Multi-year lactation and its consequences in Bornean orangutans (Pongo pygmaeus wurmbii)

In most mammals, females pay for reproduction by dramatically increasing net energy intake from conception to mid- or late lactation. To do this, they time their reproductive events in relation to environmental cycles so that periods of peak food availability coincide with peak demand or are used to build energy stores. This timing is not possible in species with slow development in which lactation is prolonged over a multi-year period with fluctuating food availability. Here, mothers are expected to sustain a stable but generally lower level of nutrient transfer. In a sample of over 1,050 complete follow days of eight mother–infant pairs collected over 7 years, we document maternal effort for wild Bornean orangutans (Pongo pygmaeus wurmbii) over their average 6.5-year lactation period. As predicted, maternal feeding time was independent of the age of her growing offspring, indicating a stable sustained “plateau” effort of ≤ 25 % above baseline level, instead of a short peak lactation as seen in seasonal breeders. Infant orangutans started to regularly supplement milk with self-harvested food when they were 1–1.5 years old, indicating milk intake was insufficient from this age onwards, even though maternal effort did not decrease. We expect the same regulation of sustained maternal effort in other large and large-brained mammals with slow infant development. We also predict that mother–infant conflict over suckling may show another peak at the onset of the milk?+?solid food phase, in addition to the well-known conflict around the endpoint of lactation (weaning), which is reached after a long and gradual increase in solid food intake by the infant.     

Food abundance affects energy intake and reproduction in frugivorous female Assamese macaques

In most mammals, female fertility and reproduction are strongly influenced by nutritional status and, therefore, by foraging conditions. Here, we investigate the relationship between food resources, feeding competition, energy intake and reproduction in a group of wild female Assamese macaques (Macaca assamensis) in northeastern Thailand. Over 2,100 h of data on feeding behaviour, energy intake and activity budgets were combined with data on resource characteristics, female reproduction and physical condition. We found that an increase in food availability had a positive effect on female energy intake and conception rates. In addition, it appeared that females incurred energetic costs during lactation and that females with a better physical condition during the mating season were more likely to conceive. The annual birth season occurred a few months before the annual peak in food availability, causing peak lactation to coincide with a period of high food availability. This suggests that females use the increased food abundance to compensate for the energetic costs of lactation. Neither energy intake rates nor activity budgets were influenced by female dominance rank, even during periods when the levels of contest competition were predicted to be high. In line with this, we found no evidence for rank-related differences in reproduction. The apparently limited influence of feeding competition in female Assamese macaques adds to the debate on the extent to which patterns in feeding competition and fitness can reliably be predicted based on ecological conditions. We suggest that this may partially be resolved by including potential competition-reducing mechanisms into the predictive framework.     

Why do most tropical animals reproduce seasonally? Testing hypotheses on an Australian snake

Most species reproduce seasonally, even in the tropics where activity occurs year-round. Squamate reptiles provide ideal model organisms to clarify the ultimate (adaptive) reasons for the restriction of reproduction to specific times of year. Females of almost all temperate-zone reptile species produce their eggs or offspring in the warmest time of the year, thereby synchronizing embryogenesis with high ambient temperatures. However, although tropical reptiles are freed from this thermal constraint, most do not reproduce year-round. Seasonal reproduction in tropical reptiles might be driven by biotic factors (e.g., peak periods of predation on eggs or hatchlings, or food for hatchlings) or abiotic factors (e.g., seasonal availability of suitably moist incubation conditions). Keelback snakes (Tropidonophis mairii, Colubridae) in tropical Australia reproduce from April to November, but with a major peak in May-June. Our field studies falsify hypotheses that invoke biotic factors as explanations for this pattern: the timing of nesting does not minimize predation on eggs, nor maximize food availability or survival rates for hatchlings. Instead, our data implicate abiotic factors: female keelbacks nest most intensely soon after the cessation of monsoonal rains when soils are moist enough to sustain optimal embryogenesis (wetter nests produce larger hatchlings, that are more likely to survive) but are unlikely to become waterlogged (which is lethal to eggs). Thus, abiotic factors may favor seasonal reproduction in tropical as well as temperate-zone animals.     

