Heritability of corticosterone response and changes in life history traits during selection in the zebra finch

Vertebrates respond to environmental stressors through the neuro-endocrine stress response, which involves the production of glucocorticoids. We have selected independent, duplicate divergent lines of zebra finches for high, low and control corticosterone responses to a mild stressor. This experiment has shown that over the first four generations, the high lines have demonstrated a significant realized heritability of about 20%. However, the low lines have apparently not changed significantly from controls. This asymmetry in response is potentially because of the fact that all birds appear to be showing increased adaptation to the environment in which they are housed, with significant declines in corticosterone response in control lines as well as low lines. Despite the existence of two- to threefold difference in mean corticosterone titre between high and low lines, there were no observed differences in testosterone titre in adult male birds from the different groups. In addition, there were no consistent, significant differences between the lines in any of the life history variables measured – number of eggs laid per clutch, number of clutches or broods produced per pair, number of fledglings produced per breeding attempt, nor in any of egg, nestling and fledgling mortality. These results highlight the fact that the mechanisms that underlie variation in the avian physiological system can be modified to respond to differences between environments through selection. This adds an additional level of flexibility to the avian physiological system, which will allow it to respond to environmental circumstances.

Ecology of epifaunal caridean shrimps in the Hopkins River estuary, and the role of estuaries in the life history of the atyid "Paratya Australiensis" Kemp, 1917 in south-eastern Australia

This study examined the factors affecting the distribution and abundance of epifaunal caridean shrimps in seagrass meadows of the Hopkins River estuary in south-western Victoria, Australia, and investigated the life history patterns of the freshwater Parana australiensis, found for the first time in estuaries. Adult and sub-adult shrimps were surveyed in seagrass meadows along the estuary over two years, and their planktonic larvae were surveyed in adjacent waters. Three species were collected. The marine Palaemon serenus occurred only near the mouth, summer to autumn, in high salinities. The marine/estuarine Macrobrachium intermedium occurred throughout the estuary. Adults were most abundant in late autumn, and least abundant in summer (unlike trends reported in marine meadows). Densities were higher and less variable in downstream meadows. P. australiensis occurred in the upper estuary all year, most abundantly in spring, due to migration from the river after peak discharge. Ovigerous females dominated, while males, showing less migration into the estuary, dominated above estuarine influence. Adults disappeared from the estuary in summer as salinity rose. Breeding period for P. australiensis was briefer in the estuary (September-December) than upstream (July-April). M. intermedium began breeding later in the upper estuary (November/December-March) than in the lower estuary (October-March), probably reflecting a physiological response to lower salinity, rather than an interaction with P. australiensis. No ovigerous P. serenus were found in the estuary. Larvae of P. australiensis and M intermedium occurred abundantly throughout the estuary, but P. serenus larvae did not. P. australiensis was an early coloniser to the plankton after peak discharge (November-December). Larvae concentrated in the deep saline layer at the head of the intruding salt wedge, thus probably maintaining longitudinal position. Diurnal vertical migrations were evident within the salt wedge, and in a deep pool above tidal influence. M. intermedium larvae occurred October-May in the lower estuary and November-April in the upper estuary, peaking in abundance one to two months after P. australiensis. They were associated with low surface flows and surface salinities greater than 10, over an anoxic deeper layer. All three species exhibited extended development of euryhaline larvae in the laboratory. Tolerances and optimal salinities of larvae of the three species reflected their distributions. M. intermedium was the most euryhaline species. P. australiensis larvae were tolerant of higher salinities than juveniles of adults: capable of developing in salinity of at least 15. Most P. australiensis juveniles recruited to the estuary November-December, after which numbers declined dramatically. After settlement, most recruits probably migrated upstream out of the estuary. Two cohorts of M. intermedium recruited to the estuary from larvae in summer (December and February), but some juveniles also migrated from adjacent coastal waters. Post-larval migration was at least as important a determinant of abundance as direct recruitment from estuarine, planktonic larvae in all three species. Distributions among seagrass meadows along the estuary were determined primarily by physico-chemical patterns driven by hydrological changes. Seasonal variations in salinity and temperature were strongly associated with seasonal variations in shrimp abundance. Salinity tolerances of adults of the three species reflected their distribution patterns. Biotic interactions were more important in determining distributions within meadows. P. australiensis, when abundant, were associated with seagrass biomass. M. intermedium were also, but when seagrass was sparsest and least extensive. The two species apparently partitioned the seagrass meadow according to depth in early summer. Laboratory experiments suggested P. australiensis was displaced from deeper water by M. intermedium. Preference for vegetative complexity and competition for position within meadows suggest the underlying importance of predation in regulating shrimp populations. A survey of south-eastern Australian estuaries found P. australiensis larvae abundant in all stable, open, well-developed, salt-wedge estuaries where adults were abundant. Adults were most abundant in low salinities among submerged leafy macrophytes. Reproductive traits of P. australiensis were compared in estuarine and fresh reaches of three rivers. Early in the breeding season, egg size was smaller, and (size-specific) egg number larger in estuaries than upstream. A trade-off between egg size and egg number resulted in no difference in total (size-specific) reproductive investment between locations. Reproductive investment tended to decrease at some locations over the breeding season, and this decrease was a result of decreased egg size in most cases. The decrease in reproductive investment probably reflected reduced food availability for the adult, while the reduced egg size was probably a response to improved conditions for larval development. In the Hopkins River, larger egg size at upstream sites was reflected in larger early stage larvae. Later stage larvae were larger in the estuary, suggesting more favourable conditions for larval development. Allozyme electrophoresis showed the P. australiensis populations in each of the three rivers to be distinct. Allozyme frequencies were not different within the Hopkins River, but upstream and estuarine locations in the Curdies and Gellibrand were different. Although some variation in reproductive traits within catchments may have been due to genotypic differences, trade-offs between egg size and number, and decreases in egg size over summer were probably due to plastic responses to environmental cues. It is proposed P. australiensis inhabits and reproduces in both estuarine and freshwater environments by plastic response to environmental conditions. Recruitment to estuaries is dependent on the presence of suitable adult, littoral habitat, and a stable salt wedge for larval retention. Estuaries are important recruitment sites for P. australiensis, potentially allowing an extra brood each year before riverine recruitment. Estuarine broods could constitute a large part of the total fecundity of P. australiensis females. Euryhaline larvae and estuarine recruitment of P. australiensis suggest marine transport of larvae between estuaries as a possible dispersal mechanism for Paratya species.

