Influence of environmental conditions on foraging behaviour and its consequences on reproductive performance in little penguins

During the breeding season, seabirds are central place foragers and have to adapt their foraging behaviour in response to environmental variation to maximize efficiency and reproductive output. Due to its small size and swimming mode of transport, the little penguin (Eudyptula minor) is expected to be greatly susceptible to such fluctuations. The links between local-, meso- and macro-scale environmental conditions and inter-annual variation in foraging behaviour and reproductive performance of little penguins were investigated during three consecutive breeding seasons at two colonies in south-eastern Australia marked by contrasting oceanographic conditions. At a local scale, foraging effort was correlated positively with wind direction and negatively with wave height. At a regional scale, foraging effort of individuals from both colonies was negatively correlated with higher sea surface temperature (SST) off the Bonney Coast in the previous Austral summer, suggesting a weaker Bonney Upwelling event and a cascade of effects throughout the Bass Strait region. At a larger scale, the El Niño Southern Oscillation was also found to correlate with foraging behaviour, with lower foraging effort being observed during La Niña event. Although individuals increased their foraging effort during years with poorer conditions, they were not able to maintain high breeding success. In addition, peak egg-laying was found to coincide with a decrease in local SST and a peak of sea surface chlorophyll-a concentration. In conclusion, these results highlight how different environmental conditions could influence foraging behaviour and ultimately reproductive success of little penguins. It also showed that under certain circumstances, these individual strategies were not sufficient to cope with environmental variability.

The bright incubate at night: sexual dichromatism and adaptive incubation division in an open-nesting shorebird

Ornamentation of parents poses a high risk for offspring because it reduces cryptic nest defence. Over a century ago, Wallace proposed that sexual dichromatism enhances crypsis of open-nesting females although subsequent studies found that dichromatism per se is not necessarily adaptive. We tested whether reduced female ornamentation in a sexually dichromatic species reduces the risk of clutch depredation and leads to adaptive parental roles in the red-capped plover Charadrius ruficapillus, a species with biparental incubation. Males had significantly brighter and redder head coloration than females. During daytime, when visually foraging predators are active, colour-matched model males incurred a higher risk of clutch depredation than females, whereas at night there was no difference in depredation risk between sexes. In turn, red-capped plovers maintained a strongly diurnal/nocturnal division of parental care during incubation, with males attending the nest largely at night when visual predators were inactive and females incubating during the day. We found support for Wallace's conclusion that reduced female ornamentation provides a selective advantage when reproductive success is threatened by visually foraging predators. We conclude that predators may alter their prey's parental care patterns and therefore may affect parental cooperation during care.

Foraging strategies and reproductive success in two sympatric seabird species: influence of environmental variability

 This work focused on flexibility in foraging strategies in little penguins and short-tailed shearwaters, which represents an important mechanism enabling them to cope with highly dynamic ecosystems and provides a buffer against spatial and temporal changes in prey availability over the breeding period.

The ecology and evolution of beak and feather disease virus in Platycercus elegans

 This thesis explores the complex ecological and evolutionary interactions between beak and feather disease virus and one of its hosts the crimson rosella. The work identifies several factors that predict viral infection in wild birds and determines how host population structure influences viral evolution.

Artificial selection for food colour preferences

Colour is an important factor in food detection and acquisition by animals using visually based foraging. Colour can be used to identify the suitability of a food source or improve the efficiency of food detection, and can even be linked to mate choice. Food colour preferences are known to exist, but whether these preferences are heritable and how these preferences evolve is unknown. Using the freshwater fish Poecilia reticulata, we artificially selected for chase behaviour towards two different-coloured moving stimuli: red and blue spots. A response to selection was only seen for chase behaviours towards the red, with realized heritabilities ranging from 0.25 to 0.30. Despite intense selection, no significant chase response was recorded for the blue-selected lines. This lack of response may be due to the motion-detection mechanism in the guppy visual system and may have novel implications for the evolvability of responses to colour-related signals. The behavioural response to several colours after five generations of selection suggests that the colour opponency system of the fish may regulate the response to selection.

