Habitat selection and behaviour of a reintroduced passerine: linking experimental restoration, behaviour and habitat ecology.

Habitat restoration can play an important role in recovering functioning ecosystems and improving biodiversity. Restoration may be particularly important in improving habitat prior to species reintroductions. We reintroduced seven brown treecreeper (Climacteris picumnus) social groups into two nature reserves in the Australian Capital Territory in south-eastern Australia. This study provided a unique opportunity to understand the interactions between restoration ecology, behavioural ecology and habitat ecology. We examined how experimental restoration treatments (addition of coarse woody debris, variations in ground vegetation cover and nest box installation) influenced the behaviour and microhabitat use of radio-tracked individuals to evaluate the success of restoration treatments. The addition of coarse woody debris benefited the brown treecreeper through increasing the probability of foraging on a log or on the ground. This demonstrated the value of using behaviour as a bio-indicator for restoration success. Based on previous research, we predicted that variations in levels of ground vegetation cover would influence behaviour and substrate use, particularly that brown treecreepers would choose sites with sparse ground cover because this allows better access to food and better vigilance for predators. However, there was little effect of this treatment, which was likely influenced by the limited overall use of the ground layer. There was also little effect of nest boxes on behaviour or substrate use. These results somewhat confound our understanding of the species based on research from extant populations. Our results also have a significant impact regarding using existing knowledge on a species to inform how it will respond to reintroduction and habitat restoration. This study also places great emphasis on the value of applying an experimental framework to ecological restoration, particularly when reintroductions produce unexpected outcomes.

Habitat selection and post-release movement of reintroduced brown treecreeper individuals in restored temperate woodland.

It is essential to choose suitable habitat when reintroducing a species into its former range. Habitat quality may influence an individual's dispersal decisions and also ultimately where they choose to settle. We examined whether variation in habitat quality (quantified by the level of ground vegetation cover and the installation of nest boxes) influenced the movement, habitat choice and survival of a reintroduced bird species. We experimentally reintroduced seven social groups (43 individuals) of the brown treecreeper (Climacteris picumnus) into two nature reserves in south-eastern Australia. We radio-tracked 18 brown treecreepers from release in November 2009 until February 2010. We observed extensive movements by individuals irrespective of the release environment or an individual's gender. This indicated that individuals were capable of dispersing and actively selecting optimum habitat. This may alleviate pressure on wildlife planners to accurately select the most optimum release sites, so long as the species' requirements are met. There was significant variation in movement between social groups, suggesting that social factors may be a more important influence on movement than habitat characteristics. We found a significant effect of ground vegetation cover on the likelihood of settlement by social groups, with high rates of settlement and survival in dry forests, rather than woodland (where the species typically resides), which has implications for the success of woodland restoration. However, overall the effects of variation in habitat quality were not as strong as we had expected, and resulted in some unpredicted effects such as low survival and settlement in woodland areas with medium levels of ground vegetation cover. The extensive movement by individuals and unforeseen effects of habitat characteristics make it difficult to predict the outcome of reintroductions, the movement behaviour and habitat selection of reintroduced individuals, particularly when based on current knowledge of a species' ecology.

Animal movement in dynamic landscapes: interaction between behavioural strategies and resource distributions.

Most ecological and evolutionary processes are thought to critically depend on dispersal and individual movement but there is little empirical information on the movement strategies used by animals to find resources. In particular, it is unclear whether behavioural variation exists at all scales, or whether behavioural decisions are primarily made at small spatial scales and thus broad-scale patterns of movement simply reflect underlying resource distributions. We evaluated animal movement responses to variable resource distributions using the grey teal (Anas gracilis) in agricultural and desert landscapes in Australia as a model system. Birds in the two landscapes differed in the fractal dimension of their movement paths, with teal in the desert landscape moving less tortuously overall than their counterparts in the agricultural landscape. However, the most striking result was the high levels of individual variability in movement strategies, with different animals exhibiting different responses to the same resources. Teal in the agricultural basin moved with both high and low tortuosity, while teal in the desert basin primarily moved using low levels of tortuosity. These results call into question the idea that broad-scale movement patterns simply reflect underlying resource distributions, and suggest that movement responses in some animals may be behaviourally complex regardless of the spatial scale over which movement occurs.

Positive effects of helpers on reproductive success in the brown treecreeper and the general importance of future benefits.

1. Numerous studies of cooperatively breeding species have tested for effects of helpers on reproductive success to evaluate hypotheses for the evolution of cooperation, but relatively few have used experimental or statistical approaches that control for the confounding effects of breeder and territory quality. 2. In the brown treecreeper Climacteris picumnus, most helpers are male offspring of the breeding pair that have delayed dispersal. We analysed 5 years of data (97 territory-years) using hierarchical linear modelling to test for effects of helpers on reproductive success while controlling for confounding factors. 3. The number of helpers was related positively to reproductive success even after controlling for differences between territories and breeders. A threshold effect was observed, with success increasing most with the presence of a second helper (i.e. at group size of four). 4. Feeding at the nest was one mechanism responsible for this effect, as larger groups had higher total feeding rates at all nesting stages. Higher total feeding rates, as well as higher feeding rates by helpers, were correlated in turn with greater reproductive success. 5. An analysis of the effects of helper feeding rate on reproductive success in groups with just one helper produced only weak support for a positive effect of helpers. Controlled comparisons of this kind utilize only a small fraction of the total data available and thus have limited statistical power compared to hierarchical or mixed-modelling. 6. A number of hypotheses to explain the evolution and maintenance of helping behaviour are consistent with our results for brown treecreepers including kin selection and hypotheses based on future direct benefits. 7. A previous synthesis of studies of helper effects that controlled for confounding factors suggested a pattern in which male helpers rarely have positive effects on reproductive success. However, revising that synthesis to include recent hierarchical or mixed-modelling studies suggests that helpers of both sexes usually have positive effects, and that the relative importance of future direct benefits may have been underestimated.

Dispersal range analysis: quantifying individual variation in dispersal behaviour.

A complete understanding of animal dispersal requires knowledge not only of its consequences at population and community levels, but also of the behavioural decisions made by dispersing individuals. Recent theoretical work has emphasised the importance of this dispersal process, particularly the phase in which individuals search the landscape for breeding opportunities. However, empirical advances are currently hampered by a lack of tools for quantifying these dispersal search tactics. Here, we review existing methods for quantifying movement that are appropriate for the dispersal search process, describe several new techniques that we developed for characterising movement and behaviour through an individual's dispersal range, and illustrate their use with data from Australasian treecreepers (Climacteridae). We also describe how the quantitative parameters we discuss are calculated in a freely available computer software package that we designed. Specifically, we present methods for calculating the area searched during dispersal, search rate, thoroughness, intensity, philopatry of search, timing of exploration, and surreptitiousness. When we applied this approach to the study of dispersal in treecreepers, we found that search area, philopatry and timing of exploration showed the greatest individual variation. Furthermore, search area, search rate, thoroughness and philopatry of search were all correlated, suggesting they may be useful parameters for further research on the causes and consequences of different dispersal search tactics. Finally, we make recommendations for modifying radiotracking protocols to facilitate more accurate assessment of individual variation in the dispersal process, and suggest future directions for this type of empirical work at the interface of population and behavioural ecology.