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Journal: The Journal of animal ecology


1. Ecologists have traditionally focused on herbivore carcasses as study models in scavenging research. However, some observations of scavengers avoiding feeding on carnivore carrion suggest that different types of carrion may lead to differential pressures. Untested assumptions about carrion produced at different trophic levels could therefore lead ecologists to overlook important evolutionary processes and their ecological consequences. 2. Our general goal was to investigate the use of mammalian carnivore carrion by vertebrate scavengers. In particular, we aimed to test the hypothesis that carnivore carcasses are avoided by other carnivores, especially at the intra-specific level, most likely to reduce exposure to parasitism. 3. We take a three-pronged approach to study this principle by: i) providing data from field experiments, ii) carrying out evolutionary simulations of carnivore scavenging strategies under risks of parasitic infection, and iii) conducting a literature-review to test two hypotheses regarding parasite life-history strategies. 4. First, our field experiments showed that the mean number of species observed feeding at carcasses and the percentage of consumed carrion biomass were substantially higher at herbivore carcasses than at carnivore carcasses. This occurred even though the number of scavenger species visiting carcasses and the time needed by scavengers to detect carcasses were similar between both types of carcasses. In addition, we did not observe cannibalism. Second, our evolutionary simulations demonstrated that a risk of parasite transmission leads to the evolution of scavengers with generally low cannibalistic tendencies, and that the emergence of cannibalism-avoidance behavior depends strongly on assumptions about parasite-based mortality rates. Third, our literature review indicated that parasite species potentially able to follow a carnivore-carnivore indirect cycle, as well as those transmitted via meat consumption, are rare in our study system. 5. Our findings support the existence of a novel coevolutionary relation between carnivores and their parasites, and suggest that carnivore and herbivore carcasses play very different roles in food webs and ecosystems. This article is protected by copyright. All rights reserved.

Concepts: Scientific method, Ecology, Parasitism, Observation, Hypothesis, Omnivore, Scavenger, Carnivore


Natural river floodplains are among Earth’s most biologically diverse and productive ecosystems but face a range of critical threats due to human disturbance. Understanding the ecological processes that support biodiversity and productivity in floodplain rivers is essential for their future protection and rehabilitation. Fish assemblage structure on tropical river floodplains is widely considered to be driven by dispersal limitation during the wet season and by environmental filtering and inter-specific interactions during the dry season. However, the individual-level movement behaviours (e.g., site attachment, nomadism, homing) that regulate dispersal of fish on floodplains are poorly understood. We combined radio-telemetry and remote sensing to examine the movement behaviour of two large-bodied fishes (barramundi Lates calcarifer, forktail catfish Neoarius leptaspis) over the flood cycle in a tropical river-floodplain system in northern Australia to: 1) quantify movement responses in relation to dynamic habitat resources at a landscape scale; and 2) determine the extent of spatial ‘reshuffling’ of individual fish following the wet season. Both species altered their behaviour rapidly in response to changes in the availability and distribution of aquatic habitat, with most individuals undertaking extensive movements (up to ~ 27 km from the tagging location) on the inundated floodplain during the wet season. Although there was considerable individual variation in movement patterns, overall barramundi distributions closely tracked the extent of floodplain primary productivity, whereas catfish distributions were most closely associated with the extent of flooded area. Most individuals of both species exhibited homing back to previously occupied dry season refugia during the wet-to-dry transition, even though other potential refugia were available in closer proximity to wet season activity areas. We postulate that homing behaviour modulates temporal variation in fish assemblage composition and abundance and limits the transfer of aquatic-derived energy and nutrients into terrestrial food webs by reducing fish mortality on drying floodplains. Our study demonstrates the importance of quantifying individual-level behaviour across the three stages of dispersal (emigration, inter-patch movement, immigration) for our understanding of how animal movement influences energetic subsidies and other large-scale ecosystem processes.


