ALBERT

Description: Background: Partial migration, where one portion of a population conducts seasonal migrations while the other remains on a single range, is common in wild ungulate populations. However the relative costs and benefits associated with the distinct strategies adopted by coexisting migrant and resident individuals have rarely been investigated. Here we compare the body condition of migrants and residents in a partially migratory population of impalas (Aepyceros melampus) in Zimbabwe. The study was conducted during two consecutive years with highly contrasted population densities (16.4 and 8.6 indiv/km2) due to harvesting. Results: We first identify a population substructure with a north--south sub-division in two spatial units related to distinct soils and vegetation cover. Impalas in the north range had a consistently higher diet quality and body condition than those in the south range. At the beginning of the dry season about one third of the individuals migrated from the lower (i.e. south) to the higher (i.e. north) diet quality range. This partial migration pattern was consistent between the consecutive years, and most individuals showed constancy to their moving strategy (migrant or resident). In both years, these migrants had a significantly higher body condition at the end of the dry season than the south residents that remained year-round in the lower diet quality range. Diet quality and body condition of impalas were higher in the year of lower density; however we did not detect any evidence for density-dependence in migration propensity, at the individual or population levels, nor in the benefit associated with migration. Conclusions: Our findings provide rare evidence for a significant relationship between body condition and seasonal migration strategy in wild ungulates in relation to a difference in the quality of resources acquired between distinct seasonal ranges. This study also constitutes rare evidence of partial migration in a tropical ungulate population.

Description: Background: The marine-terrestrial transfer of salmon (Oncorhynchus spp.) provides a substantial pulse of nutrients to receiving ecosystems along the Pacific coast of North America and has been shown to enhance productivity and isotopic signatures of conifers and other riparian vegetation. An explicitly spatial, within-watershed investigation of the influence of salmon on conifers has never been previously investigated. In a small salmon-bearing watershed in Haida Gwaii, Canada, the transfer and distributional pattern of salmon carcasses into the riparian zone by black bears provided a spatial basis for investigating the influence of salmon on Sitka spruce tree ring growth and nitrogen isotopic signatures (delta15N) across a gradient of salmon carcass densities in relation to salmon escapement. Results: Annual growth was found to be highest in the high salmon carcass zone and delta15N signatures closely tracked the known distribution of salmon carcasses at distances into the forest and upstream. Tree diameter demonstrated a positive relationship with delta15N signatures for trees with and without salmon carcass influence. Using an information theoretics approach with general linear mixed models (GLMMs), we show that salmon abundance, mean annual temperature and the interaction terms salmon abundance*temperature and salmon abundance*distance into the forest best predict tree growth. In addition, spatial variables (distance into forest and upstream) and their interaction are the strongest predictors of delta15N signatures. However patterns observed in individual trees, particularly those at increased distance into the forest, suggest positive relationships with historical salmon abundance. Conclusions: Using a replicated spatial sampling design across a sharp gradient in salmon nutrient loading, our study provides clear evidence that the temporal pattern in an allochthonous nutrient source and an interaction with temperature and spatial location influences conifer growth. Although salmon abundance has been previously linked to annual conifer growth and delta15N levels, our approach demonstrates the need to incorporate additional predictors including tree size and opens up the prospect of their dual use as historical proxies for salmon abundance.

Description: Background: The important greenhouse gas (GHG) methane is produced naturally in anaerobic wetland soils. By affecting the production, oxidation and transport of methane to the atmosphere, plants have a major influence upon the quantities emitted by wetlands. Different species and functional plant groups have been shown to affect these processes differently, but our knowledge about how these effects are influenced by abiotic factors such as water regime and temperature remains limited. Here we present a mesocosm experiment comparing eight plant species for their effects on internal transport and overall emissions of methane under contrasting hydrological conditions. To quantify how much methane was transported internally through plants (the chimney effect), we blocked diffusion from the soil surface with an agar seal. Results: We found that graminoids caused higher methane emissions than forbs, although the emissions from mesocosms with different species were either lower than or comparable to those from control mesocosms with no plant (i.e. bare soil). Species with a relatively greater root volume and a larger biomass exhibited a larger chimney effect, though overall methane emissions were negatively related to plant biomass. Emissions were also reduced by lowering the water table. Conclusions: We conclude that plant species (and functional groups) vary in the degree to which they transport methane to the atmosphere. However, a plant with a high capacity to transport methane does not necessarily emit more methane, as it may also cause more rhizosphere oxidation of methane. A shift in plant species composition from graminoids to forbs and/or from low to high productive species may lead to reduction of methane emissions.

Description: Background: Landscape complexity can mitigate or facilitate host dispersal, influencing patterns of pathogen transmission. Spatial transmission of pathogens through landscapes, therefore, presents an important but not fully elucidated aspect of transmission dynamics. Using an agent-based model (LiNK) that incorporates GIS data, we examined the effects of landscape information on the spatial patterns of host movement and pathogen transmission in a system of long-tailed macaques and their gut parasites. We first examined the role of the landscape to identify any individual or additive effects on host movement. We then compared modeled dispersal distance to patterns of actual macaque gene flow to both confirm our model's predictions and to understand the role of individual land uses on dispersal. Finally, we compared the rate and the spread of two gastrointestinal parasites, Entamoeba histolytica and E. dispar, to understand how landscape complexity influences spatial patterns of pathogen transmission. Results: LiNK captured emergent properties of the landscape, finding that interaction effects between landscape layers could mitigate the rate of infection in a non-additive way. We also found that the inclusion of landscape information facilitated an accurate prediction of macaque dispersal patterns across a complex landscape, as confirmed by Mantel tests comparing genetic and simulated dispersed distances. Finally, we demonstrated that landscape heterogeneity proved a significant barrier for a highly virulent pathogen, limiting the dispersal ability of hosts and thus its own transmission into distant populations. Conclusions: Landscape complexity plays a significant role in determining the path of host dispersal and patterns of pathogen transmission. Incorporating landscape heterogeneity and host behavior into disease management decisions can be important in targeting response efforts, identifying cryptic transmission opportunities, and reducing or understanding potential for unintended ecological and evolutionary consequences. The inclusion of these data into models of pathogen transmission patterns improves our understanding of these dynamics, ultimately proving beneficial for sound public health policy.

Description: Background: The influence of introduction history and post-introduction dynamics on genetic diversity and structure has been a major research focus in invasion biology. However, genetic diversity and structure in the invasive range can also be affected by human-mediated processes in the native range prior to species introductions, an aspect often neglected in invasion biology. Here we aim to trace the native provenance of the invasive tree Acacia pycnantha by comparing the genetic diversity and structure between populations in the native Australian range and the invasive range in South Africa. This approach also allowed us to explore how human actions altered genetic structure before and after the introduction of A. pycnantha into South Africa. We hypothesized that extensive movement and replanting in A. pycnantha's Australian range prior to its introduction to South Africa might result in highly admixed genotypes in the introduced range, comparable genetic diversity in both ranges, and therefore preclude an accurate determination of native provenance(s) of invasive populations. Results: In the native range Bayesian assignment tests identified three genetic clusters with substantial admixture and could not clearly differentiate previously identified genetic entities, corroborating admixture as a result of replantings within Australia. Assignment tests that included invasive populations from South Africa indicated similar levels of admixture compared to Australian populations and a lack of genetic structure. Invasive populations of A. pycnantha in South Africa are as genetically diverse as native populations, and could not be assigned to particular native range regions. Conclusions: Our results indicate that the genetic structure of A. pycnantha in Australia has been greatly altered through various planting initiatives. Specifically, there is little geographic structure and high levels of admixture. While numerous introduction history scenarios may explain the levels of admixture observed in South Africa, planting records of A. pycnantha in Australia suggest that populations were probably already admixed before propagules were introduced to South Africa. These findings have important implications for the management of invasive A. pycnantha populations in South Africa, especially for classical biological control, and more broadly, for studies that aim to understand the evolutionary dynamics of the invasion process.