Continuous growth facilitates feeding and reproduction: impact of size on energy allocation patterns for organisms with indeterminate growth

Body size has great influence on feeding, reproduction, and ecological importance. This study measures growth, reproduction, and feeding for several northeastern Pacific intertidal invertebrates that have indeterminate growth. In all species studied, linear size (length, diameter) showed asymptotic growth fit by the von Bertalanffy growth function, supporting the notion that less energy is allocated to growth with age because of increased reproduction. However, these same species displayed a continuous, roughly linear increase in volume with age. Both reproductive output and food intake were shown to scale proportionally with volume. This indicates that some species with indeterminate growth do not reduce energy allocation to growth with age but instead display continuous volumetric growth that facilitates increases in feeding rate and reproductive output with age and size. A simple allometric model is proposed to describe constant volumetric growth rates and linear increases in reproduction with age.     

Landscape-scale resources promote colony growth but not reproductive performance of bumble bees

Variation in the availability of food resources over space and time is a likely driver of how landscape structure and composition affect animal populations. Few studies, however, have directly assessed the spatiotemporal variation in resource availability that arises from landscape pattern, or its effect on populations and population dynamic parameters. We tested the effect of floral resource availability at the landscape scale on the numbers of worker, male, and queen offspring produced by bumble bee, Bombus vosnesen?kii, colonies experimentally placed within complex agricultural-natural landscapes. We quantified flower densities in all land use types at different times of the season and then used these data to calculate spatially explicit estimates of floral resources surrounding each colony. Floral availability strongly correlated with landscape structure, and different regions of the landscape showed distinct seasonal patterns of floral availability. The floral resources available in the landscape surrounding a colony positively affected the number of workers and males it produced. Production was more sensitive to early- than to later-season resources. Floral resources did not significantly affect queen production despite a strong correlation between worker number and queen number among colonies. No landscape produced high floral resources during both the early and late season, and seasonal consistency is likely required for greater queen production. Floral resources are important determinants of colony growth and likely affect the pollination services provided by bumble bees at a landscape scale. Spatiotemporal variation in floral resources across the landscape precludes a simple relationship between resources and reproductive success as measured by queens, but nonetheless likely influences the total abundance of bumble bees in our study region.     

Behavioral games involving a clever prey avoiding a clever predator: An individual-based model of dusky dolphins and killer whales

Faced with an intermittent but potent threat, animals exhibit behavior that allows them to balance foraging needs and avoid predators and over time, these behaviors can become hard-wired adaptations with both species trying to maximize their own fitness. In systems where both predator and prey share similar sensory modalities and cognitive abilities, such as with marine mammals, the dynamic nature of predator-prey interactions is poorly understood. The costs and benefits of these anti-predator adaptations need to be evaluated and quantified based on the dynamic engagement of predator and prey. Many theoretic models have addressed the complexity of predator-prey relationships, but few have translated into testable mechanistic models. In this study, we developed a spatially-explicit, geo-referenced, individual-based model of a prototypical adult dusky dolphin off Kaikoura, New Zealand facing a more powerful, yet infrequent predator, the killer whale. We were interested in two primary objectives, (1) to capture the varying behavioral game between a clever prey and clever predator based on our current understanding of the Kaikoura system, (2) to compare evolutionary costs vs. benefits (foraging time and number of predator encounters) for an adult non-maternal dusky dolphin at various levels of killer whale-avoidance behaviors and no avoidance rules. We conducted Monte Carlo simulations to address model performance and parametric uncertainty. Mantel tests revealed an 88% correlation (426 × 426 distance matrix, km²) between observed field sightings of dusky dolphins with model generated sightings for non-maternal adult dusky dolphin groups. Simulation results indicated that dusky dolphins incur a 2.7% loss in feeding time by evolving the anti-predator behavior of moving to and from the feeding grounds. Further, each evolutionary strategy we explored resulted in dolphins incurring an additional loss of foraging time. At low killer whale densities (appearing less than once every 3 days), each evolutionary strategy simulated converged towards the evolutionary cost of foraging, that is, the loss in foraging time approached the 2.7% loss experienced by evolving near shore-offshore movement behavior. However, the highest level of killer whale presence resulted in 38% decreases in foraging time. The biological significance of these losses potentially incurred by a dusky dolphin is dependent on various factors from dolphin group foraging behavior and individual energy needs to dolphin prey availability and behavior.     

