Showing papers in "Journal of Animal Ecology in 2008"
TL;DR: This study provides a working guide to boosted regression trees (BRT), an ensemble method for fitting statistical models that differs fundamentally from conventional techniques that aim to fit a single parsimonious model.
Abstract: Summary 1 Ecologists use statistical models for both explanation and prediction, and need techniques that are flexible enough to express typical features of their data, such as nonlinearities and interactions 2 This study provides a working guide to boosted regression trees (BRT), an ensemble method for fitting statistical models that differs fundamentally from conventional techniques that aim to fit a single parsimonious model Boosted regression trees combine the strengths of two algorithms: regression trees (models that relate a response to their predictors by recursive binary splits) and boosting (an adaptive method for combining many simple models to give improved predictive performance) The final BRT model can be understood as an additive regression model in which individual terms are simple trees, fitted in a forward, stagewise fashion 3 Boosted regression trees incorporate important advantages of tree-based methods, handling different types of predictor variables and accommodating missing data They have no need for prior data transformation or elimination of outliers, can fit complex nonlinear relationships, and automatically handle interaction effects between predictors Fitting multiple trees in BRT overcomes the biggest drawback of single tree models: their relatively poor predictive performance Although BRT models are complex, they can be summarized in ways that give powerful ecological insight, and their predictive performance is superior to most traditional modelling methods 4 The unique features of BRT raise a number of practical issues in model fitting We demonstrate the practicalities and advantages of using BRT through a distributional analysis of the short-finned eel ( Anguilla australis Richardson), a native freshwater fish of New Zealand We use a data set of over 13 000 sites to illustrate effects of several settings, and then fit and interpret a model using a subset of the data We provide code and a tutorial to enable the wider use of BRT by ecologists
TL;DR: This work compared chemical extraction and mathematical correction methods for freshwater and marine fishes and aquatic invertebrates to better understand impacts of correction approaches on carbon (delta(13)C) and nitrogen (d delta(15)N) stable isotope data.
Abstract: Summary 1. Lipids have more negative δ 13 C values relative to other major biochemical compounds in plant and animal tissues. Although variable lipid content in biological tissues alters results and conclusions of δ 13 C analyses in aquatic food web and migration studies, no standard correction protocol exists. 2. We compared chemical extraction and mathematical correction methods for freshwater and marine fishes and aquatic invertebrates to better understand impacts of correction approaches on carbon ( δ 13 C) and nitrogen ( δ 15 N) stable isotope data. 3. Fish and aquatic invertebrate tissue δ 13 C values increased significantly following extraction for almost all species and tissue types relative to nonextracted samples. In contrast, δ 15 N was affected for muscle and whole body samples from only a few freshwater and marine species and had a limited effect for the entire data set. 4. Lipid normalization models, using C : N as a proxy for lipid content, predicted lipid-corrected δ 13 C for paired data sets more closely with parameters specific to the tissue type and species to which they were applied. 5. We present species- and tissue-specific models based on bulk C : N as a reliable alternative to chemical extraction corrections. By analysing a subset of samples before and after lipid extraction, models can be applied to the species and tissues of interest that will improve estimates of dietary sources using stable isotopes.
TL;DR: To this knowledge, this is the first time that empirical data on foraging distances and forage availability, at this resolution and scale, have been collected and combined for bumblebees.
Abstract: 1. Foraging range is a key aspect of the ecology of 'central place foragers'. Estimating how far bees fly under different circumstances is essential for predicting colony success, and for estimating bee-mediated gene flow between plant populations. It is likely to be strongly influenced by forage distribution, something that is hard to quantify in all but the simplest landscapes; and theories of foraging distance tend to assume a homogeneous forage distribution. 2. We quantified the distribution of bumblebee Bombus terrestris L. foragers away from experimentally positioned colonies, in an agricultural landscape, using two methods. We mass-marked foragers as they left the colony, and analysed pollen from foragers returning to the colonies. The data were set within the context of the 'forage landscape': a map of the spatial distribution of forage as determined from remote-sensed data. To our knowledge, this is the first time that empirical data on foraging distances and forage availability, at this resolution and scale, have been collected and combined for bumblebees. 3. The bees foraged at least 1.5 km from their colonies, and the proportion of foragers flying to one field declined, approximately linearly, with radial distance. In this landscape there was great variation in forage availability within 500 m of colonies but little variation beyond 1 km, regardless of colony location. 4. The scale of B. terrestris foraging was large enough to buffer against effects of forage patch and flowering crop heterogeneity, but bee species with shorter foraging ranges may experience highly variable colony success according to location.
TL;DR: The presence of O. brumata outbreaks in regions previously affected solely by E. autumnata outbreaks is likely to increase the effective duration of local outbreaks, and hence have profound implications for the subarctic birch forest ecosystem.
Abstract: 1. Range expansions mediated by recent climate warming have been documented for many insect species, including some important forest pests. However, whether climate change also influences the eruptive dynamics of forest pest insects, and hence the ecological and economical consequences of outbreaks, is largely unresolved. 2. Using historical outbreak records covering more than a century, we document recent outbreak range expansions of two species of cyclic geometrid moth, Operophtera brumata Bkh. (winter moth) and Epirrita autumnata L. (autumnal moth), in subarctic birch forest of northern Fennoscandia. The two species differ with respect to cold tolerance, and show strikingly different patterns in their recent outbreak range expansion. 3. We show that, during the past 15-20 years, the less cold-tolerant species O. brumata has experienced a pronounced north-eastern expansion into areas previously dominated by E. autumnata outbreaks. Epirrita autumnata, on the other hand, has expanded the region in which it exhibits regular outbreaks into the coldest, most continental areas. Our findings support the suggestion that recent climate warming in the region is the most parsimonious explanation for the observed patterns. 4. The presence of O. brumata outbreaks in regions previously affected solely by E. autumnata outbreaks is likely to increase the effective duration of local outbreaks, and hence have profound implications for the subarctic birch forest ecosystem.
TL;DR: The relative size of predators and prey had a pervasive structuring influence on biomass fluxes within this large-mammal food web, and predation become a negligible cause of mortality for megaherbivores substantially exceeding 1000 kg in adult body mass.