In biological invasions, rates of range expansion tend to accelerate through time. What kind of benefits to more rapidly dispersing organisms might impose natural selection for faster rates of dispersal, and hence the evolution of range-edge acceleration? We can answer that question by comparing fitness-relevant ecological traits of individuals at the invasion front compared with conspecifics in the same area a few years post-invasion. In tropical Australia, the rate of invasion by cane toads (Rhinella marina) has increased substantially over recent decades, due to shifts in heritable traits. Our data on field-collected cane toads at a recently invaded site in the Australian wet-dry tropics span a 5-year period beginning with toad arrival. Compared with conspecifics that we monitored in the same sites post-invasion, toads in the invasion vanguard exhibited higher feeding rates, larger energy stores, better body condition and faster growth. Three processes may have contributed to this pattern: (i) higher prey availability at the front (perhaps due to reduced competition from conspecifics); (ii) the lack of viability-reducing parasites and pathogens in invasion-front toads; and (iii) distinctive (active, fast-growing) phenotypes of the invasion-front toads. Nutritional benefits to individuals in the invasion vanguard (whether because of higher prey availability, or lower pathogen levels) thus may have conferred a selective advantage to accelerated dispersal in this system.

Concepts: Natural selection, Evolution, Biology, Acceleration, Invasion, Toads, Invasions, Dwayne Johnson


Warning signals are expected to evolve towards conspicuousness and monomorphism, and thereby hamper the evolution of multiple colour morphs. Here, we test fitness responses to different rearing densities to explain colour polymorphism in aposematic wood tiger moth (Parasemia plantaginis) males. We used larval lines sired by white or yellow adult males selected for small or large melanization patterns of coloration. We reared these selected lines either solitarily (favourable conditions) or in aggregations (challenged conditions), and followed their performance to adult stage. We tested whether differences in larval density affected life-history traits, adult melanin expression, adult morph (white or yellow) survival and immunological responses. We found that the aggregated environment increased mortality of larvae, but decreased larval developmental time and pupa weight. Adult wing melanin pigmentation was dependent on larval melanin expression but not rearing density. We also confirmed that adult wing coloration had a genetic basis (h(2)  = 0·42) and was not influenced by larval growth density. Adult yellow males survived better from aggregations in comparison with white males, which may be related to differences in immune defence. White males had better encapsulation ability, whereas yellow males had increased lytic activity of haemolymph in the aggregations. Our main results highlight, that morph-linked immune responses mediated by differential growth density may facilitate the maintenance of colour polymorphism in aposematic species. In nature, risk of diseases and parasites vary spatially and temporally. Therefore, both yellow and white adult morphs can be maintained due to their differential investment in immune defence in heterogeneous environments.

Concepts: Immune system, Insect, Color, White, Lepidoptera, Arctiidae, Parasemia plantaginis, Parasemia


The manner in which patterns of variation and interactions among demographic rates contribute to population growth rate (λ) are key to understanding how animal populations will respond to changing climatic conditions. Migratory species are likely to be particularly sensitive to climatic conditions as they experience a range of different environments throughout their annual cycle. However, few studies have provided fully integrated demographic analyses of migratory populations in response to changing climatic conditions. Here, we employed integrated population models (IPM) to demonstrate that the environmental conditions experienced during a short, but critical period, play a central role in the demography of a long-distance migrant, the light-bellied Brent goose (Branta bernicla hrota). Female survival was positively associated with June North Atlantic Oscillation (NAO) values, whereas male survival was not. In contrast, breeding productivity was negatively associated with June NAO, suggesting a trade-off between female survival and reproductive success. Both adult female and adult male survival showed low temporal variation, whereas there was high temporal variation in recruitment and breeding productivity. In addition, while annual population growth was positively correlated with annual breeding productivity a sensitivity analysis revealed that population growth was most sensitive to changes in adult survival. Our results demonstrate that the environmental conditions experienced during a relatively short-time window at the start of the breeding season play a critical role in shaping the demography of a long-distant Arctic migrant. Crucially, different demographic rates responded in opposing directions to climatic variation, emphasizing the need for integrated analysis of multiple demographic traits when understanding population dynamics. This article is protected by copyright. All rights reserved.

Concepts: Reproduction, Demography, Population, Population ecology, Demographic economics, World population, Carrying capacity, Population growth


