annual reproductive output is greater in tropical populations because they can produce more clutches per year in an extended breeding season. Eggs that were incubated at warmer temperatures hatched faster and produced smaller hatchlings. Incubation temperatures above 30�C increased egg mortality and hatchling deformities, suggesting this is above the optimum developmental temperature for E. m. macquarii. Hatchling size was positively related to egg size, hence hatchling sizes was on average larger in the Murray and Brisbane rivers. However, population differences remained in hatchling size after adjustments were made for egg size. For example, hatchlings from the Hunter River were smaller than those from the Macleay River despite the egg size being the same. These differences were most likely due to the shorter incubation periods of hatchlings from the Hunter River. Nepean River The Nepean River population of E. m. macquarii is at the southern coastal limit of its range. This is a locally rare population, which is believed to be declining. This study aimed at determining the distribution, abundance, and population dynamics to assess whether any conservation management actions were required. E. m. macquarii in the Nepean River was mainly concentrated between Penrith and Nortons Basin, although even here it was found at a very low density (10.6 - 12.1 per hectare). The largest male caught was 227 mm while the largest female was 260.4 mm. Males generally mature between 140 - 150 mm in carapace length and at four or five years of age. Females mature at 185 -195 mm and at six to seven years of age. Compared with other populations of E. macquarii, Nepean River turtles grow rapidly, mature quickly, are dominated by juveniles, have a male bias and have a high reproductive output. Far from being a population on the decline, the life history traits suggest a population that is young and expanding. There are considered to be two possible scenarios as to why the Nepean River population is at such a low density when it appears to be thriving. The first scenario is that the distribution of the population on the edge of its range may mean that a small and fluctuating population size may be a natural feature due to sub-optimal environmental conditions. A second scenario is that the population in the Nepean River has only recently become established from dumped pet turtles.

Most studies of wetlands tend to focus on the biotic and abiotic interactions within the aquatic habitat. Though wetlands and associated biota may appear to be somewhat isolated from the influence of the wider landscape, wetland habitats are critically linked with adjacent terrestrial habitats and other wetlands through the two-way flows of energy and nutrients and provision of structure. While an understanding of these inter-habitat linkages is breaking down the perceived boundaries between "aquatic" and "terrestrial" ecosystems, there is more limited knowledge on the ecology of wetland animals that must meet critical needs in both aquatic and terrestrial habitats at some time during their life or seasonal cycles. Here, I examine the terrestrial ecology of a freshwater turtle, the eastern long-necked turtle (Chelodina longicollis) in the temporally dynamic and heterogeneous landscape of Booderee national park in south-east Australia by 1) providing a description of terrestrial behaviours, 2) identifying the factors driving terrestrial behaviour and its functional significance, 3) examining factors that may limit or constrain terrestrial behaviour and 4) demonstrating how various terrestrial behaviours can factor prominently in the overall biology of a nominally aquatic animal. Chelodina longicollis used terrestrial habitats for reasons other than nesting, including aestivation and movements between wetlands. Radio-telemetry of 60 turtles revealed that nearly 25 % of all locations were in terrestrial habitats up to 505 m from the wetland, where turtles remained for extended periods (up to 480 consecutive days) buried under sand and leaf litter in the forest. Individuals also maintained an association with a permanent lake and at least one temporary wetland within 1470 m, though some inter-wetland dispersal movements were much longer (5248 m). As a result of their associations with several wetlands and terrestrial aestivation sites, C. longicollis traversed large areas and long distances (13.8 +/- 2.8 ha home range, 2608 +/- 305 m moved), indicating that this species is highly vagile. In fact, a three-year capture-mark-recapture study conducted in 25 wetlands revealed that 33% of the population moved overland between wetlands. After scaling this rate to the number of generations elapsed during the study, C. longicollis moved between discrete water bodies at a rate of 88-132% per generation. This rate is not only high for freshwater turtles, but is among the highest rates of inter-patch movement for any vertebrate or invertebrate. Chelodina longicollis demonstrated an impressive capacity for individual variation in nearly every aspect of its behaviour examined. Most of the variation in space use, movements, terrestrial aestivation and activity could be attributed to extrinsic local and landscape factors, seasonal influences and rainfall, whereas intrinsic attributes of the individual such as sex, body size, body condition and maturity status were less important. Turtles increased movement distance and home range size in regions where inter-wetland distances were farther and with increasing wetland size. Individuals spent more time in terrestrial habitats with decreasing wetland hydroperiod and increasing distance to the nearest permanent lake. Overland movements between wetlands were correlated with rainfall, but the directionality of these movements and the frequency with which they occurred varied according to the prevalent rainfall patterns; movements were to permanent lakes during drought, but turtles returned to temporary wetlands en masse upon the return of heavy rainfall. However, deteriorating conditions in drying wetlands forced turtles to move even in the absence of rainfall. Captures at a terrestrial drift fence revealed that immature turtles as small as 72.3 mm plastron length may move overland between wetlands with similar frequency as larger adults. Taken together, these results suggest that C. longicollis