Is life history a barrier to dispersal? Contrasting patterns of genetic differentiation along an oceanographically complex coast

Extreme variation in early life-history strategies is considered a moderately good predictor of genetic subdivision and hence dispersal for a range of marine species. In reality, however, a good deal of population differentiation must reflect historical effects, more subtle variation in life histories, and, particularly, the interaction of larvae with oceanographic processes. Using a combination of allozyme and microsatellite markers, we show that the large-scale genetic structure of populations of three species (direct and planktonically developing cushion stars and a planktonic developing sea anemone that is also asexually viviparous) varies consistently, in line with the predicted capacity for dispersal within three geographic regions. We detected high levels of genetic subdivision for the direct developing cushion star (FST = 0.6), low levels for the planktonically developing cushion star (FST = 0.009), and intermediate levels for the sexual/asexual sea anmone (FST = 0.19). These patterns are exhibited despite the highly variable patterns of current movement and the presence of biogeographic barriers. Our results suggest that, although there is large scale genetic differentiation for two species, patterns of population connectivity are remarkably consistent within major regions and do not reflect variation in major oceanographic processes or genetic discontinuity coincident with biogeographic boundaries.

Ecological and life-history factors influencing the evolution of maternal antibody allocation : a phylogenetic comparison

Maternally derived yolk antibodies provide neonates with immune protection in early life at negligible cost to mothers. However, developmental effects on the neonate's future immunity are potentially costly and thus could limit yolk antibody deposition. The benefits to neonatal immunity must be balanced against costs, which may depend on neonate vulnerability to pathogens, developmental trajectories and the immunological strategies best suited to a species' pace of life. We measured yolk antibodies and life-history features of 23 species of small Neotropical birds and assessed the evidence for each of several hypotheses for life history and ecological effects on the evolution of yolk antibody levels. Developmental period and yolk antibodies are negatively related, which possibly reflect the importance of humoral immune priming through antigen exposure, and selection to avoid autoimmunity, in species with a slower pace of life. There is also a strong relationship between body size and yolk antibody concentration, suggesting that larger species are architecturally equipped to produce and transfer higher concentrations of antibodies. These results suggest that developmental effects of maternally derived antibodies, such as imprinting effects on B-cell diversity or autoimmune effects, are important and deserve more consideration in future research.