The timing of bird migration has shifted in response to climate change. However, few studies have linked the potential consequences of any phenological shift on individual fitness and even fewer have disentangled the role of plasticity and microevolution in the observed shifts. The arrival date and breeding success of white storks (Ciconia ciconia) have been recorded since the 1880s in Slovakia. We used data for two periods (1895-1913 and 1977-2007), which were considered, respectively, as populations before and after the start of climate warming. About 4000 male and 2500 female arrival dates along with 3000 breeding attempts were studied. Mean arrival dates did not differ between the two periods. During 1977-2007, males tended towards a slight delay for most fractions of arrival distribution. Protandry was reduced by 30% (1·44 days). In both sexes, the early percentiles of the arrival distribution arrived later those years with warmer temperatures at the African wintering grounds, while late percentiles advanced their arrival when temperatures were higher in the European areas flown over during migration. Mean breeding success of the Slovakian population has not changed since 1977. However, fecundity selection for arrival date reduced over the years: at the end of 1970s and 1980s, early breeders had more success than late breeders, but this seasonal trend disappeared towards the end of the study period. An early arrival and territory acquisition may have become less of an advantage due to the enhancement of feeding opportunities during the breeding season in recent decades. A century ago, stork arrival varied spatially, with earlier arrivals at low altitudes, southern slopes and warmer and drier regions. This spatial variation mostly vanished, and at present, we found little correlations with topographical and climatic gradients. We showed that long-term temporal changes in the timing of biological events may be complex because each fraction of a population and sex may show different temporal trends in their arrival dates. In addition, the effect of biotic and abiotic factors may change consistently in space and time, and thereby phenotypes' value depends on the circumstances that are expressed due to its variable fitness consequences.

Concepts: Bird, Sex, Climate change, Bird migration, Stork, White Stork, Ciconia, Pfeilstorch


Terrestrial food webs are woven from complex interactions, often underpinned by plant-mediated interactions between herbivores and higher trophic groups. Below- and above-ground herbivores can influence one another via induced changes to a shared host plant, potentially shaping the wider community. However, empirical evidence linking laboratory observations to natural field populations has so far been elusive. This study investigated how root-feeding weevils (Otiorhynchus sulcatus) influence different feeding guilds of herbivore (phloem-feeding aphids, Cryptomyzus galeopsidis, and leaf-chewing sawflies, Nematus olfaciens) in both controlled and field conditions. We hypothesized that root herbivore-induced changes in plant nutrients (C, N, P and amino acids) and defensive compounds (phenolics) would underpin the interactions between root and foliar herbivores, and ultimately populations of natural enemies of the foliar herbivores in the field. Weevils increased field populations of aphids by ca. 700%, which was followed by an increase in the abundance of aphid natural enemies. Weevils increased the proportion of foliar essential amino acids, and this change was positively correlated with aphid abundance, which increased by 90% on plants with weevils in controlled experiments. In contrast, sawfly populations were 77% smaller during mid-June and adult emergence delayed by >14 days on plants with weevils. In controlled experiments, weevils impaired sawfly growth by 18%, which correlated with 35% reductions in leaf phosphorus caused by root herbivory, a previously unreported mechanism for above-ground-below-ground herbivore interactions. This represents a clear demonstration of root herbivores affecting foliar herbivore community composition and natural enemy abundance in the field via two distinct plant-mediated nutritional mechanisms. Aphid populations, in particular, were initially driven by bottom-up effects (i.e. plant-mediated effects of root herbivory), but consequent increases in natural enemies triggered top-down regulation.

Concepts: Amino acid, Nutrition, Insect, Plant, Predation, Ecology, Ant, Herbivore


1.Intraguild predation (IGP) is a commonly recognised mechanism influencing the community structure of predators, but the complex interactions are notoriously difficult to disentangle. The mesopredator suppression hypothesis predicts that a superpredator may either simultaneously repress two mesopredators, restrain the dominant one and thereby release the subdominant mesopredator, or elicit different responses by both mesopredators. 2.We show the outcome arising from such conditions in a three-level predator assemblage (Eurasian eagle owl Bubo bubo L., northern goshawk Accipiter gentilis L., and common buzzard Buteo buteo L.) studied over 25 years. In the second half of the study period, the eagle owl re-colonised the study area, thereby providing a natural experiment of superpredator introduction. We combined this setup with detailed GIS-analysis of habitat use and a field experiment simulating intrusion by the superpredator into territories of the subdominant mesopredator, the buzzard. 3.Although population trends were positive for all three species in the assemblage, the proportion of failed breeding attempts increased significantly in both mesopredators after the superpredator re-colonised the area. 4.We predicted that superpredator-induced niche shifts in the dominant mesopredator may facilitate mesopredator coexistence in superpredator-free refugia. We found significant changes in nesting habitat choice in goshawk, but not in buzzard. Since competition for enemy-free refugia and the rapid increase in population density may have constrained niche shifts of the subdominant mesopredator, we further predicted behavioural changes in response to the superpredator. The field experiment indeed showed a significant increase in aggressive response of buzzards toward eagle owl territory intrusion over the course of ten years, probably due to phenotypic plasticity in the response towards superpredation risk. 5.Overall, our results show that intraguild predation can be a powerful force of behavioural change, simultaneously influencing habitat use and aggressiveness in predator communities. These changes might help to buffer mesopredator populations against the negative effects of intraguild predation. This article is protected by copyright. All rights reserved.