Abstract: 1. Size relationships are central in structuring trophic linkages within food webs, leading to suggestions that the dietary niche of smaller carnivores is nested within that of larger species. However, past analyses have not taken into account the differing selection shown by carnivores for specific size ranges of prey, nor the extent to which the greater carcass mass of larger prey outweighs the greater numerical representation of smaller prey species in the predator diet. Furthermore, the top-down impact that predation has on prey abundance cannot be assessed simply in terms of the number of predator species involved. 2. Records of found carcasses and cause of death assembled over 46 years in the Kruger National Park, South Africa, corrected for under-recording of smaller species, enabled a definitive assessment of size relationships between large mammalian carnivores and their ungulate prey. Five carnivore species were considered, including lion (Panthera leo), leopard (Panthera pardus), cheetah (Acinonyx jubatus), African wild dog (Lycaon pictus) and spotted hyena (Crocuta crocuta), and 22 herbivore prey species larger than 10 kg in adult body mass. 3. These carnivores selectively favoured prey species approximately half to twice their mass, within a total prey size range from an order of magnitude below to an order of magnitude above the body mass of the predator. The three smallest carnivores, i.e. leopard, cheetah and wild dog, showed high similarity in prey species favoured. Despite overlap in prey size range, each carnivore showed a distinct dietary preference. 4. Almost all mortality was through the agency of a predator for ungulate species up to the size of a giraffe (800-1200 kg). Ungulates larger than twice the mass of the predator contributed substantially to the dietary intake of lions, despite the low proportional mortality inflicted by predation on these species. Only for megaherbivores substantially exceeding 1000 kg in adult body mass did predation become a negligible cause of mortality. 5. Hence, the relative size of predators and prey had a pervasive structuring influence on biomass fluxes within this large-mammal food web. Nevertheless, the large carnivore assemblage was dominated overwhelmingly by the largest predator, which contributed the major share of animals killed across a wide size range.
TL;DR: It is suggested that guillemots specialize on a single foraging strategy regardless of whether chick-provisioning and self-feeding and the development of individual specialization may be an important precursor to diversification among seabirds.
Abstract: Summary 1. We studied chick diet in a known-age, sexed population of a long-lived seabird, the Brunnich’s guillemot ( Uria lomvia ), over 15 years ( N = 136; 1993‐2007) and attached time‐depth‐temperature recorders to examine foraging behaviour in multiple years ( N = 36; 2004‐07). 2. Adults showed specialization in prey fed to offspring, described by multiple indices calculated over 15 years: 27% of diet diversity was attributable to among-individual variation (withinindividual component of total niche width = 0·73); average similarity of an individual’s diet to the overall diet was 65% (mean proportional similarity between individuals and population = 0·65); diet was significantly more specialized than expected for 70% of individuals (mean likelihood = 0.53). These indices suggest higher specialization than the average for an across-taxa comparison of 49 taxa. 3. Foraging behaviour varied along three axes: flight time, dive depth and dive shape. Individuals showed specialized individual foraging behaviour along each axis. These foraging strategies were reflected in the prey type delivered to their offspring and were maintained over scales of hours to years. 4. Specialization in foraging behaviour and diet was greater over short time spans (hours, days) than over long time spans (years). Regardless of sex or age, the main component of variation in foraging behaviour and chick diet was between individuals. 5. Plasma stable isotope values were similar across years, within a given individual, and variance was low relative to that expected from prey isotope values, suggesting adult diet specialized across years. Stable isotope values were similar among individuals that fed their nestlings similar prey items and there was no difference in trophic level between adults and chicks. We suggest that guillemots specialize on a single foraging strategy regardless of whether chick-provisioning and self-feeding. With little individual difference in body mass and physiology, specialization likely represents learning and memorizing optimal feeding locations and behaviours. 6. There was no difference in survival or reproductive success between specialists and generalists, suggesting these are largely equivalent strategies in terms of evolutionary fitness, presumably because different strategies were advantageous at different levels of prey abundance or predictability. The development of individual specialization may be an important precursor to diversification among seabirds.
TL;DR: This energetic foraging tactic focused on a single or few prey items has not been observed previously in deep-diving mammals but resembles the high-risk/high-gain strategy of some terrestrial hunters such as cheetahs.
Abstract: Summary 1 Empirical testing of optimal foraging models for breath-hold divers has been difficult Here we report data from sound and movement recording DTags placed on 23 short-finned pilot whales off Tenerife to study the foraging strategies used to catch deep-water prey 2 Day and night foraging dives had a maximum depth and duration of 1018 m and 21 min Vocal behaviour during dives was consistent with biosonar-based foraging, with long series of echolocation clicks interspersed with buzzes Similar buzzes have been associated with prey capture attempts in other echolocating species 3 Foraging dives seemed to adapt to circadian rhythms Deep dives during the day were deeper, but contained fewer buzzes (median 1), than night-time deep dives (median 5 buzzes) 4 In most deep (540‐1019 m) daytime dives with buzzes, a downward directed sprint reaching up to 9 m s ‐1 occurred just prior to a buzz and coincided with the deepest point in the dive, suggestive of a chase after escaping prey 5 A large percentage (10‐36%) of the drag-related locomotion cost of these dives (15 min long) is spent in sprinting (19‐79 s) This energetic foraging tactic focused on a single or few prey items has not been observed previously in deep-diving mammals but resembles the high-risk/high-gain strategy of some terrestrial hunters such as cheetahs 6 Deep sprints contrast with the expectation that deep-diving mammals will swim at moderate speeds optimized to reduce oxygen consumption and maximize foraging time at depth Pilot whales may have developed this tactic to target a deep-water niche formed by large/calorific/fast moving prey such as giant squid
TL;DR: The especially high coral association and narrower niche breadth of juveniles suggest that the presence of live coral is crucial for many species during early life history, and that disturbance-induced coral loss may have serious flow-on effects on adult abundance.
Abstract: 1. The impact of environmental disturbance and habitat loss on associated species is expected to be dependent on a species' level of specialization. We examined habitat use and specialization of coral reef fish from the diverse and ecologically important family Pomacentridae, and determined which species are susceptible to declines in coral cover due to disturbance induced by crown-of-thorns seastar (COTS, Acanthaster planci L.). 2. A high proportion of pomacentrid species live in association with live coral as adults (40%) or juveniles (53%). Adults of many species had strong affiliations with branching corals, while juveniles favoured plating growth forms, reflecting the sizes of refuge provided by coral types. 3. Juveniles of species that associated with coral had narrower niche breadths than adult conspecifics, due to associations with specific coral types. The especially high coral association and narrower niche breadth of juveniles suggest that the presence of live coral is crucial for many species during early life history, and that disturbance-induced coral loss may have serious flow-on effects on adult abundance. 4. Microhabitat availability was a poor predictor of fish species abundance. Significant correlations between coverage of coral types and abundance of five adults and two juvenile species were detected; however, these relationships explained <35% and <10% of the variation in abundance of adult and juvenile species, respectively. 5. Niche breadth explained 74% of the variation in species' mean response to coral decline and it is clear that disturbance has a greater impact on resource specialists, suggesting that increasing frequency and intensity of coral loss will cause reef fish communities to become dominated by habitat generalists at the expense of coral-dwelling specialists.