Description: Background: Japanese black bears, a large-bodied omnivore, frequently create small gaps in the tree crown during fruit foraging. However, there are no previous reports of black bear-created canopy gaps. To characterize physical canopy disturbance by black bears, we examined a number of parameters, including the species of trees in which canopy gaps were created, gap size, the horizontal and vertical distribution of gaps, and the size of branches broken to create gaps. The size of black bear-created canopy gaps was estimated using data from branches that had been broken and dropped on the ground. Results: The disturbance regime was characterized by a highly biased distribution of small canopy gaps on ridges, a large total overall gap area, a wide range in gap height relative to canopy height, and diversity in gap size. Surprisingly, the annual rate of bear-created canopy gap formation reached 141.3 m2 ha--1 yr--1 on ridges, which were hot spots in terms of black bear activity. This rate was approximately 6.6 times that of tree-fall gap formation on ridges at this study site. Furthermore, this rate was approximately two to three times that of common tree-fall gap formation in Japanese forests, as reported in other studies. Conclusions: Our findings suggest that the ecological interaction between black bears and fruit-bearing trees may create a unique light regime, distinct from that created by tree falls, which increases the availability of light resources to plants below the canopy.

Description: Background Developing a quick and reliable technique to estimate floral cover in deserts will assist in monitoring and management. The present attempt was to estimate plant cover in the UAE desert using both digital photography and field sampling. Digital photographs were correlated with field data to estimate floral cover in moderately (Al-Maha) and heavily (DDCR) grazed areas. The Kruskal-Wallis test was also used to assess compatibility between the two techniques within and across grazing intensities and soil substrates.Results Results showed that photographs could be a reliable technique within the sand dune substrate under moderate grazing (r = 0.69). The results were very poorly correlated (r =-0.24) or even inversely proportional (r =-0.48) when performed within DDCR. Overall, Chi-square values for Al-Maha and DDCR were not significant at P 〉 0.05, indicating similarities between the two methods. At the soil type level, the Kruskal-Wallis analysis was not significant (P 〉 0.05), except for gravel plains (P 〈 0.05). Across grazing intensities and soil substrates, the two techniques were in agreement in ranking most plant species, except for Lycium shawii.Conclusions Consequently, the present study has proven that digital photography could not be used reliably to asses floral cover, while further testing is required to support such claim. An image-based sampling approach of plant cover at the species level, across different grazing and substrate variations in desert ecosystems, has its uses, but results are to be cautiously interpreted.

Description: Background: Herbivore coexistence on the Late Cretaceous island continent of Laramidia has been a topic of great interest, stemming from the paradoxically high diversity and biomass of these animals in relation to the relatively small landmass available to them. Various hypotheses have been advanced to account for these facts, of which niche partitioning is among the most frequently invoked. However, despite its wide acceptance, this hypothesis has not been rigorously tested. This study uses the fossil assemblage from the Dinosaur Park Formation of Alberta as a model to investigate whether niche partitioning facilitated herbivorous dinosaur coexistence on Laramidia. Specifically, the question of feeding height stratification is examined in light of the role it plays in facilitating modern ungulate coexistence. Results: Most herbivorous dinosaur species from the Dinosaur Park Formation were restricted to feeding no higher than approximately 1 m above the ground. There is minimal evidence for feeding height partitioning at this level, with ceratopsids capable of feeding slightly higher than ankylosaurs, but the ecological significance of this is ambiguous. Hadrosaurids were uniquely capable of feeding up to 2 m quadrupedally, or up to 5 m bipedally. There is no evidence for either feeding height stratification within any of these clades, or for change in these ecological relationships through the approximately 1.5 Ma record of the Dinosaur Park Formation. Conclusions: Although we cannot reject the possibility, we find no good evidence that feeding height stratification, as revealed by reconstructed maximum feeding heights, played an important role in facilitating niche partitioning among the herbivorous dinosaurs of Laramdia. Most browsing pressure was concentrated in the herb layer, although hadrosaurids were capable of reaching shrubs and low-growing trees that were out of reach from ceratopsids, ankylosaurs, and other small herbivores, effectively dividing the herbivores in terms of relative abundance. Sympatric hadrosaurids may have avoided competing with one another by feeding differentially using bipedal and quadrupedal postures. These ecological relationships evidently proved to be evolutionarily stable because they characterize the herbivore assemblage of the Dinosaur Park Formation through time. If niche partitioning served to facilitate the rich diversity of these animals, it may have been achieved by other means in addition to feeding height stratification. Consideration of other feeding height proxies, including dental microwear and skull morphology, may help to alleviate problems of underdetermination identified here.

Description: Background: Pathogens and parasites can have major impacts on host population dynamics, both through direct mortality and via indirect effects. Both types of effect may be stronger in species whose populations are already under pressure. We investigated the potential for blood parasites to impact upon their hosts at the immunological, physiological and population level during the non-breeding season using a declining population of yellowhammers Emberiza citrinella as a model. Results: Yellowhammers infected by Haemoproteus spp. showed both a reduced heterophil to lymphocyte (H:L) ratio, and an elevated standardised white blood cell (WBC) count compared to uninfected birds, indicating an immunological response to infection. Infected birds had shorter wings during the first winter of sampling but not during the second, colder, winter; survival analysis of 321 birds sampled across four winters indicated that increased wing length conferred a survival advantage. Conclusions: We suggest that the potential impacts of blood parasite infections on over-wintering birds may have been underestimated. Further research should consider the potential impacts of sub-clinical parasite infections on the dynamics of vulnerable populations, and we suggest using declining populations as model systems within which to investigate these relationships as well as examining interactions between sub-clinical disease and other environmental stressors.JEL Classification Code: Q5

Description: Background: Adaptive behavioural strategies promoting co-occurrence of competing species are known to result from a sympatric evolutionary past. Strategies should be different for indirect resource competition (exploitation, e.g., foraging and avoidance behaviour) than for direct interspecific interference (e.g., aggression, vigilance, and nest guarding). We studied the effects of resource competition and nest predation in sympatric small mammal species using semi-fossorial voles and shrews, which prey on vole offspring during their sensitive nestling phase. Experiments were conducted in caged outdoor enclosures. Focus common vole mothers (Microtus arvalis) were either caged with a greater white-toothed shrew (Crocidura russula) as a potential nest predator, with an herbivorous field vole (Microtus agrestis) as a heterospecific resource competitor, or with a conspecific resource competitor. Results: We studied behavioural adaptations of vole mothers during pregnancy, parturition, and early lactation, specifically modifications of the burrow architecture and activity at burrow entrances. Further, we measured pre- and postpartum faecal corticosterone metabolites (FCMs) of mothers to test for elevated stress hormone levels. Only in the presence of the nest predator were prepartum FCMs elevated, but we found no loss of vole nestlings and no differences in nestling body weight in the presence of the nest predator or the heterospecific resource competitor. Although the presence of both the shrew and the field vole induced prepartum modifications to the burrow architecture, only nest predators caused an increase in vigilance time at burrow entrances during the sensitive nestling phase. Conclusion: Voles displayed an adequate behavioural response for both resource competitors and nest predators. They modified burrow architecture to improve nest guarding and increased their vigilance at burrow entrances to enhance offspring survival chances. Our study revealed differential behavioural adaptations to resource competitors and nest predators.