behaviour is in part conditional or state-dependent (i.e., plastic) and shaped by the spatiotemporal variation and heterogeneity of the landscape. Perhaps the most surprising aspect of individual variation was the alternate responses to wetland drying. Turtles either aestivated in terrestrial habitats (for variable lengths of time), or moved to other wetlands. Movement to other wetlands was the near universal strategy when only a short distance from permanent lakes, but the proportion of individuals that aestivated terrestrially increased with distance to the nearest permanent lake. When long distances must be travelled, both behaviours were employed by turtles in the same wetland, suggesting that individuals differentially weigh the costs and benefits of residing terrestrially versus those of long-distance movement. I propose that diversity in response to wetland drying in the population is maintained by stochastic fluctuations in resource quality. The quality of temporary wetlands relative to permanent wetlands at our study site varies considerably and unpredictably with annual rainfall and with it the cost-benefit ratio of each strategy or tactic. Residency in or near temporary wetlands is more successful during wet periods due to production benefits (high growth, reproduction and increased body condition), but movement to permanent wetlands is more successful, or least costly, during dry periods due to the fitness benefits of increased survival and body condition. I used the doubly-labelled water (DLW) method to provide the first estimates of water and energy costs of aestivation and overland movement for any freshwater turtle behaving naturally in the field. Chelodina longicollis remained hydrated while terrestrial with water flux rates (14.3-19.3 ml kg-1 d-1) on par with those of strictly terrestrial turtles, but field metabolic rate during aestivation (20.0-24.6 kJ kg-1 d-1) did not indicate substantial physiological specializations in metabolism during aestivation. Energy reserves, but not water, are predicted to limit survival in aestivation to an estimated 49-261 days, which is in close agreement with the durations of natural aestivation. The energy costs of overland movement were 46-99 kJ (kg d)-1, or 1.6-1.7 times more expensive than aestivation. When a wetland dries, a turtle that foregoes movement to other wetlands can free sufficient energy to fuel up to 134 days in aestivation. The increasing value of this energy "trade-off" with travel distance fits our behavioural observations of variance in response to wetland drying. Taken together, this evidence indicates that terrestrial habitats provide more than just organic and structural inputs and filtering services and that nearby wetlands are important for reasons other than potential sources of occasional colonists to a population. Terrestrial habitats are used for aestivation in response to wetland drying and different wetlands are diverse in their functions of meeting the annual or life-cycle requirements of C. longicollis in temporally dynamic wetland systems. As overland movements between these various habitat types are in response to spatiotemporal variation in habitat quality and associated shifts in the fitness gradient between them, I suggest that terrestrial and different aquatic habitats in Booderee offer complementary resources contributing to regional carrying capacity and population persistence of the turtle population. Thus, important ecological processes regulating C. longicollis in a focal wetland should not be viewed as operating independently of other nearby wetlands and their adjacent terrestrial habitats. Collectively, these findings highlight the complex and dynamic associations between a population of freshwater turtles and the wider terrestrial and aquatic landscape, demonstrating that turtle populations and the factors that impact them can extend well beyond the boundaries of a focal wetland.

Unlike the majority of vertebrates, the freshwater turtle (Trachemys scripta) can survive anoxia for hours at warm acclimation temperatures and weeks at cold acclimation temperatures. During anoxia exposure, the turtle heart continues its role in internal convection, but systemic cardiac output and systemic cardiac power output are massively reduced, primarily due to a decreased heart rate (ƒH). Therefore, control of ƒH is critical to cardiac energy management during anoxia. This thesis investigated what extrinsic, autocrine/paracrine and intrinsic mechanisms control the cardiovascular system of anoxic freshwater turtles. Cardiac control was examined at the level of the whole animal, organ (isolated heart chambers), and cell (isolated cardiac myocytes). A 2 x 2 exposure design allowed comparisons between 21°C- and 5°C-acclimated turtles under normoxia and anoxia. For warm-acclimated turtles, I discovered a re-setting of intrinsic ƒH that accounts for up to 57% of the anoxic bradycardia, which, when combined with cholinergic cardiac inhibition (previously known), fully explains the depression of ƒH with anoxia. Interestingly, prolongation of ventricular action potential duration (APD) by 47% with anoxia was proportional to the reduction in intrinsic ƒH. My thesis also revealed how cold acclimation prepared cardiac muscle for winter anoxic conditions. Cold temperature decreased intrinsic ƒH, prolonged cardiac APDs and reduced the chronotropic sensitivity to extracellular anoxia and acidosis. Also, the decreased peak densities of ventricular I[sub Na] and I[sub Ca] and conductance of i[sub KI] observed in 5°C-acclimated turtle hearts compared with 21°C-acclimated turtle hearts could serve to conserve the ATP cost of ion pumping. When cold-acclimated turtles were exposed to anoxia, during which cholinergic cardiovascular control is blunted, my thesis discovered that a re-setting of intrinsic ƒH accounts for up to 66% of the anoxic bradycardia. However, contrary to 21°C, there was no prolongation of cardiac APDs. No evidence was found for either α-adrenergic or adenosinergic cardiac inhibition in cold-acclimated anoxic turtles, yet their reduction of in vivo cardiac activity correlated with alterations in myocardial high-energy phosphate metabolism, intracellular pH and free energy of ATP hydrolysis. These novel insights point to the importance of future studies on pacemaker currents and cardiac refractoriness.