The effect of reproductive condition on the foraging behavior of female hoary bats,Lasiurus cinereus

Summary Female mammals experience larg changes in time and energy budgets associated with reproduction and these may influence the foraging strategies of individuals. I studied the changes in foraging behavior associated with reproduction in female hoary bats, Lasiurus cinereus. As lactation progressed, individuals departed to forage earlier in the evening and spent more time foraging per night and less time roosting with their young. Foraging time increased by at least 73% between early lactation and fledging and then declined as the young became independent. Females with two young foraged for longer than did those with one and females with pre- and postfledging young foraged in different habitats. The changes in foraging time suggest that foraging activity of female L. cinereus is constrained and individuals act as time minimizers, adjusting their foraging behavior to meet current energy demand. Predation risk is unlikely to constrain the behavior of these bats. However, maximizing energy intake throughout lactation may not be the optimal strategy because storing excess energy increases flight cost and may reduce foraging efficiency. The need to keep newborn young warm may also influence foraging time. Such constraints, causing changes in foraging activity, may alter the availability of habitats and prey and must be considered when modelling foraging strategies. In addition, changes in flight time may significantly alter the energy budgets of bats in different stages of reproduction.     

Killer whale predation on marine mammals at Punta Norte,Argentina; food sharing,provisioning and foraging strategy

Summary The social dynamics of killer whales (Orcinus orca) that hunt marine mammals are apparently highly flexible, though strong individual associations do exist. The killer whales at Punta Norte offer an unusually detailed view of association patterns and foraging behaviour, and suggest a pattern of behaviour that optimizes hunting efficiency with exception only to strong associations between some individuals and the provisioning and training of offspring. The main points from this paper are as follows: First, hunting effort was concentrated where the capture rate was greatest. All pods selectively attacked the prey type for which they had the highest capture rate. The amount of southern sea lion prey captured was approximately equal to the estimated minimum energetic requirement for killer whales based on weight. Secondly, one whale in each pod did the majority of the hunting, and then provisioned the others in the pod. It was clear on numerous occasions that food was shared. A review of reported incidences of killer wales taking marine mammal prey suggests that it is common for a subset of the individuals in a pod to hunt. These results are discussed in the context of the evolution of foraging behaviour.Offprint requests to: A.R. Hoelzel at the first address     

Phytoplankton food quality determines time windows for successful zooplankton reproductive pulses

Recruitment success at the early life stages is a critical process for zooplankton demography. Copepods often dominate the zooplankton in marine coastal zones and are prey of the majority of fish larvae. Hypotheses interpreting variations of copepod recruitment are based on the concepts of "naupliar predation," "nutritional deficiency," and "toxic effect" of diatom diets. Contradictory laboratory and field studies have reached opposite conclusions on the effects of diatoms on copepod reproductive success, blurring our view of marine food-web energy flow from diatoms to higher consumers by means of copepods. Here we report estimates of copepod feeding selectivity and reproduction in response to seasonally changing phytoplankton characteristics measured in a highly productive coastal upwelling area off the coast of central Chile. The variable phytoplankton diversity and changing food quality had a strong and highly significant impact on the feeding selectivity, reproduction, and larval survival of three indigenous copepod species. Seasonal changes in copepod feeding behavior were related to the alternating protozoan-diatom diets, mostly based on dinoflagellates and ciliates during winter and autumn (low highly unsaturated fatty acids [HUFA]/polyunsaturated fatty acids [PUFA] availability), but switched to a diet of centric and chain-forming diatoms (high HUFA/PUFA availability) during the spring/summer upwelling period. Ingestion of diatom cells induced a positive effect on egg production. However, a negative relationship was found between egg hatching success, naupliar survival, and diatom ingestion. Depending on the phytoplankton species, diets had different effects on copepod reproduction and recruitment. In consequence, it seems that the classical marine food web model does not apply to some coastal upwelling systems.     

Modelling the distribution and effect of heavy metals in an aquatic ecosystem

Models are reviewed describing the distribution and effect of heavy metals in an aquatic ecosystem. Since a model used for an impact statement should give the maximum concentration level rather than the seasonal variation, a model focussing on this situation is suggested. The basic differential equations describe (1) the variation in concentration of the toxicant per biomass dry matter in a given trophic level, and (2) the exchange of toxicant between sediment and water. Furthermore, since a substantial part of the heavy metal in an aquatic ecosystem is bound to suspended matter, an equation describing the equilibrium between dissolved and suspended matter must be included.A literature review has been carried out on the parameters used in the above mentioned equations and a demonstration, showing how it is possible to find approximate values for such parameters as excretion coefficient and uptake coefficient on the basis of a relationship between these two parameters and the size of an organism, is given.     