TL;DR: This study is one of the few to demonstrate that the interaction between resource availability and infectious disease is important for shaping host population dynamics and emphasizes that multiple factors may drive oscillations in wild animal populations.
Abstract: 1. Populations of white-footed mice Peromyscus leucopus and deer mice Peromyscus maniculatus increase dramatically in response to food availability from oak acorn masts. These populations subsequently decline following this resource pulse, but these crashes cannot be explained solely by resource depletion, as food resources are still available as population crashes begin. 2. We hypothesized that intestinal parasites contribute to these post-mast crashes; Peromyscus are infected by many intestinal parasites that are often transmitted by density-dependent contact and can cause harm to their hosts. To test our hypothesis, we conducted a factorial experiment in natural populations by supplementing food to mimic a mast and by removal of intestinal nematodes with the drug, ivermectin. 3. Both food supplementation and the removal of intestinal nematodes lessened the rate and magnitude of the seasonal population declines as compared with control populations. However, the combination of food supplementation and removal of intestinal nematodes prevented seasonal population crashes entirely. 4. We also showed a direct effect on the condition of individuals. Faecal corticosterone levels, an indicator of the stress response, were significantly reduced in populations receiving both food supplementation and removal of intestinal nematodes. This effect was observed in autumn, before the overwinter crash observed in control populations, which may indicate that stress caused by the combination of food limitation and parasite infection is a physiological signal that predicts low winter survival and reproduction. 5. This study is one of the few to demonstrate that the interaction between resource availability and infectious disease is important for shaping host population dynamics and emphasizes that multiple factors may drive oscillations in wild animal populations.
TL;DR: It is suggested that climate variability strongly affects local elephant population dynamics through changes in surface-water availability, and views that a metapopulation framework should be endorsed for elephant management in open contexts are supported.
Abstract: Summary 1. An emerging perspective in the study of density dependence is the importance of the spatial and temporal heterogeneity of resources. Although this is well understood in temperate ungulates, few studies have been conducted in tropical environments where both food and water are limiting resources. 2. We studied the regulation of one of the world’s largest elephant populations in Hwange National Park, Zimbabwe. The study period started in 1986 when the population was released from culling. Using census data we investigated changes in elephant abundance with respect to rainfall and density across the entire park and across waterholes. 3. The population more than doubled since culling stopped. The population increased continuously during the first 6 years, and then fluctuated widely at about 30 000 individuals. Immigration processes must have been involved in the increase of the population size. 4. Population growth rates were negatively related to previous population density by a convex relationship, and negatively related to the ratio of previous population density on annual rainfall by a linear relationship. However, only this latter model (i.e. assuming a fluctuating carrying capacity related to annual rainfall) produced realistic dynamics. Overall, population decreased during dry years when the elephant density was high. 5. During dry years there were fewer waterholes retaining water during the dry season and consequently elephant numbers at waterholes increased, while their aggregation level across waterholes decreased. On the long-run elephant numbers increased only at the less crowded waterholes. 6. We suggest that the interaction between population size and the available foraging range determined by the number of active waterholes during the dry season controls the park population. 7. Our results emphasize the need to understand how key-resource areas cause resource-based aggregation, which ultimately influences the strength of density dependence. More specifically, this study suggests that climate variability strongly affects local elephant population dynamics through changes in surface-water availability. Finally, as dispersal is likely to be an important driver of the dynamics of this population, our results support views that a metapopulation framework should be endorsed for elephant management in open contexts.
TL;DR: It is suggested that habitat differences in nestling development or adaptive divergence of sparrow populations due to distinct environmental conditions (such as differing predation pressure) may account for the differences along the urbanization gradient.
Abstract: Summary 1 Urbanized habitats differ from natural ones in several ecological features, including climate, food availability, strength of predation and competition. Although the effects of urbanization on avian community composition are well known, there is much less information about how individual birds are affected by these human-generated habitat differences. 2 In this study we investigated the relationships between the morphological characteristics and the degree of habitat urbanization in house sparrows, Passer domesticus (Linne 1758) . We collected data for more than 1000 non-breeding adult birds in Hungary between 1997 and 2006, from seven sites including farmlands, suburban areas and city centres. 3 We found that the body mass, tarsus length and body condition of free-living sparrows differed among the sites: birds in more urbanized habitats were consistently smaller and in worse condition than birds in more rural habitats. A composite measure of habitat urbanization (based on building density, road density and vegetation cover) explained over 75% of variance between sites in the studied traits, after we controlled for the effects of sex, year, season and time of capture. 4 The difference in body mass between rural and urban sparrows was significant when birds were kept in aviaries under identical conditions, with constant ad libitum food availability. It is therefore unlikely that the reduced body size and condition of urban sparrows are a consequence of reduced access to food for adults (e.g. due to strong competition), or their short-term responses to high food predictability (e.g. by strategic mass regulation). 5 We suggest that habitat differences in nestling development or adaptive divergence of sparrow populations due to distinct environmental conditions (such as differing predation pressure) may account for the differences along the urbanization gradient.
TL;DR: This work estimated first-year, second-year and adult survival based on 43 years of ringing and dead recovery data from the Isle of May, Scotland, using recent methods to quantify temporal process variance and identify aspects of winter weather linked to survival.