Science, Faculty of
Zoology, Department of
Graduate

In aquatic turtles, females select nest sites that have a high degree of solar exposure, and exploit recently tilled agricultural fields for nesting, presumably because of increased solar exposure and/or easier nest excavation, and the importance of incubation temperature on survival and offspring phenotype. These same disturbed sites are often contaminated by pollutants and turtles can incorporate high levels of pollutants into their eggs which negatively impact hatch success. For my M.S. research, I investigated turtle nest site selection in a system dominated by agricultural and industrial land use, the impact of crop growth on the thermal and hydric dynamics of turtle nests, and I used paired field and laboratory experiments to examine the individual and interactive impacts of agricultural land use and Hg contamination on hatch success and offspring phenotype in Chelydra serpentina. Of the 150 turtle nests found during this research, 84% were located in human-disturbed soils. Nest site characteristics were similar among nests found in Hg contaminated and reference areas. Agriculture and control nests did not differ in temperature at the time of nesting, but temperatures diverged as crops grew, with temperatures in nests in agricultural fields averaging 2.5 °C lower than control nests over the course of incubation. Similarly, despite no initial difference, nest moisture levels diverged throughout incubation and moisture averaged 107 kPa lower in agricultural than control soils throughout incubation. In my field and laboratory experiments, I found that in comparison to turtles from control incubation conditions (i.e., warmer), turtles incubated under agricultural thermal regimens (i.e., colder) took longer to hatch, hatched at smaller structural body sizes, lost more mass after hatching, had lower post-hatching structural growth rates, and were more likely to be male. Additionally, thermal conditions associated with agricultural land use interacted with high levels of mercury to impact hatching success and offspring sex ratios. My thesis research provides one of the first documentations of negative interactive effects of mercury pollution and habitat quality on early vertebrate development and highlights the importance of examining the combined influence of multiple global changes on biological systems.
Master of Science

I studied aspects of the ecology of the Murray River turtle, Emydura macquarii, to determine the impact of the introduced red fox, Vulpes vulpes. The fox is one of Australia's worst vertebrate pests through its predation on livestock and native mammals, but their impact on reptilian communities is not known. I conducted a large-scale mark-recapture study to evaluate population growth of E. macquarii in the Albury region of the upper Murray River by determining growth, reproduction and survival. The study was conducted downstream of the first, and largest, impoundment on the Murray River, Lake Hume. Emydura macquarii predominantly inhabit the lagoons in the upper Murray River, as the mainstream and Lake are possibly too cool to maintain metabolic processes. They are easily captured in hoop traps and the use of live decoys maximises trap success. Over 2000 hatchling turtles were marked and released into two lagoons between January 1997 and January 1998. Growth of these individuals is rapid over the first few years but declines towards maturity, and is indeterminate after maturity. Although growth annuli are not well defined, even on young individuals, the von Bertalanffy model describes the growth of both male and female E. macquarii. Male turtles mature at 5-6 years and females mature at 10-12 years. Female turtles may maximise reproductive potential by delaying maturity and producing one relatively large clutch (mean = 21 eggs) per year, which is positively correlated with body size (PL). Although primarily related to body size, clutch size varies annually because of environmental conditions. If winter and summer rainfalls are below average and temperatures are above average, E. macquarii may reduce clutch size to increase the chance of the eggs surviving. Nesting predominantly occurs during the first major rain-bearing depression in November. Habitat variables, including distance from water, nearest nest, and tree, and soil type were measured for each nest to determine characteristics that attract predators. Nests close to the shoreline and trees are heavily preyed on, and nests constructed in sand are less likely to be destroyed by predators. Foxes detect nests through a combination of chemical cues from eggs and slight soil disturbances, whereas birds only destroy nests observed being constructed during the day. Female turtles alter nesting behaviour and construct nests much further away from water when foxes were removed and as a result, nests are less dense and away from trees. Thus in high predation risk areas, turtles minimise emergence and search times to reduce the risk of direct predation by foxes. Predation is reduced when nests are in lower densities and away from trees, because predators increase search efforts when nests are in higher densities and birds are more likely to destroy nests close to trees. Reproductive success is further reduced in high predation risk areas because more nests are constructed in sandy substrates where clutch success is reduced