A life history model for the giant crab Pseudocarcinus gigas

This study examined P. gigas populations from Cape Naturaliste (Western Australia) to Flinders Island (Tasmania), investigated reproduction, developed a unique tag to study their movement and growth, and collected new environmental information that allowed the development of a model of the life history for this iconic endemic Australian species.

Aspects of the life history of skates from southeastern Australia

The age and growth, reproduction and diet of several skate species in south-east Australia were studied. From studying these life history characteristics it was determined that skates have a relatively low productivity and may therefore be at risk of local extinction if increasingly exploited without appropriate management.

Evolutionary consequences of life history variation in galaxiid fish

This thesis examined genetic variation among populations of galaxiid fish of different life histories. Evidence of incipient speciation was found among several landlocked populations; however, little genetic differentiation was found among Australian coastal populations. Evidence was found to suggest a marine ancestry for galaxiids and several taxonomic revisions were suggested.

Population structure and life history characteristics of euastacus bispinosus and cherax destructor (Parastacidae) in the Grampians National Park, Australia

Despite being the most widely distributed and cultured freshwater crayfish species in Australia, relatively little is known about the ecology of Cherax destructor outside of captivity. Similarly, few ecological studies have been conducted on the large and threatened stream dwelling Australian freshwater crayfish Euastacus bispinosus. A series of seasonal sampling surveys over two years investigated the population structure, life history and reproductive timing of C. destructor in fire dam habitats, and of E. bispinosus in channel (stream) habitats, in the Grampians National Park in south-west Victoria, Australia. Cherax destructor individuals in the largest size class (50 – 59.95 mm OCL) were not abundant during the study, while those belonging to the 0 – 9.95 mm OCL size class were more frequent in summer than at other times of year, suggesting synchronous recruitment in fire dam habitats. Individuals in the reproductively active mid-size classes were also frequent in summer. For E. bispinosus, sex ratios in spring always favoured females, although there were no clear trends for other times of year. Gravid E. bispinosus females were found in winter and spring throughout the study, and were sexually mature at a smaller size than has previously been reported.

Comparison of life-history characteristics of island and mainland populations of the swamp antechinus

The reproductive ecology of the swamp antechinus Antechinus minimus, a small dasyurid (Dasyuridae) marsupial with obligate male semelparity, was investigated in populations inhabiting the mainland coast and on a nearby offshore island in south-eastern Australia. The size and sex ratios of litters, individual body mass and size, timing of births and female longevity were determined from live-trapped animals. The island population had significantly smaller litter sizes and greater adult body mass in comparison with the mainland population. This is consistent with features of the ‘island syndrome’, which predicts directional selection for these traits in high-density populations with reduced extrinsic mortality. However, inter-annual variability in litter sizes in the island population suggests that litter size is more responsive to fluctuating local conditions, such as population density, which is likely to affect food availability, rather than directional genetic changes. In contrast with other antechinus species, biased sex ratios were not evident. In addition, large variations of the timing of births were estimated at both sites and these appear to be related to seasonal conditions such as autumn rainfall and female body mass before mating.

Identifying crucial gaps in our knowledge of the life-history of avian influenza viruses - An Australian perspective

We review our current knowledge of the epidemiology and ecology of avian influenza viruses (AIVs) in Australia in relation to the ecology of their hosts. Understanding the transmission and maintenance of low-pathogenic avian influenza (LPAI) viruses deserves scientific scrutiny because some of these may evolve to a high-pathogenic AIV (HPAI) phenotype. That the HPAI H5N1 has not been detected in Australia is thought to be a result of the low level of migratory connectivity between Asia and Australia. Some AIV strains are endemic to Australia, with Australian birds acting as a reservoir for these viruses. However, given the phylogenetic relationships between Australian and Eurasian strains, both avian migrants and resident birds within the continent must play a role in the ecology and epidemiology of AIVs in Australia. The extent to which individual variation in susceptibility to infection, previous infections, and behavioural changes in response to infection determine AIV epidemiology is little understood. Prevalence of AIVs among Australian avifauna is apparently low but, given their specific ecology and Australian conditions, prevalence may be higher in little-researched species and under specific environmental conditions.