Abstract: 1. Most scenarios for future climate change predict increased variability and thus increased frequency of extreme weather events. To predict impacts of climate change on wild populations, we need to understand whether this translates into increased variability in demographic parameters, which would lead to reduced population growth rates even without a change in mean parameter values. This requires robust estimates of temporal process variance, for example in survival, and identification of weather covariates linked to interannual variability. 2. The European shag Phalacrocorax aristotelis (L.) shows unusually large variability in population size, and large-scale mortality events have been linked to winter gales. We estimated first-year, second-year and adult survival based on 43 years of ringing and dead recovery data from the Isle of May, Scotland, using recent methods to quantify temporal process variance and identify aspects of winter weather linked to survival. 3. Survival was highly variable for all age groups, and for second-year and adult birds process variance declined strongly when the most extreme year was excluded. Survival in these age groups was low in winters with strong onshore winds and high rainfall. Variation in first-year survival was not related to winter weather, and process variance, although high, was less affected by extreme years. A stochastic population model showed that increasing process variance in survival would lead to reduced population growth rate and increasing probability of extinction. 4. As in other cormorants, shag plumage is only partially waterproof, presumably an adaptation to highly efficient underwater foraging. We speculate that this adaptation may make individuals vulnerable to rough winter weather, leading to boom-and-bust dynamics, where rapid population growth under favourable conditions allows recovery from periodic large-scale weather-related mortality. 5. Given that extreme weather events are predicted to become more frequent, species such as shags that are vulnerable to such events are likely to exhibit stronger reductions in population growth than would be expected from changes in mean climate. Vulnerability to extreme events thus needs to be accounted for when predicting the ecological impacts of climate change.
TL;DR: It is found that the current upper elevation limit to A. crataegi is below 900 m, and that the absence of host plants from high elevations sets the upper limit, which contrasts with the frequent assumption that biotic interactions typically determine warm range margins, and thermal limitation cool margins.
Abstract: 1. The ranges of many species have expanded in cool regions but contracted at warm margins in response to recent climate warming, but the mechanisms behind such changes remain unclear. Particular debate concerns the roles of direct climatic limitation vs. the effects of interacting species in explaining the location of low latitude or low elevation range margins. 2. The mountains of the Sierra de Guadarrama (central Spain) include both cool and warm range margins for the black-veined white butterfly, Aporia crataegi, which has disappeared from low elevations since the 1970s without colonizing the highest elevations. 3. We found that the current upper elevation limit to A. crataegi's distribution coincided closely with that of its host plants, but that the species was absent from elevations below 900 m, even where host plants were present. The density of A. crataegi per host plant increased with elevation, but overall abundance of the species declined at high elevations where host plants were rare. 4. The flight period of A. crataegi was later at higher elevations, meaning that butterflies in higher populations flew at hotter times of year; nevertheless, daytime temperatures for the month of peak flight decreased by 6.2 degrees C per 1 km increase in elevation. 5. At higher elevations A. crataegi eggs were laid on the south side of host plants (expected to correspond to hotter microclimates), whereas at lower sites the (cooler) north side of plants was selected. Field transplant experiments showed that egg survival increased with elevation. 6. Climatic limitation is the most likely explanation for the low elevation range margin of A. crataegi, whereas the absence of host plants from high elevations sets the upper limit. This contrasts with the frequent assumption that biotic interactions typically determine warm range margins, and thermal limitation cool margins. 7. Studies that have modelled distribution changes in response to climate change may have underestimated declines for many specialist species, because range contractions will be exacerbated by mismatch between the future distribution of suitable climate space and the availability of resources such as host plants.
TL;DR: Across 70 Neotropical bird species, the results suggest that, despite probably widespread differences in the intensity and diversity of pathogen exposure, species-level variation in constitutive immune defences is understandable within the context of life-history theory.
Abstract: 1. It has been suggested that immune defences are shaped by life history and ecology, but few general patterns have been described across species. We hypothesized that 'fast' life-history traits (e.g. short development times, large clutch sizes) would be associated with developmentally inexpensive immune defences, minimizing the resource demands of young animals' immune systems during periods of rapid growth. Conversely, 'slow' life histories should be associated with well developed antibody-mediated defences, which are developmentally costly. 2. We therefore predicted that 'fast-living' species would exhibit higher levels of complement proteins, a component of non-specific innate defence, but lower levels of constitutive ('natural') antibodies. Additionally, we predicted that constitutive immune defences in general would be higher in species with ecological characteristics that might increase exposure to pathogens, such as open nests, omnivorous diets, gregariousness, and closed forested habitat. 3. Across 70 Neotropical bird species, we found a strongly positive relationship between incubation period and natural antibody levels in adult birds, suggesting that longer developmental times might allow the production of a more diverse and/or more reactive adaptive immune system. Complement activity was positively, although weakly, correlated with clutch size, providing some support for the hypothesis that faster-living species rely more on innate defences, such as complement. Unexpectedly, solitary species had higher natural antibody titres than species that frequently join flocks. 4. Our results suggest that, despite probably widespread differences in the intensity and diversity of pathogen exposure, species-level variation in constitutive immune defences is understandable within the context of life-history theory.
TL;DR: Seasonal variation in Plasmodium infection in a blue tit Cyanistes caeruleus population over 3 years is examined and Beaudoin et al.'s model of seasonal variation in avian malaria prevalence in temperate areas is rejected as it does not survive closer scrutiny of the complexities of seasonal variations among Plas modium morphospecies and host age classes.
Abstract: 1. Seasonal variation in environmental conditions is ubiquitous and can affect the spread of infectious diseases. Understanding seasonal patterns of disease incidence can help to identify mechanisms, such as the demography of hosts and vectors, which influence parasite transmission dynamics. 2. We examined seasonal variation in Plasmodium infection in a blue tit Cyanistes caeruleus population over 3 years using sensitive molecular diagnostic techniques, in light of Beaudoin et al.'s (1971; Journal of Wildlife Diseases, 7, 5-13) model of seasonal variation in avian malaria prevalence in temperate areas. This model predicts a within-year bimodal pattern of spring and autumn peaks with a winter absence of infection. 3. Avian malaria infections were mostly Plasmodium (24.4%) with occasional Haemoproteus infections (0.8%). Statistical nonlinear smoothing techniques applied to longitudinal presence/absence data revealed marked temporal variation in Plasmodium prevalence, which apparently showed a within-year bimodal pattern similar to Beaudoin et al.'s model. However, of the two Plasmodium morphospecies accounting for most infections, only the seasonal pattern of Plasmodium circumflexum supported Beaudoin et al.'s model. On closer examination there was also considerable age structure in infection: Beaudoin et al.'s seasonal pattern was observed only in first year and not older birds. Plasmodium relictum prevalence was less seasonally variable. 4. For these two Plasmodium morphospecies, we reject Beaudoin et al.'s model as it does not survive closer scrutiny of the complexities of seasonal variation among Plasmodium morphospecies and host age classes. Studies of host-parasite interactions should consider seasonal variation whenever possible. We discuss the ecological and evolutionary implications of seasonal variation in disease prevalence.