Description: Background: Individual body growth is controlled in large part by the spatial and temporal heterogeneity of, and competition for, resources. Grizzly bears (Ursus arctos L.) are an excellent species for studying the effects of resource heterogeneity and maternal effects (i.e. silver spoon) on life history traits such as body size because their habitats are highly variable in space and time. Here, we evaluated influences on body size of grizzly bears in Alberta, Canada by testing six factors that accounted for spatial and temporal heterogeneity in environments during maternal, natal and 'capture' (recent) environments. After accounting for intrinsic biological factors (age, sex), we examined how body size, measured in mass, length, and body condition, was influenced by: (a) population density; (b) regional habitat productivity; (c) inter-annual variability in productivity (including silver spoon effects); (d) local habitat quality; (e) human footprint (disturbances); and (f) landscape change. Results: We found sex and age explained the most variance in body mass, condition and length (R2 from 0.48--0.64). Inter-annual variability in climate the year before and of birth (silver spoon effects) had detectable effects on the three-body size metrics (R2 from 0.04--0.07); both maternal (year before birth) and natal (year of birth) effects of precipitation and temperature were related with body size. Local heterogeneity in habitat quality also explained variance in body mass and condition (R2 from 0.01--0.08), while annual rate of landscape change explained additional variance in body length (R2 = 0.03). Human footprint and population density had no observed effect on body size. Conclusions: These results illustrated that body size patterns of grizzly bears, while largely affected by basic biological characteristics (age and sex), were also influenced by regional environmental gradients the year before, and of, the individual's birth thus illustrating silver spoon effects. The magnitude of the silver spoon effects was on par with the influence of contemporary regional habitat productivity, which showed that both temporal and spatial influences explain in part body size patterns in grizzly bears. Because smaller bears were found in colder and less-productive environments, we hypothesize that warming global temperatures may positively affect body mass of interior bears, provided there is sufficient snow-cover during the denning periods.

Description: Background: Building reference libraries of DNA barcodes is relatively straightforward when specifically designed primers are available to amplify the COI-5P region from a relatively narrow taxonomic group (e.g. single class or single order). DNA barcoding marine communities have been comparatively harder to accomplish due to the broad taxonomic diversity and lack of consistently efficient primers. Although some of the so-called "universal" primers have been relatively successful, they still fail to amplify COI-5P of many marine animal groups, while displaying random success even among species within each group. Here we propose a new pair of primers designed to enhance amplification of the COI-5P region in a wide range of marine organisms. Results: Amplification tests conducted on a wide range of marine animal taxa, rendered possible the first--time sequencing of DNA barcodes from eight separated phyla (Annelida, Arthropoda, Chordata, Cnidaria, Echinodermata, Mollusca, Nemertea and Platyhelminthes), comprising a total of 14 classes, 28 orders, 57 families, 68 genus and 76 species. Conclusions: These primers demonstrated to be highly cost-effective, which is of key importance for DNA barcoding procedures, such as for building comprehensive DNA barcode libraries of marine communities, where the processing of a large numbers of specimens from a wide variety of marine taxa is compulsory.

Description: Background: Stable isotope ratios (13C/12C and 18O/16O) in fossil teeth and bone provide key archives for understanding the ecology of extinct horses during the Plio-Pleistocene in South America; however, what happened in areas of sympatry between Equus (Amerhippus) and Hippidion is less understood. Results: Here, we use stable carbon and oxygen isotopes preserved in 67 fossil tooth and bone samples for seven species of horses from 25 different localities to document the magnitude of the dietary shifts of horses and ancient floral change during the Pliocene to Pleistocene. Dietary reconstructions inferred from stable isotopes of both genera of horses present in South America document dietary separation and environmental changes in ancient ecosystems, including C3/C4 transitions. Stable isotope data demonstrate changes in C4 grass consumption, inter-species dietary partitioning and variation in isotopic niche breadth of mixed feeders with latitudinal gradient. Conclusions: The data for Hippidion indicate a preference varying from C3 plants to mixed C3-C4 plants in their diet. Equus (Amerhippus) shows three different patterns of dietary partitioning Equus (A.) neogeus from the province of Buenos Aires indicate a preference for C3 plants in the diet. Equus (A.) andium from Ecuador and Equus (A.) insulatus from Bolivia show a preference for to a diet of mixed C3-C4 plants, while Equus (A.) santaeelenae from La Carolina (sea level of Ecuador) and Brazil are mostly C4 feeders. These results confirm that ancient feeding ecology cannot always be inferred from dental morphology. While the carbon isotope composition of horses skeletal material decreased as latitude increased, we found evidence of boundary between a mixed C3/C4 diet signal and a pure C4 signal around 32o S and a change from a mixed diet signal to an exclusively C3 signal around 35oS.We found that the horses living at high altitudes and at low to middle latitude still have a C4 component in their diet, except the specimens from 4000m, which have a pure C3 diet. The change in altitudinal vegetation gradients during the Pleistocene is one of several possibilities to explain the C4 dietary component in horses living at high altitudes. Other alternative explanations imply that the horses fed partially at lower altitudes.

Description: Background: Parasites that manipulate host behavior can provide prominent examples of extended phenotypes: parasite genomes controlling host behavior. Here we focus on one of the most dramatic examples of behavioral manipulation, the death grip of ants infected by Ophiocordyceps fungi. We studied the interaction between O. unilateralis s.l. and its host ant Camponotus leonardi in a Thai rainforest, where infected ants descend from their canopy nests down to understory vegetation to bite into abaxial leaf veins before dying. Host mortality is concentrated in patches (graveyards) where ants die on sapling leaves ca. 25 cm above the soil surface where conditions for parasite development are optimal. Here we address whether the sequence of ant behaviors leading to the final death grip can also be interpreted as parasite adaptations and describe some of the morphological changes inside the heads of infected workers that mediate the expression of the death grip phenotype. Results: We found that infected ants behave as zombies and display predictable stereotypical behaviors of random rather than directional walking, and of repeated convulsions that make them fall down and thus precludes returning to the canopy. Transitions from erratic wandering to death grips on a leaf vein were abrupt and synchronized around solar noon. We show that the mandibles of ants penetrate deeply into vein tissue and that this is accompanied by extensive atrophy of the mandibular muscles. This lock-jaw means the ant will remain attached to the leaf after death. We further present histological data to show that a high density of single celled stages of the parasite within the head capsule of dying ants are likely to be responsible for this muscular atrophy. Conclusions: Extended phenotypes in ants induced by fungal infections are a complex example of behavioral manipulation requiring coordinated changes of host behavior and morphology. Future work should address the genetic basis of such extended phenotypes.