compared to incubation in more dense substrates. Where predators are a significant source of mortality, prey may use indirect methods, such as chemical recognition, to avoid encounters. Nesting turtles did not avoid areas where fox odour was present, suggesting that they assess predation pressure from foxes by other mechanisms, such as visual recognition. However, an innate response occurs to the odour of a once common predator on the Murray River, the eastern quoll (Dasyurus viverrinus), whereby turtles recognise and avoid nesting in areas where quoll odour is present. Therefore nesting turtles show a similar avoidance response to two different predators, using different mechanisms of detection. Similarly, predation risk may influence hatching times and nest emergence. The rate of embryonic development of E. macquarii may increase or eggs may hatch early so that the clutch hatches synchronously, thereby reducing the risk of predation through group emergence from the nest. Emydura macquarii reach densities of over 100 turtles.ha-1, with the majority of the population consisting of sexually mature individuals. Emydura macquarii has a Type III survival curve where mortality is extremely high in the egg stage (93% nest predation), remaining high over the hatchling stage (minimum survival rate- 10%), but decreasing rapidly throughout the juvenile stage (~70% juvenile survival). Adult survival is extremely high, with greater than 95% of adults surviving each year. Foxes through nest predation cause most mortality but a small proportion (~3%) of nesting adult females are killed by foxes each year. A removal program evaluated the impact of foxes. In 1996, fox numbers were monitored around four lagoons by spotlighting and non-toxic bait uptake. Foxes were removed from around two of the lagoons throughout 1997 and 1998, using spotlight shooting and 1080 bait poisoning. Fox numbers were continually monitored around all four lagoons during the study. Nest predation rates remained around 90% in all sites where foxes were present, but fell to less than 50% when foxes were removed. At the same time, predation on nesting female turtles was eliminated where foxes were removed. Demographic models using staged based survival schedules, together with growth and fecundity values for E. macquarii show a decline of 4% per year in these populations. Elasticity analyses shows that survival of adult female E. macquarii has the major influence on population stability and a reduction of nest predation alone is unlikely to address the population decline. Management options, such as reducing foxes prior to nesting around key lagoons, will stabilise the population decline, and eliminating foxes completely from certain areas with high dispersal potential, will promote recruitment of juvenile E. macquarii.

Com o objetivo de obter dados relacionados às estratégias reprodutivas de Hydromedusa maximiliani no Parque Estadual Carlos Botelho, SP, fêmeas e jovens foram radiografados entre setembro de 2007 e dezembro de 2008. A estação reprodutiva estendeu-se de setembro a dezembro, com a ocorrência de desovas entre novembro e dezembro, tanto em 2007 como em 2008. Das 33 fêmeas radiografadas, 19 (58%) apresentaram ovos, e, dessas, apenas três (16%) reproduziram-se nos dois anos. O tamanho da ninhada variou entre 1 e 3 ovos, mas a maioria das fêmeas (90%) apresentou dois ovos. O volume da ninhada foi de 22128,01 ± 4939,8 mm³ (10493,422136,12 mm³; N = 44). Houve fortes relações positivas entre o tamanho corpóreo das fêmeas com o tamanho dos ovos e volume da ninhada. Os dados reprodutivos obtidos no presente estudo, em conjunto com os dados populacionais estabelecidos para H. maximiliani na mesma área de estudo por outros autores, permitiram a realização de uma análise de viabilidade da população. As simulações geradas no programa VORTEX (versão 9.92) foram baseadas em dois cenários principais, denominados População Real (PR) e População Hipotética Isolada (PHI). O primeiro foi criado com base nas informações existentes para H. maximiliani, coletadas ao longo de mais de 10 anos, a partir do qual foram feitas alterações de algumas variáveis (aumento de 10% na taxa de mortalidade; aumento em duas vezes no valor de frequência de catástrofes; inclusão de depressão por endogamia com aumento de 10% no valor default do programa e diminuição da capacidade de suporte), gerando o cenário PHI, com o objetivo de avaliar as diferenças entre uma população protegida em uma unidade de conservação e uma população isolada e sujeita a maiores influências negativas. Os resultados das simulações demonstraram existir diferença significativa (p = 0,0005) entre os dois cenários com probabilidade de extinção equivalente a 31% para o cenário real e 90% para o cenário hipotético. Testes de sensibilidade foram realizados com a finalidade de verificar a influência de variações na taxa de mortalidade, catástrofes e depressão por endogamia, e forneceram indicações de que variações em todos os parâmetros testados possuem forte influência na sobrevivência da população. Estudos de longaduração e informações a respeito das espécies de áreas sujeitas a fortes pressões antrópicas, como a Mata Atlântica, são de grande importância para a conservação, já que podem auxiliar na elaboração de análises de viabilidade populacional e no delineamento de estratégias de manejo focadas nos fatores reais que ameaçam as populações.