White sharks exploit the sun during predatory approaches

There is no conclusive evidence of any nonhuman animal using the sun as part of its predation strategy. Here, we show that the world's largest predatory fish-the white shark (Carcharodon carcharias)-exploits the sun when approaching baits by positioning the sun directly behind them. On sunny days, sharks reversed their direction of approach along an east-west axis from morning to afternoon but had uniformly distributed approach directions during overcast conditions. These results show that white sharks have sufficient behavioral flexibility to exploit fluctuating environmental features when predating. This sun-tracking predation strategy has a number of potential functional roles, including improvement of prey detection, avoidance of retinal overstimulation, and predator concealment.

Writing scientific papers, with special reference to evolutionary ecology

© 2015, Springer International Publishing Switzerland. The advancement of science, as well as scientific careers, depends upon good and clear scientific writing. Science is the most democratic of human endeavours because, in principle, anyone can replicate a scientific discovery. In order for this to continue, writing must be clear enough to be understood well enough to allow replication, either in principle or in fact. In this paper I will present data on the publication process in Evolutionary Ecology, use it to illustrate some of the problems in scientific papers, make some general remarks about writing scientific papers, summarise two new paper categories in the journal which will fill gaps that appear to be expanding in the literature, and summarise new journal policies to help mitigate existing problems. Most of the suggestions about writing would apply to any scientific journal.

Invasive carnivores alter ecological function and enhance complementarity in scavenger assemblages on ocean beaches

Species composition is expected to alter ecological function in assemblages if species traits differ strongly. Such effects are often large and persistent for nonnative carnivores invading islands. Alternatively, high similarity in traits within assemblages creates a degree of functional redundancy in ecosystems. Here we tested whether species turnover results in functional ecological equivalence or complementarity, and whether invasive carnivores on islands significantly alter such ecological function. The model system consisted of vertebrate scavengers (dominated by raptors) foraging on animal carcasses on ocean beaches on two Australian islands, one with and one without invasive red foxes (Vulpes vulpes). Partitioning of scavenging events among species, carcass removal rates, and detection speeds were quantified using camera traps baited with fish carcasses at the dune–beach interface. Complete segregation of temporal foraging niches between mammals (nocturnal) and birds (diurnal) reflects complementarity in carrion utilization. Conversely, functional redundancy exists within the bird guild where several species of raptors dominate carrion removal in a broadly similar way. As predicted, effects of red foxes were large. They substantially changed the nature and rate of the scavenging process in the system: (1) foxes consumed over half (55%) of all carrion available at night, compared with negligible mammalian foraging at night on the fox-free island, and (2) significant shifts in the composition of the scavenger assemblages consuming beach-cast carrion are the consequence of fox invasion at one island. Arguably, in the absence of other mammalian apex predators, the addition of red foxes creates a new dimension of functional complementarity in beach food webs. However, this functional complementarity added by foxes is neither benign nor neutral, as marine carrion subsidies to coastal red fox populations are likely to facilitate their persistence as exotic carnivores.

Combined use of GPS and accelerometry reveals fine scale three-dimensional foraging behaviour in the short-tailed shearwater

Determining the foraging behaviour of free-ranging marine animals is fundamental for assessing their habitat use and how they may respond to changes in the environment. However, despite recent advances in bio-logging technology, collecting information on both at-sea movement patterns and activity budgets still remains difficult in small pelagic seabird species due to the constraints of instrument size. The short-tailed shearwater, the most abundant seabird species in Australia (ca 23 million individuals), is a highly pelagic procellariiform. Despite its ecological importance to the region, almost nothing is known about its at-sea behaviour, in particular, its foraging activity. Using a combination of GPS and tri-axial accelerometer data-loggers, the fine scale three-dimensional foraging behaviour of 10 breeding individuals from two colonies was investigated. Five at-sea behaviours were identified: (1) resting on water, (2) flapping flight, (3) gliding flight, (4) foraging (i.e., surface foraging and diving events), and (5) taking-off. There were substantial intra- and inter- individual variations in activity patterns, with individuals spending on average 45.8% (range: 17.1-70.0%) of time at sea resting on water and 18.2% (range: 2.3-49.6%) foraging. Individuals made 76.4 ± 65.3 dives (range: 8-237) per foraging trip (mean duration 9.0 ± 1.9 s), with dives also recorded during night-time. With the continued miniaturisation of recording devices, the use of combined data-loggers could provide us with further insights into the foraging behaviour of small procellariiforms, helping to better understand interactions with their prey.