TL;DR: A mathematical model is provided to predict the rate of decline in bat populations in the arid West in relation to climate change models for the region.
Abstract: 1. Climate change in North America is happening at an accelerated rate, reducing availability of water resources for bats and other wildlife that require it for successful reproduction. 2. We test the water-needy lactation hypotheses directly by tracking the drinking habitats of individual lactating and non-reproductive female fringed myotis at an artificial water source located near a maternity roost. 3. We used a submerged passive integrative transponder (PIT) tag reader system designed to track fish to instead record numbers of water source visitations by tagged bats. 4. Of 24 PIT-tagged adult females, 16 (67%) were detected repeatedly by the plate antenna as they passed to drink between 18 July and 28 August 2006. 5. The total number of drinking passes by lactating females (n = 255) were significantly higher than those of non-reproductive adult females (n = 22). Overall, lactating females visited 13 times more often to drink water than did non-reproductive females. On average, lactating females visited six times more often per night. Drinking bouts occurred most frequently just after evening emergence and at dawn. 6. Drinking patterns of non-reproductive females correlated significantly with fluctuating ambient temperature and relative humidity recorded at the water source, whereas lactating females drank extensively regardless of ambient conditions. 7. We provide a mathematical model to predict the rate of decline in bat populations in the arid West in relation to climate change models for the region.
TL;DR: The longevity and per capita fecundity of naturally occurring Diadegma insulare foraging in cabbage plots with and without borders of flowering buckwheat, Fagopyrum esculentum, as well as relationships between longevity, fecundy, sugar feeding and parasitism rates on larvae of the diamondback moth, Plutella xylostella are studied.
Abstract: Summary 1. The use of floral resource subsidies to improve herbivore suppression by parasitoids requires certain trophic interactions and physiological changes to occur. While the longevity and fecundity of parasitoids are positively affected by nectar subsidies in laboratory studies, the impacts of floral subsidies on the fecundity and longevity of freely foraging parasitoids have not been studied. 2. We studied the longevity and per capita fecundity of naturally occurring Diadegma insulare foraging in cabbage plots with and without borders of flowering buckwheat, Fagopyrum esculentum, as well as relationships between longevity, fecundity, sugar feeding and parasitism rates on larvae of the diamondback moth, Plutella xylostella . 3. Relative longevity was estimated by counting broken setae on the fringe of the forewing. Floral borders increased the longevity of males and females in adjacent cabbage plots. 4. The egg maturation rate of D. insulare was estimated by comparing egg loads of females collected early in the day with egg loads of females held without hosts in field cages throughout the day. Females in buckwheat cages matured 2·7 eggs per hour while females in control cages resorbed 0·27 eggs over the same time period. 5. The fecundity of females collected in the afternoon was estimated by comparing their actual egg load to the estimated egg load in the absence of oviposition for females in a given plot. Females foraging in buckwheat plots had marginally fewer eggs remaining in their ovaries, and laid marginally more eggs than females in control plots. Females from both treatments carried 30‐60 eggs by the afternoon and therefore were time-limited rather than egg-limited. 6. Plots where a greater proportion of females had fed on sugar had longer-lived females. This suggests that feeding enhanced longevity of D. insulare . However, plots with longer-lived and more fecund females did not exhibit higher parasitism rates, although the power of these tests were low.
TL;DR: It was concluded that indirect effects of aphid community structure are most likely to be mediated by predators, a prediction supported by the available experimental evidence.
Abstract: 1. Most communities of insect herbivores are unlikely to be structured by resource competition, but they may be structured by apparent competition mediated by shared natural enemies. 2. The potential of three guilds of natural enemies (parasitoids, fungal entomopathogens and predators) to influence aphid community structure through indirect interactions is assessed. Based on the biology, we predicted that the scope for apparent competition would be greatest for the predator and least for the parasitoid guilds. 3. Separate fully quantitative food webs were constructed for 3 years for the parasitoid guild, 2 years for the pathogen guild and for a single year for the predator guild. The webs were analysed using standard food web statistics designed for binary data, and using information-theory-based metrics that make use of the full quantitative data. 4. A total of 29 aphid, 24 parasitoid, five entomopathogenic fungi and 13 aphid specialist predator species were recorded in the study. Aphid density varied among years, and two species of aphid were particularly common in different years. Omitting these species, aphid diversity was similar among years. 5. The parasitoid web showed the lowest connectance while standard food web statistics suggested the pathogen and predator webs had similar levels of connectance. However, when a measure based on quantitative data was used the pathogen web was intermediate between the other two guilds. 6. There is evidence that a single aphid species had a particularly large effect on the structure of the pathogen food web. 7. The predator and pathogen webs were not compartmentalized, and the vast majority of parasitoids were connected in a single large compartment. 8. It was concluded that indirect effects are most likely to be mediated by predators, a prediction supported by the available experimental evidence.
TL;DR: Evidence is found for a largely non-saturating relationship (relative to random expectation) between species richness and functional evenness, which suggests that increased niche specialization may have allowed species to coexist in the most species-rich communities, and indicates that the conservation of functional diversity may be vital for the maintenance of species diversity in biological communities.
Abstract: 1. Interspecific niche differences have long been identified as a major explanation for the occurrence of species-rich communities. However, much fieldwork studying variation in local species richness has focused upon physical habitat attributes or regional factors, such as the size of the regional species pool. 2. We applied indices of functional diversity and niche overlap to data on the species niche to examine the importance of interspecific niche differentiation for species richness in French lake fish communities. We combined this information with environmental data to test generalizations of the physiological tolerance and niche specialization hypotheses for species-energy relationships. 3. We found evidence for a largely non-saturating relationship (relative to random expectation) between species richness and functional evenness (evenness of spacing between species in niche space), while functional richness (volume of niche space occupied) peaked at moderate levels of species richness and niche overlap showed an initial decrease followed by saturation. This suggests that increased niche specialization may have allowed species to coexist in the most species-rich communities. 4. We tested for evidence that increased temperature, local habitat area, local habitat diversity and immigration affected species richness via increased niche specialization. Temperature explained by far the largest amount of variation in species richness, functional diversity and niche overlap. These results, combined with the largely non-saturating species richness-functional evenness relationship, suggest that increased temperature may have permitted increased species richness by allowing increased niche specialization. 5. These results emphasize the importance of niche differences for species coexistence in species-rich communities, and indicate that the conservation of functional diversity may be vital for the maintenance of species diversity in biological communities. Our approach may be applied readily to many types of community, and at any scale, thus providing a flexible means of testing niche-based hypotheses for species richness gradients.