Description: Background: Sand fly saliva can drive the outcome of Leishmania infection in animal models, and salivary components have been postulated as vaccine candidates against leishmaniasis. In the sand fly Phlebotomus papatasi, natural sugar-sources modulate the activity of proteins involved in meal digestion, and possibly influence vectorial capacity. However, only a handful of studies have assessed the variability of salivary components in sand flies, focusing on the effects of environmental factors in natural habitats. In order to better understand such interactions, we compared the expression profiles of nine P. papatasi salivary gland genes of specimens inhabiting different ecological habitats in Egypt and Jordan and throughout the sand fly season in each habitat. Results: The majority of investigated genes were up-regulated in specimens from Swaymeh late in the season, when the availability of sugar sources is reduced due to water deprivation. On the other hand, these genes were not up-regulated in specimens collected from Aswan, an irrigated area less susceptible to drought effects. Conclusion: Expression plasticity of genes involved with vectorial capacity in disease vectors may play an important epidemiological role in the establishment of diseases in natural habitats.

Description: Background: Understanding the phenotypic consequences of interactions between arthropod-borne viruses (arboviruses) and their mosquito hosts has direct implications for predicting the evolution of these relationships and the potential for changes in epidemiological patterns. Although arboviruses are generally not highly pathogenic to mosquitoes, pathology has at times been noted. Here, in order to evaluate the potential costs of West Nile virus (WNV) infection and resistance in a primary WNV vector, and to assess the extent to which virus-vector relationships are species-specific, we performed fitness studies with and without WNV exposure using a highly susceptible Culex pipiens mosquito colony. Specifically, we measured and compared survival, fecundity, and feeding rates in bloodfed mosquitoes that were (i) infected following WNV exposure (susceptible), (ii) uninfected following WNV exposure (resistant), or (iii) unexposed. Results: In contrast to our previous findings with a relatively resistant Cx. tarsalis colony, WNV infection did not alter fecundity or blood-feeding behaviour of Cx. pipiens, yet results do indicate that resistance to infection is associated with a fitness cost in terms of mosquito survival. Conclusions: The identification of species-specific differences provides an evolutionary explanation for variability in vector susceptibility to arboviruses and suggests that understanding the costs of infection and resistance are important factors in determining the potential competence of vector populations for arboviruses.

Description: Background: The North American Trichoptera larvae are poorly known at the species level, despite their importance in the understanding of freshwater fauna and critical use in biomonitoring. This study focused on morphological diagnoses for larvae occurring in the Churchill, Manitoba area, representing the largest larval association effort for the caddisflies at any given locality thus far. The current DNA barcode reference library of Trichoptera (available on the Barcode of Life Data System) was utilized to deliver larval-adult associations. Results: The present study collected an additional 23 new species records for the Churchill area, increasing the total Trichoptera richness to 91 species. We were able to associate 62 larval taxa, comprising 68.1 % of the Churchill area Trichoptera taxa. This endeavor to identify immature life stage for the caddisflies enabled the development of morphological diagnoses, production of photographs and an appropriate taxonomic key to facilitate larval species analyses in the area. Conclusions: The use of DNA for associations of unknown larvae with known adults proved rapid and successful. This method should accelerate the state-of-knowledge for North American Trichoptera larvae as well as other taxonomic lineages. The morphological analysis should be useful for determination of material from the Churchill area.

Description: Background: Previous work has shown that leaf-cutting ants prefer to cut leaf material with relatively low fungal endophyte content. This preference suggests that fungal endophytes exact a cost on the ants or on the development of their colonies. We hypothesized that endophytes may play a role in their host plants' defense against leaf-cutting ants. To measure the long-term cost to the ant colony of fungal endophytes in their forage material, we conducted a 20-week laboratory experiment to measure fungal garden development for colonies that foraged on leaves with low or high endophyte content. Results: Colony mass and the fungal garden dry mass did not differ significantly between the low and high endophyte feeding treatments. There was, however, a marginally significant trend toward greater mass of fungal garden per ant worker in the low relative to the high endophyte treatment. This trend was driven by differences in the fungal garden mass per worker from the earliest samples, when leaf-cutting ants had been foraging on low or high endophyte leaf material for only 2 weeks. At two weeks of foraging, the mean fungal garden mass per worker was 77% greater for colonies foraging on leaves with low relative to high endophyte loads. Conclusions: Our data suggest that the cost of endophyte presence in ant forage material may be greatest to fungal colony development in its earliest stages, when there are few workers available to forage and to clean leaf material. This coincides with a period of high mortality for incipient colonies in the field. We discuss how the endophyte-leaf-cutter ant interaction may parallel constitutive defenses in plants, whereby endophytes reduce the rate of colony development when its risk of mortality is greatest.

Description: EditorialDavid Pimentel is a professor of ecology and agricultural sciences at Cornell University, Ithaca, NY 14853–0901. His Ph.D. is from Cornell University and had postdoctoral research at the University of Chicago, MIT, and fellowship at Oxford University (England). He was awarded a distinguished honorary degree from the University of Massachusetts. His research spans the fields of energy, population ecology, biological pest control, pesticides, sustainable agriculture, land and water conservation, livestock, and environmental policy. Pimentel has published more than 700 scientific papers and 37 books and has served on many national and government committees including the National Academy of Sciences; President’s Science Advisory Council; U.S Department of Agriculture; U.S. Department of Energy; U.S. Department of Health, Education and Welfare; Office of Technology Assessment of the U.S. Congress; and the U.S. State Department. He is currently Editorial Advisor for BMC Ecology. In this article, he reflects on 50 years since the publication of Rachel Carson’s influential book, Silent Spring.

Description: Background: Information on the age structure within populations of an endangered species can facilitate effective management. The Blue Mountains Water Skink (Eulamprus leuraensis) is a viviparous scincid lizard that is restricted to 〈 40 isolated montane swamps in south-eastern Australia. We used skeletochronology of phalanges (corroborated by mark-recapture data) to estimate ages of 222 individuals from 13 populations. Results: These lizards grow rapidly, from neonatal size (30 mm snout-vent length) to adult size (about 70 mm SVL) within two to three years. Fecundity is low (mean 2.9 offspring per litter) and is affected by maternal body length and age. Offspring quality may decline with maternal age, based upon captive-born neonates (older females gave birth to slower offspring). In contrast to its broadly sympatric (and abundant) congener E. tympanum, E. leuraensis is short-lived (maximum 6 years, vs 15 years for E. tympanum). Litter size and offspring size are similar in the two species, but female E. leuraensis reproduce annually whereas many E. tympanum produce litters biennially. Thus, a low survival rate (rather than delayed maturation or low annual fecundity) is the key reason why E. leuraensis is endangered. Our 13 populations exhibited similar growth rates and population age structures despite substantial variation in elevation, geographic location and swamp size. However, larger populations (based on a genetic estimate of effective population size) contained older lizards, and thus a wider variance in ages. Conclusion: Our study suggests that low adult survival rates, as well as specialisation on a rare and fragmented habitat type (montane swamps) contribute to the endangered status of the Blue Mountains Water Skink.

Description: Contributing reviewersThe editors of BMC Ecology would like to thank all of our reviewers who have contributed to the journal in volume 12 (2012).

Description: Background: Movement behaviour can be influenced by a multitude of biotic and abiotic factors. Here, we investigate the speed of movement in relation to environmental and individual phenotypic properties in subadult common lizards (Lacerta vivipara). We aim to disentangle the importance of substrate, cover, humidity, basking opportunity and individual phenotype on moving tendencies in 12 treatment combinations, at which each lizard was tested. Results: We find that movement behaviour depends on the starting conditions, the physical properties of the dispersal corridor, and on the individuals' phenotype. Specifically, the presence of cover and substrate providing suitable traction in the corridor had positive effects on individual movement decisions. Additionally, we find high phenotypic variation in the propensity to move dependent on the presence of cover. Individual back patterns also strongly affected movement decisions in interaction with the physical properties of the dispersal corridor. Conclusions: Our results highlight the importance of understanding the habitat resistance for movement patterns, with humid habitats with covering vegetation providing the best conditions to initiate movement in the common lizard. In addition, population effects, differences in back pattern phenotype and individual plasticity were identified as key parameters influencing movement behaviour.