Utilisation of intensive foraging zones by female Australian fur seals

Within a heterogeneous environment, animals must efficiently locate and utilise foraging patches. One way animals can achieve this is by increasing residency times in areas where foraging success is highest (area-restricted search). For air-breathing diving predators, increased patch residency times can be achieved by altering both surface movements and diving patterns. The current study aimed to spatially identify the areas where female Australian fur seals allocated the most foraging effort, while simultaneously determining the behavioural changes that occur when they increase their foraging intensity. To achieve this, foraging behaviour was successfully recorded with a FastLoc GPS logger and dive behaviour recorder from 29 individual females provisioning pups. Females travelled an average of 118 ± 50 km from their colony during foraging trips that lasted 7.3 ± 3.4 days. Comparison of two methods for calculating foraging intensity (first-passage time and first-passage time modified to include diving behaviour) determined that, due to extended surface intervals where individuals did not travel, inclusion of diving behaviour into foraging analyses was important for this species. Foraging intensity 'hot spots' were found to exist in a mosaic of patches within the Bass Basin, primarily to the south-west of the colony. However, the composition of benthic habitat being targeted remains unclear. When increasing their foraging intensity, individuals tended to perform dives around 148 s or greater, with descent/ascent rates of approximately 1.9 m•s-1 or greater and reduced postdive durations. This suggests individuals were maximising their time within the benthic foraging zone. Furthermore, individuals increased tortuosity and decreased travel speeds while at the surface to maximise their time within a foraging location. These results suggest Australian fur seals will modify both surface movements and diving behaviour to maximise their time within a foraging patch.

Incorporating anthropogenic effects into trophic ecology: predator-prey interactions in a human-dominated landscape

Apex predators perform important functions that regulate ecosystems worldwide. However, little is known about how ecosystem regulation by predators is influenced by human activities. In particular, how important are top-down effects of predators relative to direct and indirect human-mediated bottom-up and top-down processes? Combining data on species' occurrence from camera traps and hunting records, we aimed to quantify the relative effects of top-down and bottom-up processes in shaping predator and prey distributions in a human-dominated landscape in Transylvania, Romania. By global standards this system is diverse, including apex predators (brown bear and wolf), mesopredators (red fox) and large herbivores (roe and red deer). Humans and free-ranging dogs represent additional predators in the system. Using structural equation modelling, we found that apex predators suppress lower trophic levels, especially herbivores. However, direct and indirect top-down effects of humans affected the ecosystem more strongly, influencing species at all trophic levels. Our study highlights the need to explicitly embed humans and their influences within trophic cascade theory. This will greatly expand our understanding of species interactions in human-modified landscapes, which compose the majority of the Earth's terrestrial surface.

Benefits of group foraging depend on prey type in a small marine predator, the Little Penguin

Group foraging provides predators with advantages in over-powering prey larger than themselves or in aggregating small prey for efficient exploitation. For group-living predatory species, cooperative hunting strategies provide inclusive fitness benefits. However, for colonial-breeding predators, the benefit pay-offs of group foraging are less clear due to the potential for intra-specific competition. We used animal-borne cameras to determine the prey types, hunting strategies, and success of little penguins (Eudyptula minor), a small, colonial breeding air-breathing marine predator that has recently been shown to display extensive at-sea foraging associations with conspecifics. Regardless of prey type, little penguins had a higher probability of associating with conspecifics when hunting prey that were aggregated than when prey were solitary. In addition, success was greater when individuals hunted schooling rather than solitary prey. Surprisingly, however, success on schooling prey was similar or greater when individuals hunted on their own than when with conspecifics. These findings suggest individuals may be trading-off the energetic gains of solitary hunting for an increased probability of detecting prey within a spatially and temporally variable prey field by associating with conspecifics.

Life history and the ecology of stress: how do glucocorticoid hormones influence life-history variation in animals?