Fallow deer polyandry is related to fertilization insurance

Polyandry is widespread, but its adaptive significance is not fully understood. The hypotheses used to explain its persistence have rarely been tested in the wild and particularly for large, long-lived mammals. We investigated polyandry in fallow deer, using female mating and reproduction data gathered over 10 years. Females of this species produce a single offspring (monotocous) and can live to 23 years old. Overall, polyandry was evident in 12 % of females and the long-term, consistent proportion of polyandrous females observed, suggests that monandry and polyandry represent alternative mating strategies. Females were more likely to be polyandrous when their first mate had previously achieved high numbers of matings during the rut or was relatively old. However, polyandry was not related to the following factors: female age, the stage of the rut, the dominance ranks of mates, or the number of daily matings achieved by males. Polyandrous and monandrous multiple-mating females were not more likely than single-mating females to be observed with an offspring during the following year, and there were no significant differences in offspring size between these females. These results provide support for a fertility insurance hypothesis, with females remating if fertilization from the first mating was uncertain due to possible sperm depletion. The potential for different female mating strategies among large, polygynous mammals has generally been overlooked. Our findings highlight the complexity of female reproductive strategies and the possible trade-offs between fertilization success, preferences for high-quality males, and potential costs of polyandry, particularly for monotocous species.     

Energy allocation in a reef coral under varying resource availability

An organism’s pattern of resource allocation to reproduction and growth over time critically impacts on its lifetime reproductive success. During times of low resource availability, there are two fundamental, mutually exclusive strategies of energy investment: maintenance of somatic tissues to support survival and later reproduction or investment into an immediate reproductive event at the risk of subsequent death. Here, we examine energy allocation patterns in the coral Montipora digitata to determine whether energy investment during periods of resource shortage favours growth or reproduction in a sessile, modular marine species. We manipulated light regimes (two levels of shading) on plots within a shallow reef flat habitat (Orpheus Island, Great Barrier Reef, Australia) and quantified energy uptake (rates of net photosynthesis), energy investment into reproduction (E _R ), tissue growth per unit surface area (E _T ) and energy channelled into calcification (E _C ). With declining resource availability (i.e. reduced photosynthesis), relative energy investment shifted from high (~80%) allocation to tissue growth (E _R :E _T :E _C  = 11:81:8%) to an increasing proportion channelled into reproduction and skeletal growth (20:31:49%). At the lowest light regime, calcification was maintained but reproduction was halted and thus energy content per unit surface area of tissue declined, although no mortality was observed. The changing hierarchy in energy allocation among life functions with increasing resource limitation found here for an autotrophic coral, culminating in cessation of reproduction when limitations are severe, stands in contrast to observations from annual plants. However, the strategy may be optimal for maximising fitness components (growth, reproduction and survival) through time in marine modular animals.     

Where is the best site for wave energy exploitation? Case study along the coast of northern Latium (ITALY)

The present work proposes a siting process for the detection of a suitable site for wave energy exploitation. The choice of a suitable site is based on the good agreement between energy availability, environmental sustainability, and equipped facilities to exploit wave energy. The case study in the northern Latium coast is explicative because in this area there are several activities that affect marine ecosystems, and the introduction of renewable energies promote the reduction of anthropic pressures. The nearshore wave power is studied through the numerical wave model (CMS-Wave), and available wave buoys along the coast were used to compare numerical results. In correspondence with Civitavecchia harbour, the largest nearshore wave energy was found; the large depth in front of the breakwater allows to conserve a great part of offshore wave power, with an average dissipation rate of 10 % less than offshore, with mean annual available wave energy of 25.4 MWh/m and seasonal fluctuation of 5.4 MWh/m. This area appears to be an optimal site for nearshore and shoreline wave energy device tests and installations, for energy availability (intermediate level respect Mediterreanean Sea), low potential environmental impact, easier accessibility, and policy oriented towards a larger sustainability of harbour activities.     