The reproductive strategies of Hydromedusa maximiliani at the Carlos Botelho State Park, SP, was verified by X-ray in females and juveniles between September 2007 and December 2008. Throughout the study two reproductive seasons were detected, both beginning in September. From a total of 33 X-rayed females, only 19 showed eggs, and three of them showed eggs in the two reproductive seasons. Clutch size varied from 1 to 3 eggs, but only one female presented one egg and another female presented three eggs. Clutch volume was 22128.01 ± 4939.8 mm³ (10493.422136.12 mm³; N = 44). Female body size was positively correlated to clutch volume, a pattern usually detected in chelonians. Reproductive information acquired in the present study together with population data available for H. maximiliani in the study site makes it possible a population viability analysis using the software VORTEX (version 9.92). A principal scenario (Real Population) was created based on life-history data collected throughout 10 years. In order to compare the protected area with an unprotected hypothetical area regarding the persistence of the population, this scenario was modified by altering some variables (10% increasing of mortality rate, 10% increasing of inbreeding depression, decreasing of carrying capacity), which generates a scenario named Hypothetical Isolated Population. Results were compared and sensitivity tests were carried out to verify the influence of variation in mortality ratio, catastrophes and inbreeding depression. There was a significant difference between the two scenarios (p = 0.0005). The extinction risk was 31% for Real Population and 90% for Hypothetical Isolated Population. The sensitivity analysis indicated that mortality ratio, deforestation, forest fires and inbreeding depression strongly influence the population survival. Long-term studies and biological information on species inhabiting areas subjected to anthropogenic impacts, like the Atlantic Rainforest, are vital to conservation actions, since they make it possible population viability analyses and the design of management strategies to threatened populations.

Turtles have long life spans, overlapping generations and promiscuous mating systems. Thus, they are an ideal system with which to investigate the application of conservation genetics methods and assumptions to long-lived organisms. Turtles are also one of the most threatened groups of vertebrates and conservation genetics studies are essential to effective recovery of turtle species. This thesis has two main objectives: 1) to evaluate some common population genetics assumptions with respect to turtles and other long-lived organisms, and 2) to collect important information on the population genetics of threatened turtles in Ontario, which can be used to inform species recovery. In Chapters Two and Three, I describe the development of novel microsatellite markers for the snapping turtle and spiny softshell. In Chapter Four I demonstrate significant genetic structure in populations of the endangered spotted turtle in Ontario, and find that “bottleneck tests” may fail to detect recent population declines in small turtle populations. I also show that spotted turtles do not show the typical correlation between population size and genetic diversity. In Chapter Five I use microsatellite markers developed in Chapter Two and document population structure in the widespread snapping turtle for the first time. I compare these results with results from Chapter Four to test the traditionally accepted hypothesis that genetic diversity is reduced in small, isolated populations compared to large, connected populations. As in Chapter Four, my results suggest that the usual patterns of genetic structure and loss of diversity may not apply to turtles. In Chapter Six I conduct a conservation genetics study of the endangered Blanding’s turtle. Finally, in Chapter Seven I combine results from spotted, snapping and Blanding’s turtles to test whether vagility predicts population structure, genetic diversity and significant barriers to gene flow in three species sampled across a single landscape. Analyses reveal minimal congruence in barriers to gene flow and the three species show unexpected and contrasting patterns of diversity across the landscape. Discordant patterns among species highlight areas for further research and shed light on possible cryptic behaviour, and I discuss potential further directions for research in the Summary.

Wytamma Wirth studied ranaviral infection in Australian freshwater turtles. He found several factors that influence ranaviral disease in Australian freshwater turtles and examined the distribution of these viruses in North Queensland.