Covariation in life-history traits : differential effects of diet on condition, hormones, behavior, and reproduction in genetic finch morphs

The relative contribution of genetic and environmental factors in determining variation in life-history traits is of central interest to evolutionary biologists, but the physiological mechanisms underlying these traits are still poorly understood. Here we experimentally demonstrate opposing effects of nutritional stress on immune function, endocrine physiology, parental care, and reproduction between red and black head-color morphs of the Gouldian finch (Erythrura gouldiae). Although the body condition of black morphs was largely unaffected by diet manipulation, red birds were highly sensitive to dietary changes, exhibiting considerable within-individual changes in condition and immune function. Consequently, nutritionally stressed red birds delayed breeding, produced smaller broods, and reared fewer and lower-quality foster offspring than black morphs. Differences in offspring quality were largely due to morph-specific differences in parental effort: red morphs reduced parental provisioning, whereas black morphs adaptively elevated their provisioning effort to meet the increased nutritional demands of their foster brood. Nutritionally stressed genetic morphs also exhibited divergent glucocorticoid responses. Black morphs showed reduced corticosterone-binding globulin (CBG) concentrations and increased levels of free corticosterone, whereas red morphs exhibited reduced free corticosterone levels and elevated CBG concentrations. These opposing glucocorticoid responses highlight intrinsic differences in endocrine sensitivities and plasticity between genetic morphs, which may underlie the morph-specific differences in condition, behavior, and reproduction and thus ultimately contribute to the evolution and maintenance of color polymorphism.

Are animal personality traits linked to life-history productivity?

Animal personality traits such as boldness, activity and aggressiveness have been described for many animal species. However, why some individuals are consistently bolder or more active than others, for example, is currently obscure. Given that life-history tradeoffs are common and known to promote inter-individual differences in behavior, we suggest that consistent individual differences in animal personality traits can be favored when those traits contribute to consistent individual differences in productivity (growth and/or fecundity). A survey of empirical studies indicates that boldness, activity and/or aggressiveness are positively related to food intake rates, productivity and other life-history traits in a wide range of taxa. Our conceptual framework sets the stage for a closer look at relationships between personality traits and life-history traits in animals.

Spatial and temporal drivers of small mammal distributions in a semi-arid environment : the role of rainfall, vegetation and life-history

A key task in ecology is to understand the drivers of animal distributions. In arid and semi-arid environments, this is challenging because animal populations show considerable spatial and temporal variation. An effective approach in such systems is to examine both broad-scale and long-term data. We used this approach to investigate the distribution of small mammal species in semi-arid ‘mallee’ vegetation in south-eastern Australia. First, we examined broad-scale data collected at 280 sites across the Murray Mallee region. We used generalized additive mixed models (GAMMs) to examine four hypotheses concerning factors that influence the distribution of individual mammal species at this scale: vegetation structure, floristic diversity, topography and recent rainfall. Second, we used long-term data from a single conservation reserve (surveyed from 1997 to 2012) to examine small mammal responses to rainfall over a period spanning a broad range of climatic conditions, including record high rainfall in 2011. Small mammal distributions were strongly associated with vegetation structure and rainfall patterns, but the relative importance of these drivers was species-specific. The distribution of the mallee ningaui Ningaui yvonneae, for example, was largely determined by the cover of hummock grass; whereas the occurrence of the western pygmy possum Cercartetus concinnus was most strongly associated with above-average rainfall. Further, the combination of both broad-scale and long-term data provided valuable insights. Bolam's mouse Pseudomys bolami was uncommon during the broad-scale survey, but long-term surveys showed that it responds positively to above-average rainfall. Conceptual models developed for small mammals in temperate and central arid Australia, respectively, were not, on their own, adequate to account for the distributional patterns of species in this semi-arid ecosystem. Species-specific variation in the relative importance of different drivers was more effectively explained by qualitative differences in life-history attributes among species.