Environmental context explains Lévy and Brownian movement patterns of marine predators

An optimal search theory, the so-called Lévy-flight foraging hypothesis1, predicts that predators should adopt search strategies known as Lévy flights where prey is sparse and distributed unpredictably, but that Brownian movement is sufficiently efficient for locating abundant prey2, 3, 4. Empirical studies have generated controversy because the accuracy of statistical methods that have been used to identify Lévy behaviour has recently been questioned5, 6. Consequently, whether foragers exhibit Lévy flights in the wild remains unclear. Crucially, moreover, it has not been tested whether observed movement patterns across natural landscapes having different expected resource distributions conform to the theory’s central predictions. Here we use maximum-likelihood methods to test for Lévy patterns in relation to environmental gradients in the largest animal movement data set assembled for this purpose. Strong support was found for Lévy search patterns across 14 species of open-ocean predatory fish (sharks, tuna, billfish and ocean sunfish), with some individuals switching between Lévy and Brownian movement as they traversed different habitat types. We tested the spatial occurrence of these two principal patterns and found Lévy behaviour to be associated with less productive waters (sparser prey) and Brownian movements to be associated with productive shelf or convergence-front habitats (abundant prey). These results are consistent with the Lévy-flight foraging hypothesis1, 7, supporting the contention8, 9 that organism search strategies naturally evolved in such a way that they exploit optimal Lévy patterns.

All signals are not equal: acoustic signalling of individuality, sex and breeding status in a cooperative breeder

Repeated interactions between individuals in socially living animals select for the evolution of signals that convey information identifying individuals or categories of individuals, which may enable the discrimination of familiar versus unfamiliar individuals. Such information may help animals maximize their inclusive fitness by adjusting their own behaviour, allowing them to avoid conflict, preferentially direct help and/or ignore unreliable individuals. Acoustic signals in birds provide the potential to encode individual-specific information. We examined the degree to which individual identity, sex, breeding status, group membership and genetic relatedness were related to variability in six different call types, which occurred across a variety of different behavioural contexts in the apostlebird, Struthidea cinerea, a socially living and cooperatively breeding Australian passerine. We demonstrated that not all calls reflected the same extent of information. Of the six call types, call variation was related to individual identity in three call types, breeding status in two call types and sex and group relatedness in one call type. Finally, variation in two call types was not related to any of the measured variables. Our results suggest that some, but not all, acoustic signals in apostlebirds may be selected for individual distinctiveness between individuals and categories of individuals (male versus female, breeder versus nonbreeder), and these signals may be important in determining levels of cooperation and interaction between individuals in this cooperatively breeding society. © 2014 The Association for the Study of Animal Behaviour.

Odour-based discrimination of subspecies, species and sexes in an avian species complex, the crimson rosella

Olfaction is an ancient sensory capability, and yet while it is now widely recognized that birds have olfactory mechanisms, use of the sense within a social context has been largely overlooked. In our study, we aimed to determine, for the first time, whether plumage odour may contribute to avian subspecies discrimination. We used a species complex, the crimson rosella, Platycercus elegans, which exhibits large geographical and phenotypic differences. Across 2 years in a wild population of P.elegans elegans we tested whether females at the nest could: (1) discriminate odours of conspecifics; (2) discriminate odours of subspecies; (3) discriminate odours of sexes of conspecifics; and (4) habituate at different rates to odour treatments. We found that female response differed between odours of feathers of consubspecifics, heterosubspecifics, heterospecific controls and sham controls and between odours of sexes of conspecifics. Across all odour treatments, we found habituation to the odour and the rate of habituation differed between odour treatments. Our results indicate that P.e. elegans females are able to discriminate conspecifics, consubspecifics and sexes based on plumage odour. To our knowledge, this is the first work to show that birds of a certain subspecies can discriminate the odour of its own subspecies from that of other subspecies. Our findings suggest that olfaction in birds may play a larger role than hitherto considered, and may even act as a signal to maintain or promote population divergence. © 2014 The Association for the Study of Animal Behaviour.