Roads are long linear features on the landscape that impact wildlife and their habitats. Among all forms of wildlife turtles are one of the most negatively affected by roads. Wildlife biologists and civil engineers have developed and implemented road design measures to mitigate the negative effects associated with roads. One common approach used to reduce road mortality and to facilitate movement of turtles is to construct a road mitigation system. There are currently 28 road mitigation systems for wildlife in Massachusetts, of which 14 were specifically built for turtles. We identified all known systems in Massachusetts and collected site and structural design information for each. In addition, we also examined the relative effectiveness of experimental passages for freshwater turtles. Structures were evaluated with respect to how their height, width, and position (at or below-grade), influenced the movements of painted turtles. A total of 190 turtles were exposed to the experiential trials and their behavior was characterized by 3 response variables (Total time to complete the trial, Total hesitations observed, and Success based on no hesitations and completion of the trial in less than 120 minutes). We concluded that painted turtles exposed to below-grade tunnels were less hesitant and traveled faster through them as the tunnel size increased from 0.6 m x 0.6 m to 1.2 m x 1.2 m. The 1.2 m x 1.2 m tunnel size overall proved to be the size with the fewest hesitations observed, fastest total times, and highest success rate.

Roadways are a pervasive feature of northeastern landscapes and can be a significant source of mortality for turtles. Until recently, little has been known about the design requirements for successful under-road passages for turtles and other wildlife to move safely between bisected habitat patches. At outdoor laboratories, using a factorial experimental design, we examined movements in response to varying light levels, and barrier opacity for painted turtles (Chrysemys picta, n=833), Blanding’s turtles (Emydoidea blandingii, n=49), and spotted turtles (Clemmys guttata, n=49). Additionally, we examined tunnel size, tunnel entrance design, and artificial lighting for painted turtles only. All three species responded poorly to a 0% available light treatment. As the amount of natural light transmitted through the tops of tunnels increased, successful completion of the trials increased. Furthermore, turtles generally moved at a slower rate when traveling along a translucent barrier, compared to an opaque one. Our results indicate the importance of designing road passage structures for freshwater turtles that provide adequate tunnel lighting in combination with specific entrance designs that meet the goals of the project.

Connectivity is currently a central issue in landscape management and planning for the conservation of wildlife species occupying sparse habitat patches. The European pond turtle (Emys orbicularis L., 1758) is an endangered species that has important populations in several pond systems in the SW Alentejo region of Portugal. About 56% of the ponds of this region have been destroyed from 1991 to 2009, primarily as a consequence of the agricultural intensification. This destruction possibly has severe consequences for the European pond turtle, namely in the interpond connectivity of the overall pond system. A population of E. orbicularis living in a pond system in the surroundings of a small locality (Longueira‐Almograve) has been regularly surveyed since 2003. A total of 595 captures, totalling 205 individuals, were performed during this period. Data on interpond movements was used to estimate movement probabilities among pairs of ponds separately for females, males and juveniles. The relationship between these movement probabilities and three measures of interpond distances – linear, least cost and accumulated cost distance – were tested through Mantel tests. Movement probabilities tended to be more correlated with accumulated cost distance estimates, although a significant relationship with linear distances was also found. Females had a higher proportion of successive movements between different ponds than males. The variation of the movement between pairs of ponds with distance was modelled by fitting a negative exponential function. This function was used to predict the interpond movement probabilities among all the ponds of the system, including non‐sampled or undersampled ponds. Based on these probabilities, defining several alternative threshold probabilities to settle different degrees of connectivity among ponds, several graph‐based attributes were then estimated in order to rank the ponds in terms of their importance to the overall functional connectivity for the European pond turtle in this landscape; Ecologia espacial de uma tartaruga-de-água-doce num complexo de charcos temporários Resumo: A conetividade é uma propriedade central na gestão e planeamento da paisagem para a conservação de espécies que ocupam parcelas de habitat dispersas. O cágado‐de‐carapaça‐estriada (Emys orbicularis L., 1758) é uma espécie de tartaruga de água doce considerada em perigo em Portugal, que mantém importantes populações em diversos sistemas de charcos na Costa Sudoeste Alentejana. Cerca de 56% dos charcos desta região foram destruídos entre 1991 e 2009 em consequência da intensificação agrícola no perímetro de rega do Rio Mira. Esta destruição teve certamente importantes consequências para o cágado‐de‐carapaça‐estriada, nomeadamente ao nível da perda de conetividade entre charcos de todo o sistema da região. Uma população de E. orbicularis que ocorre num sistema de charcos nas imediações de uma pequena localidade (Longueira‐Almograve) tem sido regularmente amostrada desde 2003. Um total de 595 capturas, totalizando 205 indivíduos, foram realizadas durante este período. Os dados de movimentos entre lagoas foram usados para estimar a probabilidade de movimentos entre lagoas, separadamente para fêmeas, machos e juvenis. A relação entre essas probabilidades e três medidas de distância entre lagoas – distância linear, de menor custo e de custo acumulado – foi testada através de testes de Mantel. As probabilidades de movimento tenderam a apresentar uma maior correlação com a distância de custo acumulado, apesar de se ter encontrado uma relação significativa também com a distância linear. A proporção de movimentos sucessivos entre lagoas diferentes foi significativamente superior nas fêmeas. A variação da probabilidade de movimentos com a distância foi modelada através do ajustamento de uma função exponencial negativa. Esta função foi utilizada para prever a probabilidade de movimentos entre todos os charcos do sistema, incluindo charcos não amostrados ou subamostrados. Diversos atributos baseados em grafos espaciais foram estimados, definindo diferentes probabilidades críticas para estabelecer diferentes graus de conetividade entre charcos. Esta metodologia permitiu ordenar a importância dos charcos em termos da sua contribuição para a conetividade funcional global do sistema para E. orbicularis.