Concepts: Predation, Birds of Europe, Birds of Turkey, Birds of Pakistan, Common Buzzard, Northern Goshawk, Eurasian Eagle-owl, Birds of Asia


Quantifying the complex spatial dynamics taking place at range edges is critical for understanding future distributions of species, yet very few systems have sufficient data or the spatial resolution to empirically test these dynamics. This paper reviews how data from a large-scale pest management program have provided important contributions to the fields of population dynamics and invasion biology. The invasion of gypsy moth (Lymantria dispar) is well-documented from its introduction near Boston, Massachusetts USA in 1869 to its current extent of over 900,000 km(2) in Eastern North America. Over the past two decades, the USDA Forest Service Slow the Spread (STS) program for managing the future spread of gypsy moth has produced unrivaled spatiotemporal data across the invasion front. The STS program annually deploys a grid of 60,000 - 100,000 pheromone-baited traps, currently extending from Minnesota to North Carolina. The data from this program has provided the foundation for investigations of complex population dynamics and the ability to examine ecological hypotheses previously untestable outside of theoretical venues, particularly regarding invasive spread and Allee effects. This system provides empirical data on the importance of long-distance dispersal and time lags on population establishment and spatial spread. Studies showing high rates of spatiotemporal variation of the range edge, from rapid spread to border stasis and even retraction, highlight future opportunities to test mechanisms that influence both invasive and native species ranges. The STS trap data have also created a unique opportunity to study low-density population dynamics and quantify Allee effects with empirical data. Notable contributions include evidence for spatiotemporal variation in Allee effects, demonstrating empirical links between Allee effects and spatial spread, and testing mechanisms of population persistence and growth rates at range edges. There remain several outstanding questions in spatial ecology and population biology that can be tested within this system, such as the scaling of local ecological processes to large-scale dynamics across landscapes. The gypsy moth is an ideal model of how important ecological questions can be answered by thinking more broadly about monitoring data. This article is protected by copyright. All rights reserved.

Concepts: Scientific method, Biology, Ecology, Science, Empiricism, Empirical, Lymantriidae, Adam Copeland


Understanding drivers of population change is critical for effective species conservation. In the northeast Atlantic Ocean, recent changes amongst seabird communities are linked to human and climate change impacts on food webs. Many species have declined severely, with food shortages, and increased predation reducing productivity. Arctic skua Stercorarius parasiticus, a kleptoparasite of other seabirds, is one such species. The aim of the study was to determine relative effects of bottom-up and top-down pressures on Arctic skuas across multiple colonies in a rapidly declining national population. Long-term monitoring data were used to quantify changes in population size and productivity of Arctic skuas, their hosts (black-legged kittiwake Rissa tridactyla, common guillemot Uria aalge, Atlantic puffin Fratercula arctica, Arctic tern Sterna paradisaea) and an apex predator (great skua Stercorarius skua) over 24 years (1992-2015) in Scotland. We used digital mapping and statistical models to determine relative effects of bottom-up (host productivity) and top-down (great skua density) pressures on Arctic skuas across 33 colonies, and assess variation between three colony types classified by host abundance. Arctic skuas declined by 81% and their hosts by 42%-92%, whereas at most colonies great skuas increased. Annual productivity declined in Arctic skuas and their hosts, and reduced Arctic skua breeding success was a driver of the species' population decline. Arctic skua productivity was positively associated with annual breeding success of hosts and negatively with great skua density. Intercolony variation suggested Arctic skua trends and productivity were most sensitive to top-down pressures at smaller colonies of host species where great skuas had increased most, whereas bottom-up pressures dominated at large colonies of host species. Scotland’s Arctic skua population is declining rapidly, with bottom-up and top-down pressures simultaneously reducing breeding success to unsustainably low levels. Marine food web alterations, strongly influenced by fisheries management and climate change, are driving the decline, and this study demonstrates severe vulnerability of seabirds to rapid change in human-modified ecosystems. Potential but untested conservation solutions for Arctic skuas include marine protected areas, supplementary feeding within colonies and management of great skuas.