Using repeatability to study physiological and behavioural traits: ignore time-related change at your peril

© 2015 The Association for the Study of Animal Behaviour. Broad sense repeatability, which refers to the extent to which individual differences in trait scores are maintained over time, is of increasing interest to researchers studying behavioural or physiological traits. Broad sense repeatability is most often inferred from the statistic R (the intraclass correlation, or narrow sense repeatability). However, R ignores change over time, despite the inherent longitudinal nature of the data (repeated measures over time). Here, we begin by showing that most studies ignore time-related change when estimating broad sense repeatability, and estimate R with low statistical power. Given this problem, we (1) outline how and why ignoring time-related change in scores (that occurs for whatever reason) can seriously affect estimates of the broad sense repeatability of behavioural or physiological traits, (2) discuss conditions in which various indices of R can or cannot provide reliable estimates of broad sense repeatability, and (3) provide suggestions for experimental designs for future studies. Finally, given that we already have abundant evidence that many labile traits are 'repeatable' in that broad sense (i.e. R>. 0), we suggest a shift in focus towards obtaining robust estimates of the repeatability of behavioural and physiological traits. Given how labile these traits are, this will require greater experimental (and/or statistical) control and larger sample sizes in order to detect and quantify change over time (if present).

Impacts of visitor number on Kangaroos housed in free-range exhibits

Free range exhibits are becoming increasingly popular in zoos as a means to enhance interaction between visitors and animals. However very little research exists on the impacts of visitors on animal behaviour and stress in free range exhibits. We investigated the effects of visitor number on the behaviour and stress physiology of Kangaroo Island (KI) Kangaroos, Macropus fuliginosus fuliginosus, and Red Kangaroos, Macropus rufus, housed in two free range exhibits in Australian zoos. Behavioural observations were conducted on individual kangaroos at each site using instantaneous scan sampling to record activity (e.g., vigilance, foraging, resting) and distance from the visitor pathway. Individually identifiable faecal samples were collected at the end of each study day and analysed for faecal glucocorticoid metabolite (FGM) concentration. When visitor number increased, both KI Kangaroos and Red Kangaroos increased the time spent engaged in visitor-directed vigilance and KI Kangaroos also increased the time spent engaged in locomotion and decreased the time spent resting. There was no effect of visitor number on the distance kangaroos positioned themselves from the visitor pathway or FGM concentration in either species. While there are limitations in interpreting these results in terms of fear of visitors, there was no evidence of adverse effects animal welfare in these study groups based on avoidance behaviour or stress physiology under the range of visitor numbers that we studied.

Individual differences in activity and habitat selection of juvenile queen conch evaluated using acceleration biologgers

Fine-scale differences in behaviour and habitat use have important ecological implications, but have rarely been examined in marine gastropods. We used tri-axial accelerometer loggers to estimate activity levels and movement patterns of the juvenile queen conch Lobatus gigas (n = 11) in 2 habitat types in Eleuthera, The Bahamas. In 2 manipulations in nearshore areas, queen conchs were equipped with accelerometers and released in adjacent coral rubble or seagrass habitats. Queen conchs were located approximately every 6 h during daylight by snorkeling, to measure individual differences in linear distance moved, and after 24 h they were relocated to an alternate habitat (24 h in each habitat). We found significant inter-individual variability in activity levels, but more consistent levels of activity between the 2 habitat types within individual queen conchs. Four (36%) of the individuals placed in seagrass moved back to the adjacent coral rubble habitat, suggesting selectivity for coral rubble. Individuals showed variable behavioural responses when relocated to the less preferable seagrass habitat, which may be related to differing stress-coping styles. Our results suggest that behavioural variability between individuals may be an important factor driving movement and habitat use in queen conch and, potentially, their susceptibility to human stressors. This study provides evidence of diverse behavioural (activity) patterns and habitat selectivity in a marine gastropod and highlights the utility of accelero meter biologgers for continuously monitoring animal behaviour in the wild.

