Hierarchical analysis of avian re-nesting behavior: mean, across-individual, and intra-individual responses

© 2015, Springer-Verlag Berlin Heidelberg. Anti-predator behavior is a key aspect of life history evolution, usually studied at the population (mean), or across-individual levels. However individuals can also differ in their intra-individual (residual) variation, but to our knowledge, this has only been studied once before in free-living animals. Here we studied the distances moved and changes in nest height and concealment between successive nesting attempts of marked pairs of grey fantails (Rhipidura albiscapa) in relation to nest fate, across the breeding season. We predicted that females (gender that decides where the nest is placed) should on average show adaptive behavioral responses to the experience of prior predation risk such that after an unsuccessful nesting attempt, replacement nests should be further away, higher from the ground, and more concealed compared with replacement nests after successful nesting attempts. We found that, on average, females moved greater distances to re-nest after unsuccessful nesting attempts (abandoned or depredated) in contrast to after a successful attempt, suggesting that re-nesting decisions are sensitive to risk. We found no consistent across-individual differences in distances moved, heights, or concealment. However, females differed by 53-fold (or more) in their intra-individual variability (i.e., predictability) with respect to distances moved and changes in nest height between nesting attempts, indicating that either some systematic variation went unexplained and/or females have inherently different predictability. Ignoring these individual differences in residual variance in our models obscured the effect of nest fate on re-nesting decisions that were evident at the mean level.

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.

Dingoes can help conserve wildlife and our methods can tell

Management of apex predators is among the most controversial wildlife management issues globally. In Australia, some ecologists have advocated using the dingo, Canis dingo, as a tool for conservation management, due to evidence that they suppress invasive mesopredators. Hayward & Marlow (Journal of Applied Ecology, 51, 2014 and 835) questioned the capacity of dingoes to provide benefits to native biodiversity due to their inability to eradicate foxes and cats. They also argued that indices of abundance commonly used in studies of mesopredator release by dingoes (namely, track-based indices) invalidate the conclusions of the studies. Hayward & Marlow caution conservation practitioners against incorporating dingoes into conservation programmes. Counter to their claims, we summarise research showing that the suppression of invasive mesopredators (cf. eradication) can enhance populations of native species and is therefore a meaningful conservation objective. We highlight literature supporting the hypothesis that dingoes suppress mesopredator abundance and activity, which in turn benefits native biodiversity. We show that Hayward & Marlow overlook many studies of carnivores that show track indices capture a large amount of the variation in the density of medium- and large-sized carnivores. Synthesis and applications. Practitioners cannot afford to wait to act given the perilous state of Australia's mammal species, and we argue that the evidence is sufficiently strong to justify managing dingoes for biodiversity conservation.

Predators help protect carbon stocks in blue carbon ecosystems

Predators continue to be harvested unsustainably throughout most of the Earth's ecosystems. Recent research demonstrates that the functional loss of predators could have far-reaching consequences on carbon cycling and, by implication, our ability to ameliorate climate change impacts. Yet the influence of predators on carbon accumulation and preservation in vegetated coastal habitats (that is, salt marshes, seagrass meadows and mangroves) is poorly understood, despite these being some of the Earth's most vulnerable and carbon-rich ecosystems. Here we discuss potential pathways by which trophic downgrading affects carbon capture, accumulation and preservation in vegetated coastal habitats. We identify an urgent need for further research on the influence of predators on carbon cycling in vegetated coastal habitats, and ultimately the role that these systems play in climate change mitigation. There is, however, sufficient evidence to suggest that intact predator populations are critical to maintaining or growing reserves of 'blue carbon' (carbon stored in coastal or marine ecosystems), and policy and management need to be improved to reflect these realities.

Resolving the value of the dingo in ecological restoration

There is global interest in restoring populations of apex predators, both to conserve them and to harness their ecological services. In Australia, reintroduction of dingoes (Canis dingo) has been proposed to help restore degraded rangelands. This proposal is based on theories and the results of studies suggesting that dingoes can suppress populations of prey (especially medium- and large-sized herbivores) and invasive predators such as red foxes (Vulpes vulpes) and feral cats (Felis catus) that prey on threatened native species. However, the idea of dingo reintroduction has met opposition, especially from scientists who query the dingo's positive effects for some species or in some environments. Here, we ask 'what is a feasible experimental design for assessing the role of dingoes in ecological restoration?' We outline and propose a dingo reintroduction experiment-one that draws upon the existing dingo-proof fence-and identify an area suitable for this (Sturt National Park, western New South Wales). Although challenging, this initiative would test whether dingoes can help restore Australia's rangeland biodiversity, and potentially provide proof-of-concept for apex predator reintroductions globally.