La biologie de la conservation est un domaine de recherche en pleine expansion en raison de la perte accélérée de la biodiversité à l’échelle mondiale. Pour mieux comprendre les processus et les menaces au maintien des populations de petite taille et les effets des facteurs anthropiques sur la biodiversité, la génétique est fréquemment utilisée en conservation. Des analyses génétiques peuvent, par exemple, nous informer sur les tendances à long terme, la diversité des populations et les stratégies de reproduction d’une espèce. La tortue des bois (Glyptemys insculpta) est une espèce endémique à l’Amérique du Nord qui est en danger d’extinction selon l’Union internationale pour la conservation de la nature. Dans le cadre de ma thèse, j’avais comme objectif de caractériser la diversité génétique de cette espèce menacée au Canada. À cet effet, j’ai analysé la génétique des populations de tortues des bois à plusieurs échelles spatiales et temporelles, afin de mieux cerner les processus ayant un impact sur la diversité des populations. Dans un premier temps, les relations de parentalité ont été reconstruites au sein d’une population de tortues des bois pour estimer la fréquence de paternité multiple et de paternité répétée. Les résultats de mes travaux suggèrent que l’emmagasinement de sperme chez la femelle et la reproduction multiple avec les mêmes partenaires pour plus d’une saison de reproduction pourraient expliquer ces phénomènes. Ces stratégies de reproduction pourraient dans ce cas être induites par la faible densité de la population à l’étude, ou encore par la fidélité au site d’hibernation où la majorité des évènements de copulation ont lieu. Par la suite, je me suis intéressée à la diversité génétique des populations de tortues de bois. J’ai voulu comprendre les effets de la configuration spatiale des éléments du paysage et les évènements de dispersion géographique sur la diversité des populations. À l’aide d’une approche de génétique du paysage, mes analyses montrent que la division des populations par bassins versants explique une large fraction de la diversité génétique interpopulations. Ces résultats confirment également que les bassins versants représentent des unités de gestion propices à la protection des populations de tortues des bois. Finalement, des analyses de réseaux ont été utilisées pour mieux cerner la dynamique de flux génique entre les populations de la rive nord et de la rive sud du fleuve Saint-Laurent. Plus spécifiquement, la rive nord se caractérise par un réseau robuste de populations isolées, alors que les populations de la rive sud présentent plutôt une structure de métapopulation. En utilisant les réseaux construits à partir de données génétiques, des scénarios hypothétiques furent comparés pour explorer la sélection de populations à l’aide du logiciel BRIDES. Les résultats de ces analyses ont permis de cibler l’importance de certaines populations de tortues des bois pour la connectivité du réseau. L’importance de ces populations n’aurait pu être prédite par les résultats de la diversité et de la différenciation génétique, les indices de centralité et les analyses d’élimination de nœuds. Grâce à la génétique, cette thèse apporte de nouvelles connaissances sur la tortue des bois, les stratégies de reproduction des différents sexes, le flux génique, la connectivité et l’influence du réseau hydrographique sur la diversité des populations. Ces résultats nous permettent d’avoir une meilleure compréhension des processus affectant la diversité génétique de cette espèce afin de mieux la protéger. Toutes les analyses réalisées pour cette thèse sont directement applicables à l’ensemble des autres espèces longévives avec des générations chevauchantes.