TL;DR: Surprisingly, contrary to predictions of metabolic models, this suggests that for any prey species, the per link and total energy fluxes to its largest predators are smaller than those to predators of intermediate body size.
Abstract: 1. In natural communities, populations are linked by feeding interactions that make up complex food webs. The stability of these complex networks is critically dependent on the distribution of energy fluxes across these feeding links. 2. In laboratory experiments with predatory beetles and spiders, we studied the allometric scaling (body-mass dependence) of metabolism and per capita consumption at the level of predator individuals and per link energy fluxes at the level of feeding links. 3. Despite clear power-law scaling of the metabolic and per capita consumption rates with predator body mass, the per link predation rates on individual prey followed hump-shaped relationships with the predator-prey body mass ratios. These results contrast with the current metabolic paradigm, and find better support in foraging theory. 4. This suggests that per link energy fluxes from prey populations to predator individuals peak at intermediate body mass ratios, and total energy fluxes from prey to predator populations decrease monotonically with predator and prey mass. Surprisingly, contrary to predictions of metabolic models, this suggests that for any prey species, the per link and total energy fluxes to its largest predators are smaller than those to predators of intermediate body size. 5. An integration of metabolic and foraging theory may enable a quantitative and predictive understanding of energy flux distributions in natural food webs.
TL;DR: Relationships between rainfall and changes in age- and sex-structured abundances of seven ungulate species monitored monthly for 15 years using vehicle ground counts in the Maasai Mara National Reserve suggest that rainfall underpins the dynamics of African savanna ungulates, and that changes in rainfall due to global warming may markedly alter the abundance and diversity of these mammals.
Abstract: 1. Rainfall is the prime climatic factor underpinning the dynamics of African savanna ungulates, but no study has analysed its influence on the abundance of these ungulates at monthly to multiannual time scales. 2. We report relationships between rainfall and changes in age- and sex-structured abundances of seven ungulate species monitored monthly for 15 years using vehicle ground counts in the Maasai Mara National Reserve, Kenya. 3. Abundance showed strong and curvilinear relationships with current and cumulative rainfall, with older topi, Damaliscus korrigum (Ogilby); warthog, Phacochoerus aethiopicus (Pallas); waterbuck, Kobus ellipsyprimnus (Ogilby); and impala, Aepyceros melampus (Lichtenstein) responding to longer lags than younger animals, portraying carryover effects of prior habitat conditions. 4. The abundances of newborn calves were best correlated with monthly rainfall averaged over the preceding 5-6 months for topi, waterbuck, warthog, and 2 months for the migratory zebra Equus burchelli (Gray), but with seasonal rainfall averaged over 2-5 years for giraffe, Giraffa camelopardalis (L.); impala; and kongoni, Alcelaphus busephalus (Pallas). The cumulative late wet-season rainfall was the best predictor of abundance for quarter- to full-grown animals for most species. Monthly rainfall exerted both negative and positive effects on the abundances of zebra, impala and waterbuck. Ignoring age, both sexes responded similarly to rainfall. 5. Births were strongly seasonal only for warthog and topi, but peaked between August and December for most species. Hence abundance was strongly seasonal for young topi and warthog and the migratory zebra. Pronounced seasonality in births for warthog and topi obliterated otherwise strong relationships between abundance and rainfall when both month and rainfall were included in the same model. Aggregated density produced relationships with rainfall similar to those for fully grown animals, emphasizing the necessity of demographic monitoring to reliably reveal rainfall influences on ungulate abundance in the Mara. 6. Strong relationships between abundance and rainfall suggest that rainfall underpins the dynamics of African savanna ungulates, and that changes in rainfall due to global warming may markedly alter the abundance and diversity of these mammals. Ungulates respond to rainfall fluctuations through movements, reproduction or survival, and the responses appear independent of breeding phenology and synchrony, dietary guild, or degree of water dependence. Newborns and adults have contrasting responses to rainfall. Males and females respond similarly to rainfall when age is ignored.
TL;DR: It is suggested that selection for body weight and competitive ability will be high in female meerkats, which may moderate their investment in cooperative activities.
Abstract: 1. In cooperative societies with high reproductive skew, selection on females is likely to operate principally through variation in the probability of acquiring dominant status and variation in reproductive success while dominant. Despite this, few studies of cooperative societies have investigated the factors that influence which females become dominant, and/or their reproductive output while in the dominant position. 2. Here we use long-term data from a wild meerkats population to describe variation in the breeding success of dominant female meerkats Suricata suricatta and investigate its causes. 3. Female meerkats compete intensely for breeding positions, and the probability of acquiring the breeding role depends upon a female's age in relation to competitors and her weight, both at the time of dominance acquisition and early in life. 4. Once dominant, individual differences in breeding success depend principally on the duration of dominance tenure. Females remain for longer in the dominant position if they are heavier than their competitors at the start of dominance, and if the number of adult female competitors at the start is low. 5. Female breeding success is also affected by variation in fecundity and pup survival, both of which increase with group size. After controlling for these effects, female body weight has a positive influence on breeding rate and litter size, while the number of adult female competitors reduces litter survival. 6. These findings suggest that selection for body weight and competitive ability will be high in female meerkats, which may moderate their investment in cooperative activities. We suggest that similar consequences of competition may occur among females in other cooperative societies where the benefits of attaining dominance status are high.
TL;DR: A long-term data set of individually identifiable common guillemots, Uria aalge Pont.
Abstract: 1. In long-lived animals with delayed maturity, the non-breeding component of the population may play an important role in buffering the effects of stochastic mortality. Populations of colonial seabirds often consist of more than 50% non-breeders, yet because they spend much of their early life at sea, we understand little about their impact on the demographic process. 2. Using multistate capture-mark-recapture techniques, we analyse a long-term data set of individually identifiable common guillemots, Uria aalge Pont., to assess factors influencing their immature survival and two-stage recruitment process. 3. Analysis of the distribution of ringed common guillemots during the non-breeding season, separated by age classes, revealed that all age classes were potentially at risk from four major oil spills. However, the youngest age class (0-3 years) were far more widely spread than birds 4-6 years old, which were more widely spread than birds aged 6 and over. Therefore the chance of encountering an oil spill was age-dependent. 4. A 2-year compound survival estimate for juvenile guillemots was weakly negatively correlated with winter sea-surface temperature, but was not influenced by oil spills. Non-breeder survival did not vary significantly over time. 5. In years following four oil spills, juvenile recruitment was almost double the value in non-oil-spill years. Recent work from Skomer Island showed a doubling of adult mortality associated with major oil spills, which probably reduced competition at the breeding colony, allowing increased immature recruitment to compensate for these losses. We discuss the implications of compensatory recruitment for assessing the impact of oil pollution incidents.