Multiple threats, or multiplying the threats? Interactions between invasive predators and other ecological disturbances

Invasive species have reshaped the composition of biomes across the globe, and considerable cost is now associated with minimising their ecological, social and economic impacts. Mammalian predators are among the most damaging invaders, having caused numerous species extinctions. Here, we review evidence of interactions between invasive predators and six key threats that together have strong potential to influence both the impacts of the predators, and their management. We show that impacts of invasive predators can be classified as either functional or numerical, and that they interact with other threats through both habitat- and community-mediated pathways. Ecosystem context and invasive predator identity are central in shaping variability in these relationships and their outcomes. Greater recognition of the ecological complexities between major processes that threaten biodiversity, including changing spatial and temporal relationships among species, is required to both advance ecological theory and improve conservation actions and outcomes. We discuss how novel approaches to conservation management can be used to address interactions between threatening processes and ameliorate invasive predator impacts.

Home range, habitat use and movements by the little raven (Corvus mellori) in a coastal peri-urban landscape

Context In peri-urban environments, high availability of anthropogenic resources may result in relatively high abundances of some species, with potentially negative implications for other native biota. Effective management of such impacts requires understanding of the spatial ecology of problem species. However, home range and habitat use have not been described for the little raven (Corvus mellori), a superabundant native predator that occurs in urban and natural habitats, including those where threatened shorebirds breed. Aims The aim of this study was to provide basic information on little raven home range, habitat use and movements in a coastal peri-urban landscape. Methods Between October 2011 and January 2012 we radio-tracked 20 little ravens captured in a coastal wetland (near Melbourne, Australia). Key results Little ravens were highly mobile, moving up to 9.9km in an hour (median≤2km), and had large ranges: Minimum Convex Polygons were 1664-9989ha (median≤3362ha). Although most birds used both anthropogenic and natural habitats, some birds strongly selected for coastal wetland habitat. Birds used multiple roosts during the study period, most of which occurred in grassland (58.7%) or urban (22.3%) areas. Movement of up to 8.3km (median≤2.2km) between roosts during the night was also detected. Conclusions Ravens were highly mobile and used large home ranges and a variety of habitats, with habitat preferences varying between birds. Implications Considering the large home ranges and inter-individual variation in habitat preferences of little raven populations, localised management to reduce their impacts on breeding shorebirds is unlikely to be successful. Journal compilation

Intense predation of non-colonial, ground-nesting bird eggs by corvid and mammalian predators

Context Loss of eggs to predators is a major cause of reproductive failure among birds. It is especially pronounced among ground-nesting birds because their eggs are accessible to a wide range of predators. Few studies document the main causes of clutch fate of ground-nesting birds. Aims The main objective of the present study was to identify the major egg predator of red-capped plovers (Charadrius ruficapillus). We also investigated the effectiveness of the following two primary strategies available to the plovers to avoid egg predation: (1) the placement of clutches under vegetative cover and (2) avoiding predators by nesting outside the peak season of predator occurrence. Methods Remote-sensing cameras were deployed on plover nests to identify egg predators and nests were monitored over four breeding seasons to document reproductive success and fate. An experiment using false clutches with model eggs investigated the influence of nest cover on the risk of egg predation throughout the year. Line-transect surveys were conducted to estimate the abundance of egg predators in and around the wetlands. Key results The little raven (Corvus mellori) was the major egg predator identified in 78.6% of red-capped plover clutches and in 92.4% of false clutches that were camera-monitored. The hatching success of plover eggs was not influenced by nest cover (P≤0.36), but model egg survival in false clutches improved significantly with the presence of nest cover (P≤0.02). The abundance of little ravens increased during the plover breeding season and was highly negatively correlated with false clutch survival (rpearson≤-0.768, P≤0.005). Conclusions Little ravens were the major predator of red-capped plover eggs and their abundance increased significantly during the plover breeding season. Any influence of nest cover on hatching success of eggs may have been masked by the extremely high rate of egg loss associated with the increased little raven abundance during the plover breeding season. Implications The high rate of egg predation is likely to have negative consequences on the local red-capped plover population, suggesting management is warranted. Little raven populations have expanded and, thus, their impact as egg predators needs to be investigated especially on threatened species. Journal compilation

Out of sight but not out of mind: corvids prey extensively on eggs of burrow-nesting penguins