Sex bias or equal opportunity? Patterns of maternal investment in bison

Summary In polygynous mammals, it may be adaptive for mothers to invest more in sons and/or to adjust the sex ratio of offspring in relation to body condition. Calving patterns were examined over an 8-year period (1982–1989) for a population of Bison bison in which barren females are not selectively culled. From these data, we tested predictions of the sex ratio adjustment hypothesis as well as two assumptions: (1) that offspring weight at the end of the period of parental investment (PI) is correlated with later condition, and (2) that maternal and offspring condition during the period of PI are correlated. In contrast to predictions, there was little evidence that mothers in better condition bear more sons. Short- and long-term measures of maternal condition (previous reproductive status, age, dominance status, pre-pubertal body weight, age at first reproduction, birth date, and the duration of the mother's own suckling period) were little related to offspring sex ratio, although the last calves of old females were nearly always female. Similarly, there was little evidence for sex-biased PI. Weights at about 7 months of age were greater for males than females; males also had somewhat later birth dates, suggesting either longer gestation or later conception. However, maternal reproductive costs, as measured by subsequent fecundity, weight loss, and interbirth intervals, did not vary with calf sex. Both assumptions of the model received some support. However, while maternal condition was correlated with offspring condition, there may be sex differences in investment patterns. Mothers appear better able to influence the condition of daughters than of sons. This sex difference may negate any benefit from male-biased investment.     

Seasonality of feeding and nutritional status during the austral winter in the Antarctic sea urchin Sterechinus neumayeri

The seasonal pattern of food intake and tissue energy status was measured over a 2 year period for two populations of the common Antarctic echinoid Sterechinus neumayeri living on contrasting substrata at Rothera Point, Adelaide Island, Antarctica. Food intake was estimated from faecal egestion, and both tissue mass and energy content assessed. Food availability was intensely seasonal, in that water column chlorophyll content and sediment pigment content varied markedly throughout the year. In response, both urchin populations showed an extremely seasonal cycle of faecal egestion, indicating a strong seasonality of feeding activity. Urchins from North Cove living on soft sediment fed at a considerably higher rate, and had a significantly larger Aristotle's lantern, than those in South Cove, 1 km away, living on hard substrata and taking a more cosmopolitan diet. Faecal egestion in both populations was zero for a 7 month period in the austral winter of 1997, and again for a 4 month period in 1998. During the austral summer of 1997/1998, 84% of the total annual energy intake took place in the period January to March. Significant decrease in gut tissue mass provided energy for maintenance in early winter, although progressive reduction of both gonad energy content (but without a detectable change in gonad mass) and body wall organic mass provided energy during the late winter period. The mass of reproductive tissue showed large differences between the two sites, but there was no marked decrease associated with spawning. Tissue proximate composition was assessed stoichiometrically from elemental composition and also checked by direct assay; gonad tissue was richer in lipid than gut tissue, though both were dominated by protein and contained only small amounts of carbohydrate. These data suggest that the very strong seasonality of food intake does not pose a significant energetic challenge for this species.     

Responses to threat by female bobucks,<Emphasis Type="Italic"> Trichosurus caninus</Emphasis>, during different stages of offspring development

The costs of reproduction are widely recognised as a major selective force in the evolution of various behavioural and life-history characteristics. In particular, the behaviour of reproductively active animals is likely to change when breeding increases risk of predation. We investigated the effect of an experimentally derived threat on the vigilance and escape behaviour of female Trichosurus caninus with and without dependent offspring and at different stages of offspring development. Females with offspring showed a heightened response to the threat in comparison to females without offspring. In addition, females with offspring displayed a stronger response at earlier stages of their offsprings development.Communicated by P. Bednekoff     

Renal lesions in Greenland sledge dogs ( Canis familiaris ) exposed to a natural dietary cocktail of persistent organic pollutants

Persistent organic pollutants such as Polychlorinated biphenyls (PCB) and dichloro diphenyl trichloroethane (DDT) are known to exert various adverse health effects in wildlife mammals. The impact from dietary intake of minke whale (Balaenoptera acutorostrata) blubber high in organohalogen and other chemical contaminants on renal morphology and function was investigated via a controlled study on West Greenland sledge dogs (Canis familiaris). Our results showed significantly higher frequencies of glomerular, tubular, and interstitial lesions in the exposed group. Furthermore, higher urine protein?:?creatinine ratio and plasma urea levels were found in the exposed group, which indicated a negative impact on kidney function via tubular and glomerular dysfunctions. The lesions were similar to those observed in top predator marine species–such as seals and polar bears–and humans exposed via diet to persistent environmental pollutants. The lesions may have impacts on the overall health of these animals. It is reasonable to suggest that chronic exposure to organic pollutants via a country diet of high trophic level marine organisms may have an impact on renal morphology and function in Arctic apex predators in general as well as Inuit people.     