TL;DR: Using data from a 60-year study period, a detailed description of how the northern goshawk Accipiter gentilis L.--generally considered a shy forest species--colonized the city of Hamburg, Germany is provided.
Abstract: 1. Despite the accelerating global spread of urbanized habitats and its associated implications for wildlife and humans, surprisingly little is known about the biology of urban ecosystems. 2. Using data from a 60-year study period, this paper provides a detailed description of how the northern goshawk Accipiter gentilis L.--generally considered a shy forest species--colonized the city of Hamburg, Germany. Six non-mutually exclusive hypotheses are investigated regarding the environmental factors that may have triggered this invasion. 3. The spatio-temporal analysis of 2556 goshawk chance observations (extracted from a total data set of 1 174 493 bird observations; 1946-2003) showed that hawks regularly visited the city centre decades before the first successful breeding attempts were recorded. Many observations were made in parts of the city where territories were established in later years, demonstrating that these early visitors had encountered, but not used, potential nest sites. 4. Pioneer settlement coincided with: (i) an increase in (legal) hunting pressure on goshawks in nearby rural areas; (ii) an increase in avian prey abundance in the city; and (iii) a succession of severe winters in the Greater Hamburg area. On the other hand, there was no evidence to suggest that the early stages of the invasion were due to: (i) decreasing food availability in rural areas; (ii) major habitat changes in the city; or (iii) rural intraguild dynamics forcing hawks into urban refugia. While breeding numbers of a potential rural source population were at a long-term low when the city was colonized, prior to first settlement there was a sharp increase of goshawk chance observations in the city and its rural periphery. 5. The urban population expanded rapidly, and pair numbers began to stabilize after about 10 years. Ringing data (219 ringed nestlings from 70 urban broods; 1996-2000) demonstrated that most urban recruits had fledged in the city, but also confirmed considerable gene flow between urban and rural habitats. Analysis of chance observations (as raw data or as detrended time series) suggested a tight coupling of population dynamics inside and outside the city. 6. City-colonizations such as the one described here provide a valuable opportunity to study some fundamental aspects of population ecology on a scale at which detailed monitoring is logistically feasible. Furthermore, a good understanding of urban ecology has become essential for efficient wildlife conservation in modern, human-altered environments.
TL;DR: Using a laboratory model system, the soil mite Sancassania berlesei is shown that, controlling for egg size, older mothers lay eggs that hatch later, develop more slowly, and mature at larger body sizes.
Abstract: 1. Maternal effects describe how mothers influence offspring life histories. In many taxa, maternal effects arise by differential resource allocation to young, often identified by variation in propagule size, and which affects individual traits and population dynamics. 2. Using a laboratory model system, the soil mite Sancassania berlesei, we show that, controlling for egg size, older mothers lay eggs that hatch later, develop more slowly, and mature at larger body sizes. 3. Such differences in life histories lead to marked population dynamical effects lasting for multiple generations, as evidenced by an experiment initiated with similarly sized eggs that came from young or old mothers. Differences in maturation from the initial cohort led to differences in population structure and life history that propagated the initial differences over time. 4. Maternal-age effects, which are not related to gross provisioning of the egg and are therefore phenotypically cryptic, can have profound implications for population dynamics, especially if environmental variation can affect the age structure of the adult population.
TL;DR: Five structural models predict the detailed structure of complex food webs given only two input parameters, the numbers of species and the number of feeding links among them but differ in the extent to which species' diets are required to be contiguous and the rules used to assign feeding links.
Abstract: 1. Following the development of the relatively successful niche model, several other simple structural food web models have been proposed. These models predict the detailed structure of complex food webs given only two input parameters, the numbers of species and the number of feeding links among them. 2. The models claim different degrees of success but have not been compared consistently with each other or with the empirical data. We compared the performance of five structural models rigorously against 10 empirical food webs from a variety of aquatic and terrestrial habitats containing 25-92 species and 68-997 links. 3. All models include near-hierarchical ordering of species' consumption and have identical distributions of the number of prey of each consumer species, but differ in the extent to which species' diets are required to be contiguous and the rules used to assign feeding links. 4. The models perform similarly on a range of food-web properties, including the fraction of top, intermediate and basal species, the standard deviations of generality and connectivity and the fraction of herbivores and omnivores. 5. For other properties, including the standard deviation of vulnerability, the fraction of cannibals and species in loops, mean trophic level, path length, clustering coefficient, maximum similarity and diet discontinuity, there are significant differences in the performance of the different models. 6. While the empirical data do not support the niche model's assumption of diet contiguity, models which relax this assumption all have worse overall performance than the niche model. All the models under-estimate severely the fraction of species that are herbivores and exhibit other important failures that need to be addressed in future research.
TL;DR: It is demonstrated that host ecology importantly affects parasite virulence, with implications for host-parasite dynamics in natural populations, and the virulence rank order of parasite genotypes was unaffected by host plant species.
Abstract: Summary 1. Studies have considered how intrinsic host and parasite properties determine parasite virulence, but have largely ignored the role of extrinsic ecological factors in its expression. 2. We studied how parasite genotype and host plant species interact to determine virulence of the protozoan parasite Ophryocystis elektroscirrha (McLaughlin & Myers 1970) in the monarch butterfly Danaus plexippus L. We infected monarch larvae with one of four parasite genotypes and reared them on two milkweed species that differed in their levels of cardenolides: toxic chemicals involved in predator defence. 3. Parasite infection, replication and virulence were affected strongly by host plant species. While uninfected monarchs lived equally long on both plant species, infected monarchs suffered a greater reduction in their life spans (55% vs. 30%) on the low-cardenolide vs. the high-cardenolide host plant. These life span differences resulted from different levels of parasite replication in monarchs reared on the two plant species. 4. The virulence rank order of parasite genotypes was unaffected by host plant species, suggesting that host plant species affected parasite genotypes similarly, rather than through complex plant species‐parasite genotype interactions. 5. Our results demonstrate that host ecology importantly affects parasite virulence, with implications for host‐parasite dynamics in natural populations.