Description: Background: The positive relationship between habitat area and species number is considered a fundamental rule in ecology. This relationship predicts that the link number of species interactions increases with habitat area, and structure is related to habitat area. Biological invasions can affect species interactions and area relationships. However, how these relationships change at different spatial scales has remained unexplored. We analysed understory plant-pollinator networks in seven temperate forest sites at 20 spatial scales (radius 120-2020 m) to clarify scale-associated relationships between forest area and plant-pollinator networks. Results: The pooled data described interactions between 18 plant (including an exotic) and 89 pollinator (including an exotic) species. The total number of species and the number of interaction links between plant and pollinator species were negatively correlated with forest area, with the highest correlation coefficient at radii of 1520 and 1620 m, respectively. These results are not concordant with the pattern predicted by species-area relationships. However, when associations with exotic species were excluded, the total number of species and the number of interaction links were positively correlated with forest area (the highest correlation coefficient at a radius of 820 m). The network structure, i.e., connectance and nestedness, was also related to forest area (the highest correlation coefficients at radii of 720-820 m), when associations with exotics were excluded. In the study area, the exotic plant species Alliaria petiolata, which has invaded relatively small forest patches surrounded by agricultural fields, may have supported more native pollinator species than initially expected. Therefore, this invasive plant may have altered the original relationships between forest area and plant-pollinator networks. Conclusions: Our results demonstrate scale-dependent effects of forest area on the size and structure of plant-pollinator networks. We also suggest that a single exotic plant species can impact plant-pollinator networks, even in temperate continental habitats.

Description: Background: Although inducible defences have been studied extensively, only little is known about how the presence of parasites might interfere with these anti-predator adaptations. Both parasites and predators are important factors shaping community structure and species composition of ecosystems. Here, we simultaneously exposed Daphnia magna to predator cues (released by the tadpole shrimp, Triops, or by a fish) and spores of the yeast parasite Metschnikowia sp. to determine how life history and morphological inducible defences against these two contrasting types of predators are affected by infection. Results: The parasite suppressed some Triops-induced defences: Daphnia lost the ability to produce a greater number of larger offspring, a life-history adaptation to Triops predation. In contrast, the parasite did not suppress inducible defences against fish: induction (resulting in smaller body length of the mothers as well as of their offspring) and infection acted additively on the measured traits. Thus, fish-induced defences may be less costly than inducible defences against small invertebrate predators like Triops; the latter defences could no longer be expressed when the host had already invested in fighting off the parasite. Conclusions: In summary, our study suggests that as specific inducible defences differ in their costs, some might be suppressed if a target prey is additionally infected. Therefore, adding parasite pressure to predator-prey systems can help to elucidate the costs of inducible defences.

Description: Background: Organisms modify their environment and in doing so change the quantity and possibly the quality of available resources. Due to the two-way relationship between organisms and their resource environment, and the complexity it brings to biological communities, measuring species interactions reliably in any biological system is a challenging task. As the resource environment changes, the intensity and even the sign of interactions may vary in time. We used Serratia marcescens and Novosphingobium capsulatum bacteria to study how the interaction between resource environment and organisms influence the growth of the bacterial species during circa 200 generations. We used a sterile-filtering method to measure how changes in resource environment are reflected in growth rates of the two species. Results: Changes in the resource environment caused complex time and species composition-dependent effects on bacterial growth performance. Variation in the quality of the growth medium indicated existence of temporally fluctuating within-species facilitation and inhibition, and between-species asymmetric facilitation. Conclusions: The interactions between the community members could not be fully predicted based only on the knowledge of the growth performance of each member in isolation. Growth dynamics in sterile-filtered samples of the conditioned growth medium can reveal both biologically meaningful changes in resource availability and temporally changing facilitative resource-mediated interactions between study species. This is the first study we are aware of where the filter-sterilization - growth assay method is applied to study the effect of long-term changes in the environment on species interactions.

Description: Background: Evidence is accumulating that telomere length is a good predictor of life expectancy, especially early in life, thus calling for determining the factors that affect telomere length at this stage. Here, we investigated the relative influence of early growth conditions and origin (genetics and early maternal effects) on telomere length of collared flycatchers (Ficedula albicollis) at fledging. We experimentally transferred hatchlings among brood triplets to create reduced, control (i.e. unchanged final nestling number) and enlarged broods. Results: Although our treatment significantly affected body mass at fledging, we found no evidence that increased sibling competition affected nestling tarsus length and telomere length. However, mixed models showed that brood triplets explained a significant part of the variance in body mass (18%) and telomere length (19%), but not tarsus length (13%), emphasizing that unmanipulated early environmental factors influenced telomere length. These models also revealed low, but significant, heritability of telomere length (h2 = 0.09). For comparison, the heritability of nestling body mass and tarsus length was 0.36 and 0.39, respectively, which was in the range of previously published estimates for those two traits in this species. Conclusion: Those findings in a wild bird population demonstrate that telomere length at the end of the growth period is weakly, but significantly, determined by genetic and/or maternal factors taking place before hatching. However, we found no evidence that the brood size manipulation experiment, and by extension the early growth conditions, influenced nestling telomere length. The weak heritability of telomere length suggests a close association with fitness in natural populations.

Description: Background: The outcome of male-male competition can be predicted from the relative fighting qualities of the opponents, which often depend on their age. In insects, freshly emerged and still sexually inactive males are morphologically indistinct from older, sexually active males. These young inactive males may thus be easy targets for older males if they cannot conceal themselves from their attacks. The ant Cardiocondyla obscurior is characterised by lethal fighting between wingless ("ergatoid") males. Here, we analyse for how long young males are defenceless after eclosion, and how early adult males can detect the presence of rival males. Results: We found that old ergatoid males consistently won fights against ergatoid males younger than two days. Old males did not differentiate between different types of unpigmented pupae several days before emergence, but had more frequent contact to ready-to-eclose pupae of female sexuals and winged males than of workers and ergatoid males. In rare cases, old ergatoid males displayed alleviated biting of pigmented ergatoid male pupae shortly before adult eclosion, as well as copulation attempts to dark pupae of female sexuals and winged males. Ergatoid male behaviour may be promoted by a closer similarity of the chemical profile of ready-to-eclose pupae to the profile of adults than that of young pupae several days prior to emergence. Conclusion: Young ergatoid males of C. obscurior would benefit greatly by hiding their identity from older, resident males, as they are highly vulnerable during the first two days of their adult lives. In contrast to the winged males of the same species, which are able to prevent ergatoid male attacks by chemical female mimicry, young ergatoids do not seem to be able to produce a protective chemical profile. Conflicts in male-male competition between ergatoid males of different age thus seem to be resolved in favour of the older males. This might represent selection at the colony level rather than the individual level.