Context Egg depredation is a major cause of reproductive failure among birds and can drive population declines. In this study we investigate predatory behaviour of a corvid (little raven; Corvus mellori) that has only recently emerged, leading to widespread and intense depredation of eggs of a burrow-nesting seabird (little penguin; Eudyptula minor). Aims The main objective of this study was to measure the rate of penguin egg depredation by ravens to determine potential threat severity. We also examined whether penguin burrow characteristics were associated with the risk of egg depredation. Ravens generally employ two modes of predatory behaviour when attacking penguin nests; thus we examined whether burrow characteristics were associated with these modes of attack. Methods Remote-sensing cameras were deployed on penguin burrows to determine egg predation rates. Burrow measurements, including burrow entrance and tunnel characteristics, were measured at the time of camera deployment. Key results Overall, clutches in 61% of monitored burrows (n≤203) were depredated by ravens, the only predator detected by camera traps. Analysis of burrow characteristics revealed two distinct types of burrows, only one of which was associated with egg depredation by ravens. Clutches depredated by ravens had burrows with wider and higher entrances, thinner soil or vegetation layer above the egg chamber, shorter and curved tunnels and greater areas of bare ground and whitewash near entrances. In addition, 86% were covered by bower spinach (Tetragonia implexicoma), through which ravens could excavate. Ravens used two modes to access the eggs: they attacked through the entrance (25% of burrow attacks, n≤124); or dug a hole through the burrow roof (75% of attacks, n≤124). Burrows that were subject to attack through the entrance had significantly shorter tunnels than burrows accessed through the roof. Conclusions The high rates of clutch loss recorded here highlight the need for population viability analysis of penguins to assess the effect of egg predation on population growth rates. Implications The subterranean foraging niche of a corvid described here may have implications for burrow-nesting species worldwide because many corvid populations are increasing, and they exhibit great capacity to adopt new foraging strategies to exploit novel prey. Journal compilation

Social networks in changing environments

Social network analysis (SNA) has become a widespread tool for the study of animal social organisation. However despite this broad applicability, SNA is currently limited by both an overly strong focus on pattern analysis as well as a lack of dynamic interaction models. Here, we use a dynamic modelling approach that can capture the responses of social networks to changing environments. Using the guppy, Poecilia reticulata, we identified the general properties of the social dynamics underlying fish social networks and found that they are highly robust to differences in population density and habitat changes. Movement simulations showed that this robustness could buffer changes in transmission processes over a surprisingly large density range. These simulation results suggest that the ability of social systems to self-stabilise could have important implications for the spread of infectious diseases and information. In contrast to habitat manipulations, social manipulations (e.g. change of sex ratios) produced strong, but short-lived, changes in network dynamics. Lastly, we discuss how the evolution of the observed social dynamics might be linked to predator attack strategies. We argue that guppy social networks are an emergent property of social dynamics resulting from predator–prey co-evolution. Our study highlights the need to develop dynamic models of social networks in connection with an evolutionary framework.

Effects of multiple disturbance processes on arboreal vertebrates in Eastern Australia: implications for management

Habitat loss and invasive predators increasingly threaten global biodiversity. Here we use a landscape-scale experimental approach to explore the individual and synergistic effects of logging and an invasive predator, the red fox Vulpes vulpes on two common native arboreal vertebrates (a predator and prey species) in south-eastern Australia. We used site occupancy methods to evaluate different models evaluating the effects of site specific forest logging disturbance, lethal fox baiting and forest structural elements for explaining variation in site occupancy of a large monitor lizard Varanus varius, and a marsupial prey, the common ringtail possum Pseudocheirus peregrinus across a complex forest landscape. Site occupancy of ringtail possum was influenced by habitat resources and the structural complexity of forest, which indirectly mediated predation risk. Presence of fox baiting had no direct effect on the ringtail site occupancy. In contrast, access to prey resources and fox baiting appeared to best explain site occupancy variation in monitor lizards across the landscape. While these species are affected primarily by separate disturbances, synergistic interactions between the processes may intensify their effects. Our results demonstrate that species susceptibility to disturbance processes are highly idiosyncratic. This approach makes efficient use of integrated modelling to aid conservation management at both local and landscape levels.