TL;DR: A simulation approach was used to assess the magnitude of fitness underestimation expected for dispersing individuals depending on the heritability of dispersal distance and study area size, and to list three potential ways of reducing the risk of making wrong inferences on biased fitness estimates.
Abstract: 1. Many studies investigating fitness correlates of dispersal in vertebrates report dispersers to have lower fitness than philopatric individuals. However, if dispersers are more likely to produce dispersing young or are more likely to disperse again in the next year(s) than philopatric individuals, there is a risk that fitness estimates based on local adult survival and local recruitment will be underestimated for dispersers. 2. We review the available empirical evidence on parent-offspring resemblance and individual lifelong consistency in dispersal behaviour, and relate these studies to recent studies of fitness correlates of dispersal in vertebrates. 3. Of the 12 studies testing directly for parent-offspring resemblance in dispersal propensity, five report a significant resemblance. The average effect size (r) of parent-offspring resemblance in dispersal was 0.15 [95% confidence interval (CI) = 0.07-0.22], with no difference between the sexes (average weighted effect size of 0.12 (0.08-0.16) and 0.16 (0.11-0.20) for females and males, respectively). Only three studies report data on within-individual consistency in dispersal propensity, of which two suggest dispersers to be more likely to disperse again. 4. To assess the magnitude of fitness underestimation expected for dispersing individuals depending on the heritability of dispersal distance and study area size, we used a simulation approach. Even when study area size is 10 times the mean dispersal distance, local recruitment per breeding event may be underestimated by 4-10%, generating a potential difference of 4-60% in average lifetime production of recruits between dispersing and philopatric individuals, with larger differences in long-lived species. 5. Estimates of both fitness correlates of dispersal and parent-offspring resemblance or within-individual consistency in dispersal behaviour have been reported for 11 species. Although some comparisons suggest genuine differences in fitness components between philopatric and dispersing individuals, others, based on adult and juvenile survival, are open to the alternative explanation of biased fitness estimates. 6. We list three potential ways of reducing the risk of making wrong inferences on biased fitness estimates due to such non-random dispersal behaviour between dispersing and philopatric individuals: (a) diagnosing effects of non-random dispersal, (b) reducing the effects of spatially limited study area and (c) performing controlled experiments.
TL;DR: The results emphasize the potential importance of detritivore diversity for stream ecosystem functioning, but both the effects of diversity on the studied processes, and the mechanisms underlying those effects, were specific to each assemblage and process.
Abstract: 1. The diversity of species traits in a biological assemblage varies not only with species richness, but also with species evenness and organism density, which together influence the concentration of traits within functional guilds. Potential trait diversity at local scales is also constrained by the regional species pool. Implications of such variation for spatio-temporal variability in biodiversity-ecosystem functioning relationships are likely to be complex, but are poorly understood. 2. In microcosm experiments conducted at laboratories in Sweden, Ireland and Romania, we investigated effects of species richness, evenness and density of stream-living detritivores on two related processes: detritivore leaf-processing efficiency (LPE) and growth. Assemblage composition varied among laboratories: one taxonomic order (Plecoptera) was studied in Sweden, whereas two orders, encompassing wider trait variation, were studied in Romania (Trichoptera and Plecoptera) and Ireland (Trichoptera and Isopoda). 3. Relationships between density and both LPE and growth ranged from negative to positive across the study species, highlighting the potential for density-dependent variation in process rates to alter ecosystem functioning, but indicating that such effects depend on species identity. 4. LPE varied with species diversity in the two more heterogeneous assemblages, but whereas LPE in the Romanian study was generally enhanced as richness increased, LPE in the Irish study increased only in less-even polycultures dominated by particular species. Transgressive overyielding was detected in the Irish experiment, indicating complementary resource use and/or facilitation (complementarity). These mechanisms could not be distinguished from the selection effect in the Romanian study. 5. Growth was elevated in Romanian species mixtures, reflecting positive complementarity, but lower than expected growth in some Swedish mixtures was associated with negative complementarity, indicating interspecific interference competition. 6. Our results emphasize the potential importance of detritivore diversity for stream ecosystem functioning, but both the effects of diversity on the studied processes, and the mechanisms underlying those effects, were specific to each assemblage and process. Such variability highlights challenges in generalizing impacts of diversity change for functional integrity in streams and other ecosystems in which the occurrence of important species traits fluctuates over relatively small spatio-temporal scales.
TL;DR: It is proposed that including these three surrogates of condition will substantially improve the accuracy of non-intrusive estimates of body condition, thus providing more powerful tools with direct application in a wide range of disciplines.
Abstract: 1. Body condition (defined as the relative amount of energy reserves in the body) is an animal trait with strong ecological implications. In some animal taxa (e.g. arthropods), the external volume of the body part in which most nutrients are stored (e.g. abdomen) is used interchangeably with body mass to estimate body condition, making the implicit assumption that abdomen residual volume is a good surrogate of residual mass. However, the degree of correlation between these two measures should largely depend on the density of the nutrients stored. 2. We simulated two food-supplemented experimental groups of animals, each storing a slightly different amount of lipids either in their abdomens or in their entire bodies, and explored (i) how different estimates of condition were able to detect fixed differences between the groups; and (ii) how the amount of lipids stored could affect the outcome of non-intrusive measures of condition on a dichotomous variable (e.g. survival, mating success). We found that density body condition (body mass statistically controlled for structural body size and body volume) has much greater power to detect differences between experimental groups or effects on binary response variables than do classic mass/size or volume/size condition indices. 3. Using data on Lycosa tarantula (L.), a burrowing wolf spider, we report dramatic differences among these three indices in their ability to detect sex differences in the effect of feeding treatment on body condition at maturity. In particular, a plot of residual mass against residual volume reflecting nutrient density suggests that poorly fed spiders are nutritionally unbalanced, since well-fed spiders invest in nutrients of very different density. 4. Furthermore, using data on Scathophaga stercoraria (L.), the yellow dung fly, we found that an index of density condition was better at distinguishing condition differences among three populations than were mass or volume condition estimates alone. 5. We propose that including these three surrogates of condition (mass, volume and density) will substantially improve the accuracy of non-intrusive estimates of body condition, thus providing more powerful tools with direct application in a wide range of disciplines.