Description: Background: A central question for ecologists is the extent to which anthropogenic disturbances (e.g. tourism) might impact wildlife and affect the systems under study. From a research perspective, identifying the effects of human disturbance caused by research-related activities is crucial in order to understand and account for potential biases and derive appropriate conclusions from the data. Results: Here, we document a case of biological adjustment to chronic human disturbance in a colonial seabird, the king penguin (Aptenodytes patagonicus), breeding on remote and protected islands of the Southern ocean. Using heart rate (HR) as a measure of the stress response, we show that, in a colony with areas exposed to the continuous presence of humans (including scientists) for over 50 years, penguins have adjusted to human disturbance and habituated to certain, but not all, types of stressors. When compared to birds breeding in relatively undisturbed areas, birds in areas of high chronic human disturbance were found to exhibit attenuated HR responses to acute anthropogenic stressors of low-intensity (i.e. sounds or human approaches) to which they had been subjected intensely over the years. However, such attenuation was not apparent for high-intensity stressors (i.e. captures for scientific research) which only a few individuals experience each year. Conclusions: Habituation to anthropogenic sounds/approaches could be an adaptation to deal with chronic innocuous stressors, and beneficial from a research perspective. Alternately, whether penguins have actually habituated to anthropogenic disturbances over time or whether human presence has driven the directional selection of human-tolerant phenotypes, remains an open question with profound ecological and conservation implications, and emphasizes the need for more knowledge on the effects of human disturbance on long-term studied populations.

Description: Kirtland's warblers are the rarest songbird species in North America, rarity due in part to a reliance on early successional Jack Pine forests. Habitat loss due to fire suppression led to population declines to fewer than 200 males during the 1970s. Subsequent conservation management has allowed the species to recover to over 1700 males by 2010. In this study, we directly examine the impact that low population sizes have had on genetic variation in Kirtland's warblers. We compare the molecular variation of samples collected in Oscoda County, Michigan across three time periods: 1903-1912, 1929-1955 and 2008-2009. Results: In a hierarchical rarified sample of 20 genes and one time period, allelic richness was highest in 1903-1912 sample (AR=5.96), followed by the 1929-1955 sample (AR= 5.74), and was lowest in the 2008-2009 sample (AR= 5.54). Heterozygosity measures were not different between the 1929-1955 and 2008-2009 samples, but were lower in the 1903-1912 sample. Under some models, a genetic bottleneck signature was present in the 1929-1955 and 2008-2009 samples but not in the 1903-1912 sample. Conclusions: We suggest that these temporal genetic patterns are the result of the declining Kirtland's warbler population compressing into available habitat and a consequence of existing at low numbers for several decades.

Description: Background: Theory predicts a nonlinear response of dispersal evolution to habitat fragmentation. First, dispersal will be favoured in line with both decreasing area of habitat patches and increasing inter-patch distances. Next, once these inter-patch distances exceed a critical threshold, dispersal will be counter-selected, unless essential resources no longer co-occur in compact patches but are differently scattered; colonization of empty habitat patches or rescue of declining populations are then increasingly overruled by dispersal costs like mortality risks and loss of time and energy. However, to date, most empirical studies mainly document an increase of dispersal associated with habitat fragmentation. We analyzed dispersal kernels for males and females of the common, widespread woodland butterfly Pararge aegeria in highly fragmented landscape, and for males in landscapes that differed in their degree of habitat fragmentation. Results: The male and female probabilities of moving were considerably lower in the highly fragmented landscapes compared to the male probability of moving in fragmented agricultural and deciduous oak woodland landscapes. We also investigated whether, and to what extent, daily dispersal distance in the highly fragmented landscape was influenced by a set of landscape variables for both males and females, including distance to the nearest woodland, area of the nearest woodland, patch area and abundance of individuals in the patch. We found that daily movement distance decreased with increasing distance to the nearest woodland in both males and females. Daily distances flown by males were related to the area of the woodland capture site, whereas no such effect was observed for females. Conclusion: Overall, mobility was strongly reduced in the highly fragmented landscape, and varied considerably among landscapes with different spatial resource distributions. We interpret the results relative to different cost-benefit ratios of movements in fragmented landscapes.

Description: Globally, forests cover nearly one third of the land area and they contain over 80 % of terrestrial biodiversity. Both the extent and quality of forest habitat continue to decrease and the associated loss of biodiversity jeopardizes forest ecosystem functioning and the ability of forests to provide ecosystem services. In the light of the increasing population pressure, it is of major importance not only to conserve, but also to restore forest ecosystems.Ecological restoration has recently started to adopt insights from the biodiversity-ecosystem functioning (BEF) perspective. Central is the focus on restoring the relation between biodiversity and ecosystem functioning. Here we provide an overview of important considerations related to forest restoration that can be inferred from this BEF-perspective.Restoring multiple forest functions requires multiple species. It is highly unlikely that species-poor plantations, which may be optimal for above-ground biomass production, will outperform species diverse assemblages for a combination of functions, including overall carbon storage and control over water and nutrient flows. Restoring stable forest functions also requires multiple species. In particular in the light of global climatic change scenarios, which predict more frequent extreme disturbances and climatic events, it is important to incorporate insights from the relation between biodiversity and stability of ecosystem functioning into forest restoration projects. Rather than focussing on species per se, focussing on functional diversity of tree species assemblages seems appropriate when selecting tree species for restoration. Finally, also plant genetic diversity and above - below-ground linkages should be considered during the restoration process, as these likely have prominent but until now poorly understood effects at the level of the ecosystem.The BEF-approach provides a useful framework to evaluate forest restoration in an ecosystem functioning context, but it also highlights that much remains to be understood, especially regarding the relation between forest functioning on the one side and genetic diversity and above-ground-below-ground species associations on the other. The strong emphasis of the BEF-approach on functional rather than taxonomic diversity may also be the beginning of a paradigm shift in restoration ecology, increasing the tolerance towards allochthonous species.

Description: Background: To identify the determinants of invasiveness, comparisons of traits of invasive and native species are commonly performed. Invasiveness is generally linked to higher values of reproductive, physiological and growth-related traits of the invasives relative to the natives in the introduced range. Phenotypic plasticity of these traits has also been cited to increase the success of invasive species but has been little studied in invasive tree species. In a greenhouse experiment, we compared ecophysiological traits between an invasive species to Europe, Acer negundo, and early- and late-successional co-occurring native species, under different light, nutrient availability and disturbance regimes. We also compared species of the same species groups in situ, in riparian forests. Results: Under non-limiting resources, A. negundo seedlings showed higher growth rates than the native species. However, A. negundo displayed equivalent or lower photosynthetic capacities and nitrogen content per unit leaf area compared to the native species; these findings were observed both on the seedlings in the greenhouse experiment and on adult trees in situ. These physiological traits were mostly conservative along the different light, nutrient and disturbance environments. Overall, under non-limiting light and nutrient conditions, specific leaf area and total leaf area of A. negundo were substantially larger. The invasive species presented a higher plasticity in allocation to foliage and therefore in growth with increasing nutrient and light availability relative to the native species. Conclusions: The higher level of plasticity of the invasive species in foliage allocation in response to light and nutrient availability induced a better growth in non-limiting resource environments. These results give us more elements on the invasiveness of A. negundo and suggest that such behaviour could explain the ability of A. negundo to outperform native tree species, contributes to its spread in European resource-rich riparian forests and impedes its establishment under closed-canopy hardwood forests.