Evaluating and predicting risk to a large reptile (Varanus varius) from feral cat baiting protocols

Control of introduced predators to mitigate biodiversity impacts is a pressing conservation challenge. Across Australia feral cats (Felis catus) are a major threat to terrestrial biodiversity. Currently feral cat control is hindered by the limited utility of existing predator baiting methods. Further proposed control methods include use of the novel poison para-aminopropiophenone (PAPP) which may present a hazard to some native animal populations. Here we used experimental and predictive approaches to evaluate feral cat bait take by a large native Australian predatory reptile the Lace monitor (Varanus varius). These lizards would be expected to readily detect, ingest and consume a lethal dose (depending on toxin) from surface-laid baits intended for feral cat control if a precautionary approach was not adopted when baiting. We modelled V. varius bait take using experimental and predictive biophysical modelling approaches to evaluate temporal effects of climate variables on V. varius activity and hence potential for bait removal. Finally we conducted a pre-PAPP baiting site occupancy assessment of V. varius within Wilson Promontory National Park (WPNP) to provide a basis for monitoring any longer term population effects of cat baiting. V. varius removed 7 % of deployed baits from 73 % of bait stations across another study area in Far Eastern Victoria. Daily bait removal was positively correlated with maximum temperature and solar radiation. Biophysical modelling for Far Eastern Victoria predicted that maximum temperatures <19.5 °C prevented V. varius activity and hence opportunity for bait removal. V. varius in WPNP was undetectable suggesting aerial baiting posed limited hazard to this species at this location. Depending how climate influences annual activity patterns and the specific poison, surface-laid baits could pose a significant mortality risk to V. varius. However, use of biophysical models to predict periods of V. varius inactivity may provide a novel means to reduce non-target bait take by this predator.

Stress triangle: do introduced predators exert indirect costs on native predators and prey?

Non-consumptive effects of predators on each other and on prey populations often exceed the effects of direct predation. These effects can arise from fear responses elevating glucocorticoid (GC) hormone levels (predator stress hypothesis) or from increased vigilance that reduces foraging efficiency and body condition (predator sensitive foraging hypothesis); both responses can lead to immunosuppression and increased parasite loads. Non-consumptive effects of invasive predators have been little studied, even though their direct impacts on local species are usually greater than those of their native counterparts. To address this issue, we explored the non-consumptive effects of the invasive red fox Vulpes vulpes on two native species in eastern Australia: a reptilian predator, the lace monitor Varanus varius and a marsupial, the ringtail possum Pseudocheirus peregrinus. In particular, we tested predictions derived from the above two hypotheses by comparing the basal glucocorticoid levels, foraging behaviour, body condition and haemoparasite loads of both native species in areas with and without fox suppression. Lace monitors showed no GC response or differences in haemoparasite loads but were more likely to trade safety for higher food rewards, and had higher body condition, in areas of fox suppression than in areas where foxes remained abundant. In contrast, ringtails showed no physiological or behavioural differences between fox-suppressed and control areas. Predator sensitive foraging is a non-consumptive cost for lace monitors in the presence of the fox and most likely represents a response to competition. The ringtail's lack of response to the fox potentially represents complete naiveté or strong and rapid selection to the invasive predator. We suggest evolutionary responses are often overlooked in interactions between native and introduced species, but must be incorporated if we are to understand the suite of forces that shape community assembly and function in the wake of biological invasions.

De novo assembly and characterization of the invasive Northern Pacific Seastar transcriptome

Invasive species are a major threat to global biodiversity but can also serve as valuable model systems to examine important evolutionary processes. While the ecological aspects of invasions have been well documented, the genetic basis of adaptive change during the invasion process has been hampered by a lack of genomic resources for the majority of invasive species. Here we report the first larval transcriptomic resource for the Northern Pacific Seastar, Asterias amurensis, an invasive marine predator in Australia. Approximately 117.5 million 100 base-pair (bp) paired-end reads were sequenced from a single RNA-Seq library from a pooled set of full-sibling A. amurensis bipinnaria larvae. We evaluated the efficacy of a pre-assembly error correction pipeline on subsequent de novo assembly. Error correction resulted in small but important improvements to the final assembly in terms of mapping statistics and core eukaryotic genes representation. The error-corrected de novo assembly resulted in 115,654 contigs after redundancy clustering. 41,667 assembled contigs were homologous to sequences from NCBI's non-redundant protein and UniProt databases. We assigned Gene Ontology, KEGG Orthology, Pfam protein domain terms and predicted protein-coding sequences to > 36,000 contigs. The final transcriptome dataset generated here provides functional information for 18,319 unique proteins, comprising at least 11,355 expressed genes. Furthermore, we identified 9,739 orthologs to P. miniata proteins, evaluated our annotation pipeline and generated a list of 150 candidate genes for responses to several environmental stressors that may be important for adaptation of A. amurensis in the invasive range. Our study has produced a large set of A. amurensis RNA contigs with functional annotations that can serve as a resource for future comparisons to other echinoderm transcriptomes and gene expression studies. Our data can be used to study the genetic basis of adaptive change and other important evolutionary processes during a successful invasion.