Glucocorticoids hormones (GCs) are intuitively important for mediation of age-dependent vertebrate life-history transitions through their effects on ontogeny alongside underpinning variation in life-history traits and trade-offs in vertebrates. These concepts largely derive from the ability of GCs to alter energy allocation, physiology and behaviour that influences key life-history traits involving age-specific life-history transitions, reproduction and survival. Studies across vertebrates have shown that the neuroendocrine stress axis plays a role in the developmental processes that lead up to age-specific early life-history transitions. While environmental sensitivity of the stress axis allows for it to modulate the timing of these transitions within species, little is known as to how variation in stress axis function has been adapted to produce interspecific variation in the timing of life-history transitions. Our assessment of the literature confirms that of previous reviews that there is only equivocal evidence for correlative or direct functional relationships between GCs and variation in reproduction and survival. We conclude that the relationships between GCs and life-history traits are complex and general patterns cannot be easily discerned with current research approaches and experimental designs. We identify several future research directions including: (i) integration of proximate and ultimate measures, including longitudinal studies that measure effects of GCs on more than one life-history trait or in multiple environmental contexts, to test explicit hypotheses about how GCs and life-history variation are related and (ii) the measurement of additional factors that modulate the effects of GCs on life-history traits (e.g. GC receptors and binding protein levels) to better infer neurendocrine stress axis actions. Conceptual models of HPA/I axis actions, such as allostatic load and reactive scope, to some extent explicitly predict the role of GCs in a life-history context, but are descriptive in nature. We propose that GC effects on life-history transitions, survival probabilities and fecundity can be modelled in existing quantitative demographic frameworks to improve our understanding of how GC variation influences life-history evolution and GC-mediated effects on population dynamics Lay Summary Functional Ecology

Disentangling the cause of a catastrophic population decline in a large marine mammal

Considerable uncertainties often surround the causes of long-term changes in population abundance. One striking example is the precipitous decline of southern sea lions (SSL; Otariaflavescens) at the Falkland Islands, from 80 555 pups in the mid 1930s to just 5506 pups in 1965. Despite an increase in SSL abundance over the past two decades, the population has not recovered, with the number of pups born in 2014 (minimum 4443 pups) less than 6% of the 1930s estimate. The order-of-magnitude decline is primarily attributed to commercial sealing in Argentina. Here, we test this established paradigm and alternative hypotheses by assessing (1) commercial sealing at the Falkland Islands, (2) winter migration of SSL from the Falkland Islands to Argentina, (3) whether the number of SSL in Argentina could have sustained the reported level of exploitation, and (4) environmental change. The most parsimonious hypothesis explaining the SSL population decline was environmental change. Specifically, analysis of 160 years of winter sea surface temperatures revealed marked changes, including a period of warming between 1930 and 1950 that was consistent with the period of SSL decline. Sea surface temperature changes likely influenced the distribution or availability of SSL prey and impacted its population dynamics. We suggest that historical harvesting may not always be the "smoking gun" as is often purported. Rather, our conclusions support the growing evidence for bottom-up forcing on the abundance of species at lower trophic levels (e.g., plankton and fish) and resulting impacts on higher trophic levels across a broad range of ecosystems.

Feeding ecology of King George whiting Sillaginodes punctatus (Perciformes) recruits in seagrass and unvegetated habitats. Does diet reflect habitat utilization?

This study investigated the feeding ecology of King George whiting Sillaginodes punctatus recruits to determine how diet composition varies between habitat types (seagrass and unvegetated habitats), and between sites separated by distance. Broad-scale sampling of seagrass and unvegetated habitats at nine sites in Port Phillip Bay (Australia) indicated the diet composition varied more by distance into the bay than by habitat. Near the entrance to the bay the diet was dominated by harpacticoids and gammarid amphipods, in the middle reaches of the bay the diet was completely dominated by harpacticoids, while at sites furthest into the bay, mysids and crab zoea were also important. Abundances of prey in guts was significantly higher between 1000 and 2200 hours compared with other times, indicating diurnal feeding. Laboratory determined gut evacuation rate (based on an exponential model) was estimated to be -0·54. Daily rations were highly variable among sites and habitat types. Sillaginodes punctatus recruits consumed much higher quantities of prey on unvegetated habitat than seagrass habitat at some middle reach sites; with prey consumption of harpacticoid copepods on unvegetated habitat approaching 3000 individuals per day at one site. The results of this study provide insight into why habitat associations of S. punctatus recruits within mosaics of seagrass and unvegetated habitat show high spatial variation.