Conservation biology is a rapidly expanding field of research due to the accelerating loss of global biodiversity. To better understand the processes and threats to the persistence of small populations and the effects of anthropogenic factors on biodiversity, genetic approaches are frequently used in conservation. Genetic analyzes can, for example, inform us about long-term trends, population diversity and reproductive strategies of a species. The wood turtle (Glyptemys insculpta) is a species endemic to North America that is endangered according to the International Union for the Conservation of Nature. As part of my thesis, my objective was to characterize the genetic diversity of this threatened species in Canada. In order to better understand the impact of reproductive strategy and landscape structure on population diversity, I analyzed the genetics of wood turtle populations at several spatial and temporal scales. First, parentage relationships were reconstructed in a population of wood turtles to estimate the frequency of multiple and repeated paternity. The results of my work suggest that sperm storage in females and multiple reproduction with the same partners for more than one breeding season could explain these phenomena. These reproduction strategies could in this case be induced by the low density of the study population, or by fidelity to the overwintering site where the majority of copulation events take place. Subsequently, I assessed the genetic diversity of wood turtle populations. I wanted to understand the effects of the landscape configuration and geographic dispersion events on the diversity of populations. Using a landscape genetics approach, my analyzes show that the division of populations by watershed explains a large fraction of the genetic diversity between populations. These results also confirm that watersheds represent management units conducive to the protection of wood turtle populations. Finally, network analysis was used to better understand the gene flow dynamics among populations located on the north and south shores of the St. Lawrence River. More specifically, the north shore is characterized by a robust network of isolated populations, whereas the populations on the south shore present more of a metapopulation structure. Using population graphs, hypothetical scenarios were compared to explore the node selection process using the BRIDES algorithm. The results of these analyzes made it possible to point out specific populations of wood turtles, considering their importance for network connectivity. This could have not been predicted by using genetic diversity and distinctiveness estimates, node-based metrics, and node removal analysis for these populations. Thanks to genetics, this thesis brings new knowledge on the wood turtle, the reproductive strategies of both sexes, the gene flow, the connectivity and the influence of the hydrographic network on population diversity. These results allow us to have a better understanding of the processes affecting the genetic diversity of this species in order to better protect it. All analyses performed for this thesis are directly applicable to other long-lived species with overlapping generations.

碩士
國立屏東科技大學
野生動物保育研究所
93
Ocadia sinensis is the most common turtle in Taiwan. According to the field research, it was omnivorous. The diet habit, size and growing rate were different between sexes. With the increasing of the carapace, females ingested more plants. And as they grew, the growing rate of females was higher than that of males, the size of females would be larger than that of males significantly, too. The purpose of this study is to experiment the effect of different diet compositions to O. sinensis by captivity raised. We want to know if the digestive perform will affect their diet habit and growth. At first, we kept turtles in pool individually, fed them three kinds of diets: plant, animal, both plant and animal (mix), respectively, and then collected their feces. We counted the digestibility of crude fiber and total nitrogen of these turtles. The digestibility of crude fiber and total nitrogen of males had no significant differences among the three diets, but there was significant difference between that for females. The digestibility of crude fiber and total nitrogen had no significant difference between males and females.

Anoxia is an environmental stress that few air-breathing vertebrates can tolerate for more than a few minutes before extensive neurodegeneration occurs. Some facultative anaerobes, including the freshwater western painted turtle Chrysemys picta bellii, are able to coordinately reduce ATP demand to match reduced ATP availability during anoxia, and thus tolerate prolonged insults without apparent detriment. To reduce metabolic rate, turtle neurons undergo channel arrest and spike arrest to decrease membrane ion permeability and neuronal electrical excitability, respectively. However, although these adaptations have been documented in turtle brain, the mechanisms underlying channel and spike arrest are poorly understood. The aim of my research was to elucidate the cellular mechanisms that underlie channel and spike arrest and the neuroprotection they confer on the anoxic turtle brain. Using electrophysiological and fluorescent imaging techniques, I demonstrate for the first time that: 1) the α-amino-3-hydroxy-5-methyl-4-isoxazole propionic acid receptor (AMPAR) undergoes anoxia-mediated channel arrest; 2) delta opioid receptors (DORs), and 3) mild mitochondrial uncoupling via mitochondrial ATP-sensitive K+ channels result in an increase in cytosolic calcium concentration and subsequent channel arrest of the N-methyl-D-aspartate receptor, preventing excitotoxic calcium entry, and 4) reducing nitric oxide (NO) production; 5) the cellular concentration of reactive oxygen species (ROS) decreases with anoxia and ROS bursts do not occur during reoxygenation; and 6) spike arrest occurs in the anoxic turtle cortex, and that this is regulated by increased neuronal conductance to chloride and potassium ions due to activation of γ–amino-butyric acid receptors (GABAA and GABAB respectively), which create an inhibitory electrical shunt to dampen neuronal excitation during anoxia. These mechanisms are individually critical since blockade of DORs or GABA receptors induce excitotoxic cell death in anoxic turtle neurons. Together, spike and channel arrest significantly reduce neuronal excitability and individually provide key contributions to the turtle’s long-term neuronal survival during anoxia. Since the turtle is the most anoxia-tolerant air-breathing vertebrate identified, these results suggest that multiple mechanisms of metabolic suppression acting in concert are essential to maximizing anoxia-tolerance.