Description: Background: South America is one of the most species diverse continents in the world. Within South America diversity is not distributed evenly at both local and continental scales and this has led to the recognition of various areas with unique species assemblages. Several schemes currently exist which divide the continental-level diversity into large species assemblages referred to as biomes. Here we review five currently available biome maps for South America, including the WWF Ecoregions, the Americas basemap, the Land Cover Map of South America, Morrone's Biogeographic regions of Latin America, and the Ecological Systems Map. The comparison is performed through a case study on the Seasonally Dry Tropical Forest (SDTF) biome using herbarium data of habitat specialist species. Results: Current biome maps of South America perform poorly in depicting SDTF distribution. The poor performance of the maps can be attributed to two main factors: (1) poor spatial resolution, and (2) poor biome delimitation. Poor spatial resolution strongly limits the use of some of the maps in GIS applications, especially for areas with heterogeneous landscape such as the Andes. Whilst the Land Cover Map did not suffer from poor spatial resolution, it showed poor delimitation of biomes. The results highlight that delimiting structurally heterogeneous vegetation is difficult based on remote sensed data alone. A new refined working map of South American SDTF biome is proposed, derived using the Biome Distribution Modelling (BDM) approach where georeferenced herbarium data is used in conjunction with bioclimatic data. Conclusions: Georeferenced specimen data play potentially an important role in biome mapping. Our study shows that herbarium data could be used as a way of ground-truthing biome maps in silico. The results also illustrate that herbarium data can be used to model vegetation maps through predictive modelling. The BDM approach is a promising new method in biome mapping, and could be particularly useful for mapping poorly known, fragmented, or degraded vegetation. We wish to highlight that biome delimitation is not an exact science, and that transparency is needed on how biomes are used as study units in macroevolutionary and ecological research.

Description: Background: Social insects maintain the integrity of their societies by discriminating between colony members and foreigners through cuticular hydrocarbon (CHC) signatures. Nevertheless, parasites frequently get access to social resources, for example through mimicry of host CHCs among other mechanisms. The origin of mimetic compounds, however, remains unknown in the majority of studies (biosynthesis vs. acquisition). Additionally, direct evidence is scarce that chemical mimicry is indeed beneficial to the parasites (e.g., by improving social acceptance). Results: In the present study we demonstrated that the kleptoparasitic silverfish Malayatelura ponerophila most likely acquires CHCs directly from its host ant Leptogenys distinguenda by evaluating the transfer of a stable-isotope label from the cuticle of workers to the silverfish. In a second experiment, we prevented CHC pilfering by separating silverfish from their host for six or nine days. Chemical host resemblance as well as aggressive rejection behaviour by host ants was then quantified for unmanipulated and previously separated individuals. Separated individuals showed reduced chemical host resemblance and they received significantly more aggressive rejection behaviour than unmanipulated individuals. Conclusion: Our study clarifies the mechanism of chemical mimicry in a social insect parasite in great detail. It shows empirically for the first time that social insect parasites are able to acquire CHCs from their host. Furthermore, it demonstrates that the accuracy of chemical mimicry can be crucial for social insect parasites by enhancing social acceptance and, thus, allowing successful exploitation. We discuss the results in the light of coevolutionary arms races between parasites and hosts.

Description: Background: Most species of brown macroalgae recruit exclusively sexually. However, Fucus radicans, a dominant species in the northern Baltic Sea, recruits new attached thalli both sexually and asexually. The level of asexual recruitment varies among populations from complete sexual recruitment to almost (〉90%) monoclonal populations. If phenotypic traits have substantial inherited variation, low levels of sexual activity will decrease population variation in these traits, which may affect function and resilience of the species. We assessed the level of inherited variation in nine phenotypic traits by comparing variation within and among three monoclonal groups and one group of unique multilocus genotypes (MLGs) sampled in the wild. Results: Of the nine phenotypic traits, recovery after freezing, recovery after desiccation, and phlorotannin content showed substantial inherited variation, that is, phenotypic variation in these traits were to a large extend genetically determined. In contrast, variation in six other phenotypic traits (growth rate, palatability to isopod grazers, thallus width, distance between dichotomies, water content after desiccation and photochemical yield under ambient conditions) did not show significant signals of genetic variation at the power of analyses used in the study. Averaged over all nine traits, phenotypic variation within monoclonal groups was only 68% of the variation within the group of different MLGs showing that genotype diversity does affect the overall level of phenotypic variation in this species. Conclusions: Our result indicates that, in general, phenotypic diversity in populations of Fucus radicans increases with increased multilocus genotype (MLG) diversity, but effects are specific for individual traits. In the light of Fucus radicans being a foundation species of the northern Baltic Sea, we propose that increased MLG diversity (leading to increased trait variation) will promote ecosystem function and resilience in areas where F. radicans is common, but this suggestion needs experimental support.

Description: Background: Predicting the geographic distribution of widespread species through modeling is problematic for several reasons including high rates of omission errors. One potential source of error for modeling widespread species is that subspecies and/or races of species are frequently pooled for analyses, which may mask biologically relevant spatial variation within the distribution of a single widespread species. We contrast a presence-only maximum entropy model for the widely distributed oldfield mouse (Peromyscus polionotus) that includes all available presence locations for this species, with two composite maximum entropy models. The composite models either subdivided the total species distribution into four geographic quadrants or by fifteen subspecies to capture spatially relevant variation in P. polionotus distributions. Results: Despite high Area Under the ROC Curve (AUC) values for all models, the composite species distribution model of P. polionotus generated from individual subspecies models represented the known distribution of the species much better than did the models produced by partitioning data into geographic quadrants or modeling the whole species as a single unit. Conclusions: Because the AUC values failed to describe the differences in the predictability of the three modeling strategies, we suggest using omission curves in addition to AUC values to assess model performance. Dividing the data of a widespread species into biologically relevant partitions greatly increased the performance of our distribution model; therefore, this approach may prove to be quite practical and informative for a wide range of modeling applications.

Description: Background: Group coordination is one of the greatest challenges facing animals living in groups. Obligatory trade-offs faced by group members can potentially lead to phenomena at the group level such as the emergence of a leader, consistent structure in the organization of individuals when moving, and the use of visual or acoustic communication. This paper describes the study of collective decision-making at the time of departure (i.e. initiation) for movements of two groups of wild chacma baboons (Papio ursinus). One group was composed of 11 individuals, whilst the other consisted of about 100 individuals. Results: Results for both groups showed that adult males initiated more movements even if the leadership was also distributed to adult females and young individuals. Baboons then joined a movement according to a specific order: adult males and adult females were at the front and the back of the group, sub-adults were at the back and juveniles were located in the central part of the progression. In the two groups, vocalisations, especially loud calls, were more frequently emitted just before the initiation of a group movement, but the frequency of these vocalisations did not influence the success of an initiation in any way. Conclusion: The emergence of a leadership biased towards male group members might be related to their dominance rank and to the fact that they have the highest nutrient requirements in the group. Loud calls are probably not used as recruitment signals but more as a cue concerning the motivation to move, therefore enhancing coordination between group members.

Description: Background: Experiments from a range of ecosystems have shown that insectivorous birds are important in controlling the populations of their invertebrate prey. Here, we report on a large field experiment testing the hypothesis that management for enhancing recreational values in suburban woodlands affects the intensity of bird predation on canopy-living arthropods. Bird exclosures were used in two types of management (understory clearance and dense understory) at two foraging heights in oak Quercus robur canopies and the experiment was replicated at two sites. Results: The biomass and abundance of arthropods were high on net-enclosed branches but strongly reduced on control branches in both types of management. In woods with dense understory, the effect of bird predation on arthropod abundance was about twice as high as in woods with understory clearance. The effect of bird predation on arthropod biomass was not significantly affected by management. Conclusions: Our data provide experimental evidence to support the idea that bird predation on arthropods can be affected by forest management. We suggest that the mechanism is twofold: reduction of bird abundance and shift of foraging behaviour. In urban woodlands, there may be a management trade-off between enhancing recreational values and promoting bird predation rates on arthropods.