Ontogeny of diving behaviour in the Australian sea lion: trials of adolescence in a late bloomer

1.Foraging behaviours of the Australian sea lion (Neophoca cinerea) reflect an animal working hard to exploit benthic habitats. Lactating females demonstrate almost continuous diving, maximize bottom time, exhibit elevated field metabolism and frequently exceed their calculated aerobic dive limit. Given that larger animals have disproportionately greater diving capabilities, we wanted to examine how pups and juveniles forage successfully.
2.Time/depth recorders were deployed on pups, juveniles and adult females at Seal Bay Conservation Park, Kangaroo Island, South Australia. Ten different mother/pup pairs were equipped at three stages of development (6, 15 and 23 months) to record the diving behaviours of 51 (nine instruments failed) animals.
3. Dive depth and duration increased with age. However, development was slow. At 6 months, pups demonstrated minimal diving activity and the mean depth for 23-month-old juveniles was only 44 ± 4 m, or 62% of adult mean depth.
4. Although pups and juveniles did not reach adult depths or durations, dive records for young sea lions indicate benthic diving with mean bottom times (2·0 ± 0·2 min) similar to those of females (2·1 ± 0·2 min). This was accomplished by spending higher proportions of each dive and total time at sea on or near the bottom than adults. Immature sea lions also spent a higher percentage of time at sea diving.
5. Juveniles may have to work harder because they are weaned before reaching full diving capability. For benthic foragers, reduced diving ability limits available foraging habitat. Furthermore, as juveniles appear to operate close to their physiological maximum, they would have a difficult time increasing foraging effort in response to reductions in prey. Although benthic prey are less influenced by seasonal fluctuations and oceanographic perturbations than epipelagic prey, demersal fishery trawls may impact juvenile survival by disrupting habitat and removing larger size classes of prey. These issues may be an important factor as to why the Australian sea lion population is currently at risk.

When does physiology limit the foraging behaviour of freely diving mammals?

Diving animals offer a unique opportunity to study the importance of physiological constraint and the limitation it can impose on animal's behaviour in nature. This paper examines the interaction between physiology and behaviour and its impact on the diving capability of five eared seal species (Family Otariidae; three sea lions and two fur seals). An important physiological component of diving marine mammals is the aerobic dive limit (ADL). The ADL of these five seal species was estimated from measurements of their total body oxygen stores, coupled with estimates of their metabolic rate while diving. The tendency of each species to exceed its calculated ADL was compared relative to its diving behaviour. Overall, our analyses reveal that seals which forage benthically (i.e. on the sea floor) have a greater tendency to approach or exceed their ADL compared to seals that forage epipelagically (i.e. near the sea surface). Furthermore, the marked differences in foraging behaviour and physiology appear to be coupled with a species demography. For example, benthic foraging species have smaller populations and lower growth rates compared to seal species that forage epipelagically. These patterns are relevant to the conservation and management of diving vertebrates.

Nestling testosterone is associated with begging behaviour and fledging success in the pied flycatcher, Ficedula hypoleuca

Animal signals are hypothesized to be costly in order to honestly reflect individual quality. Offspring solicitation signals given by nestling birds are thought to have evolved to advertise either need or individual quality. We tested the potential role of testosterone (T) in controlling the intensity of these signals by measuring begging behaviour as: (i) duration of the begging display and (ii) maximum height of the begging stretch, and by sampling endogenous T levels in nestling blood. We tested nestling pied flycatchers (Ficedula hypoleuca) using well-established experimental paradigm involving transient food deprivation to encourage begging behaviour and then blood-sampled nestlings at the end of these tests for T levels. Our results show that individual nestlings with the most intense begging displays had the highest circulating levels of T immediately after testing. In addition, we found substantial differences between broods in terms of circulating T. Finally, we found evidence that broods with higher levels of T showed increased fledging success, indicating a benefit for increased T production in nestlings. The potential trade-offs involved in T-mediated begging behaviour are discussed.