Selecting a binary Markov model for a precipitation process

This paper uses rth-order categorical Markov chains to model the probability of precipitation. Several stationary and non-stationary high-order Markov models are proposed and compared using BIC. The number of parameters increases exponentially by adding the Markov order. Several classes of high-order Markov models are proposed which their increase of number of parameters are modest. For example models that use the number of precipitation days in a period prior to date, temperature of the previous day and sines/cosines periodic functions (to model the seasonality) are considered. The theory of partial likelihood is used to estimate the parameters. Parsimonious non-stationary first order Markov models with few seasonal terms are found optimal using BIC and temperature does not turn out to be a useful covariate. However BIC seems to underestimate the number of seasonal terms. We have also compared the results with AIC in some cases which tends to pick parsimonious models with more seasonal terms and higher order. We also show that ignoring seasonal terms result in picking higher order Markov chains. Finally we apply the methods to build confidence intervals for the probability of periods with no precipitation or low number of precipitation days in Calgary using historical data from 1980 to 2000.     

Public Willingness to Pay for Recovering and Downlisting Threatened and Endangered Marine Species

Abstract: Nonmarket valuation research has produced economic value estimates for a variety of threatened, endangered, and rare species around the world. Although over 40 value estimates exist, it is often difficult to compare values from different studies due to variations in study design, implementation, and modeling specifications. We conducted a stated‐preference choice experiment to estimate the value of recovering or downlisting 8 threatened and endangered marine species in the United States: loggerhead sea turtle (Caretta caretta), leatherback sea turtle (Dermochelys coriacea), North Atlantic right whale (Eubalaena glacialis), North Pacific right whale (Eubalaena japonica), upper Willamette River Chinook salmon (Oncorhynchus tshawytscha), Puget Sound Chinook salmon (Oncorhynchus tshawytscha), Hawaiian monk seals (Monachus schauinslandi), and smalltooth sawfish (Pristis pectinata). In May 2009, we surveyed a random sample of U.S. households. We collected data from 8476 households and estimated willingness to pay for recovering and downlisting the 8 species from these data. Respondents were willing to pay for recovering and downlisting threatened and endangered marine taxa. Willingness‐to‐pay values ranged from $40/household for recovering Puget Sound Chinook salmon to $73/household for recovering the North Pacific right whale. Statistical comparisons among willingness‐to‐pay values suggest that some taxa are more economically valuable than others, which suggests that the U.S. public's willingness to pay for recovery may vary by species.     

Contrasting diel patterns in vertical movement and locomotor activity of whale sharks at Ningaloo Reef

Activity patterns of animals often relate to environmental variables such as food availability and predation pressure. Technological advances are providing us with new tools to monitor and better understand these activity patterns. We used animal-attached data loggers recording acceleration and depth to compare activity patterns and vertical habitat use of whale sharks (Rhincodon typus) at Ningaloo Reef, Western Australia. Whale sharks showed a moderate reverse diel vertical migration but exhibited a clear crepuscular pattern in locomotory activity. Peak activity occurred at sunset, whereas vertical movement peaked prior to this. Typical ram surface filter feeding could be identified and occurred primarily during sunset and the first hours of night. At such times, direct observations indicated whale sharks were feeding on tropical krill swarms. Kinematic analysis of postural data and data from vertical movement suggests that whale sharks at Ningaloo spend ~8 min per day actively ram surface filter feeding. Considering the high biomass present in krill schools, it is estimated that whale sharks at Ningaloo have a similar energy intake as those at other aggregation sites. Diel patterns in activity and diving behaviour suggest that whale sharks have tuned their diving behaviour in anticipation of the formation of these high-density patches which appear to only be periodically, but predictably available at sunset. Our results confirm that diel patterns in vertical habitat selection and vertical movements do not necessarily reflect patterns in activity and foraging behaviour. Direct quantification of activity and behaviour is required in gaining accurate representation of diel activity patterns.