Typescript (photocopy) / viii, 193 leaves, [5] leaves of plates : ill. ; 30 cm. / Title page, contents and abstract only. The complete thesis in print form is available from the University Library. / Thesis (Ph.D.)--University of Adelaide, Dept. of Zoology, 1984

Thesis (Ph.D.)--University of Adelaide, Dept. of Zoology, 1984. / Typescript (photocopy).

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.

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.

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.