Influence of stocking rate and mixed grazing of Angora goats and Merino sheep on animal and pasture production in southern Australia : 1 : botanical composition, sward characteristics and availability of components of annual temperate pastures

The effects of animal species (AS; Angora goats, Merino sheep or goats and sheep mixed grazed together at ratio 1:1) and stocking rate (SR; 7.5, 10 and 12.5 animals/ha) on the availability, botanical composition and sward characteristics of annual temperate pastures under continuous grazing were determined in a replicated experiment from 1981 to 1984. AS and SR had significant effects on pasture availability and composition and many AS SR interactions were detected. The pastures grazed by sheep had significantly reduced content and proportion of subterranean clover and more undesirable grasses compared with those grazed by goats. There were no differences in dry matter availabilities between goat- and sheep-grazed pastures at 7.5/ha, but at 10 and 12.5/ha goat pastures had significantly increased availabilities of green grass, dead and green clover and less weeds compared with sheep pastures. There was a significant AS SR interaction for the density of seedlings in May following pasture germination. Between July and January, the height of pastures was greater under goats than sheep but from January to March pasture height declined more on goat-grazed than on sheep-grazed pastures. There was an AS SR interaction for incidence of bare ground. Increasing the SR increased bare ground in pastures grazed by sheep but no change occurred on pastures grazed by goats. Changes in pasture characteristics due to increased SR were minimised on pastures grazed by goats but the grazing of sheep caused larger and faster changes and the pastures were damaged at the highest SR. Goats did not always select the same herbage material as sheep, changed their selection between seasons and were not less selective than sheep. Angora goats were flexible grazers and continually adapted their grazing behaviour to changing herbage conditions. Goat grazing led to an increase in subterranean clover, an accumulation of dead herbage at the base of the sward, reduced bare ground, taller pastures in spring and a more stable botanical composition. Mixed-grazed pasture characteristics were altered with SR. With careful management Angora goats on sheep farms may be used to manipulate pasture composition, to speed up establishment of subterranean clover, to decrease soil erosion and to reduce weed invasion.

Energy metabolism and animal personality

In this paper we show how animal personality could explain some of the large inter-individual variation in resting metabolic rate (MR) and explore methodological and functional linkages between personality and energetics. Personality will introduce variability in resting MR measures because individuals consistently differ in their stress response, exploration or activity levels, all of which influence MR measurements made with respirometry and the doubly-labelled water technique. Physiologists try to exclude these behavioural influences from resting MR measurements, but animal personality research indicates that these attempts are unlikely to be successful. For example, because reactive animals “freeze” when submitted to a stress, their MR could be classified as “resting” because of immobility when in fact they are highly stressed with an elevated MR. More importantly, recent research demonstrating that behavioural responses to novel and highly artificial stimuli are correlated with both behaviour and fitness under more natural circumstances calls into question the wisdom of excluding these behavioural influences on MR measurements. The reason that intra-specific variation in resting MR are so weakly correlated with daily energy expenditure (DEE) and fitness, may be that the latter two measures fully incorporate personality while the former partially excludes its influence. Because activity, exploration, boldness and aggressiveness are energetically costly, personality and metabolism should be correlated and physiological constraints may underlie behavioural syndromes. We show how physiological ecologists can better examine behavioural linkages between personality and metabolism, as required to better understand the physiological correlates of personality and the evolutionary consequences of metabolic variability.

Factors affecting the foraging behaviour of the European shag : implications for seabird tracking studies

Seabird tracking has become an ever more popular tool to aid environmental procedures such as the designation of marine protected areas and environmental impact assessments. However, samples used are usually small and little consideration is given to experimental design and sampling protocol. European shags Phalacrocorax aristotelis were tracked using GPS technology over three breeding seasons and the following foraging trip characteristics: trip duration, trip distance, maximum distance travelled from the colony, size of area used and direction travelled from colony were determined for each foraging trip. The effect of sex, year of study, breeding site, number and age of chicks and the timing of tracking on foraging behaviour were investigated using a General Estimation Equation model. A range of sampling scenarios reflecting likely field sampling were also tested to compare how foraging behaviour differed depending on composition of the sample of birds tracked. Trip distance, trip duration, maximum distance travelled and size of area used were all significantly affected by the breeding site, and the number of chicks a tracked adult was raising. The effect of sex was also seen when examining trip distance, trip duration and the maximum distance travelled. The direction travelled on a foraging trip was also significantly affected by breeding site. This study highlights the importance of sampling regime and the influence that year, sex, age, number of chicks and breeding site can have on the foraging trip characteristics for this coastal feeding seabird. Given the logistical and financial constraints in tracking large numbers of individuals, this study identifies the need for researchers to consider the composition of their study sample to ensure any identified foraging areas are as representative as possible of the whole colony's foraging area.