Description: Background: The Argentine ant, Linepithema humile, is a widespread invasive ant species that has successfully established in nearly all continents across the globe. Argentine ants are characterised by a social structure known as unicoloniality, where territorial boundaries between nests are absent and intraspecific aggression is rare. This is particularly pronounced in introduced populations and results in the formation of large and spatially expansive supercolonies. Although it is amongst the most well studied of invasive ants, very little work has been done on this ant in South Africa. In this first study, we investigate the population structure of Argentine ants in South Africa. We use behavioural (aggression tests) and chemical (CHC) approaches to investigate the population structure of Argentine ants within the Western Cape, identify the number of supercolonies and infer number of introductions. Results: Both the aggression assays and chemical data revealed that the Western Cape Argentine ant population can be divided into two behaviourally and chemically distinct supercolonies. Intraspecific aggression was evident between the two supercolonies of Argentine ants with ants able to discriminate among conspecific non-nestmates. This discrimination is linked to the divergence in cuticular hydrocarbon profiles of ants originating from the two supercolonies. Conclusions: The presence of these two distinct supercolonies is suggestive of at least two independent introductions of this ant within the Western Cape. Moreover, the pattern of colonisation observed in this study, with the two colonies interspersed, is in agreement with global patterns of Argentine ant invasions. Our findings are of interest because recent studies show that Argentine ants from South Africa are different from those identified in other introduced ranges and therefore provide an opportunity to further understand factors that determine the distributional and spread patterns of Argentine ant supercolonies.

Description: Background: The genetic structure of populations can be influenced by geographic isolation (including physical distance) and ecology. We examined these effects in Leptopilina boulardi, a parasitoid of Drosophila of African origin and widely distributed over temperate and (sub) tropical climates. Results: We sampled 11 populations of L. boulardi from five climatic zones in Iran and measured genetic differentiation at nuclear (Amplified Fragment Length Polymorphism; AFLP) and mitochondrial (Cytochrome Oxidase I; COI) loci. An Analysis of Molecular Variance (AMOVA) for the AFLP data revealed that 67.45 % of variation resided between populations. No significant variation was observed between climatic zones. However, a significant difference was detected between populations from the central (dry) regions and those from the wetter north, which are separated by desert. A similarly clear cut genetic differentiation between populations from the central part of Iran and those from the north was observed by UPGMA cluster analysis and Principal Coordinates Analysis (PCO). Both UPGMA and PCO further separated two populations from the very humid western Caspian Sea coast (zone 3) from other northern populations from the temperate Caspian Sea coastal plain (zone 2), which are connected by forest. One population (Nour) was genetically intermediate between these two zones, indicating some gene flow between these two groups of populations. In all analyses a mountain population, Sorkhabad was found to be genetically identical to those from the nearby coastal plain (zone 2), which indicates high gene flow between these populations over a short geographical distance. One population from the Caspian coast (Astaneh) was genetically highly diverged from all other populations. A partial Mantel test showed a highly significant positive correlation between genetic and geographic distances, as well as separation by the deserts of central Iran. The COI sequences were highly conserved among all populations. Conclusion: The Iranian populations of L. boulardi showed clear genetic structure in AFLP profiles, but not in COI sequence data. The transfer of fruits containing Drosophila larvae parasitized by L. boulardi appears to have caused some unexpected gene flow and changed the genetic composition of populations, particularly in urban areas. Nevertheless, our results suggest that climate, geographic distance and physical barriers may all have contributed to the formation of genetically distinct populations of L. boulardi. Inevitably, there will be overlap between the portions of variance explained by these variables. Disentangling the relative contributions of climate and geography to the genetic structure of this species will require additional sampling.

Description: Background: In colonially breeding birds, the ability to discriminate between individuals is often essential. During post-fledging care, parents have to recognize their own offspring among many other unrelated chicks in the breeding colony. It is well known that fledglings and food-provisioning parents of many bird species use contact calls to convey their identity. These calls are also often used as hunger-related signals of need in young birds. Here, we investigate how such calls incorporate signals of need and at the same time act as reliable indicators of each chick's identity. Results: In a field study, we experimentally manipulated the hunger level of colonially breeding Jackson's golden-backed weaver (Ploceus jacksoni) nestlings close to fledging and investigated its effects on acoustic call parameters. Some acoustic parameters that were related to the time-frequency pattern showed high individuality and were largely unaffected by a nestling's state of hunger. However, the majority of call parameters were significantly affected by hunger. Interestingly, most of these acoustic parameters showed both consistent changes with hunger and high between-individual differences, i.e. potential for individual recognition. Conclusion: The results indicate that individual recognition processes can be based on static, hunger-independent call parameters, but also on dynamic hunger-related parameters that show high individuality. Furthermore, these signal properties suggest that the assessment of signals of need can be improved if the signal value is referenced to a chick's vocal spectrum.

Description: Background: A key challenge for conservation biologists is to determine the most appropriate demographic and genetic management strategies for wildlife populations threatened by disease. We explored this topic by examining whether genetic background and previous pathogen exposure influenced survival of translocated animals when captive-bred and free-ranging bighorn sheep (Ovis canadensis) were used to re-establish a population that had been extirpated in the San Andres Mountains in New Mexico, USA. Results: Although the free-ranging source population had significantly higher multi-locus heterozygosity at 30 microsatellite loci than the captive bred animals, neither source population nor genetic background significantly influenced survival or cause of death. The presence of antibodies to a respiratory virus known to cause pneumonia was associated with increased survival, but there was no correlation between genetic heterozygosity and the presence of antibodies to this virus. Conclusions: Although genetic theory predicts otherwise, increased heterozygosity was not associated with increased fitness (survival) among translocated animals. While heterosis or genetic rescue effects may occur in F1 and later generations as the two source populations interbreed, we conclude that previous pathogen exposure was a more important marker than genetic heterozygosity for predicting survival of translocated animals. Every wildlife translocation is an experiment, and whenever possible, translocations should be designed and evaluated to test hypotheses that will further improve our understanding of how pathogen exposure and genetic variability influence fitness.

Description: Background: Understanding the effects of anthropogenically-driven changes in global temperature, atmospheric carbon dioxide and biodiversity on the functionality of marine ecosystems is crucial for predicting and managing the associated impacts. Coastal ecosystems are important sources of carbon (primary production) to shelf waters and play a vital role in global nutrient cycling. These systems are especially vulnerable to the effects of human activities and will be the first areas impacted by rising sea levels. Within these coastal ecosystems, microalgal assemblages (microphytobenthos: MPB) are vital for autochthonous carbon fixation. The level of in situ production by MPB mediates the net carbon cycling of transitional ecosystems between net heterotrophic or autotrophic metabolism. In this study, we examine the interactive effects of elevated atmospheric CO2 concentrations (370, 600, and 1000 ppmv), temperature (6degreesC, 12degreesC, and 18degreesC) and invertebrate biodiversity on MPB biomass in experimental systems. We assembled communities of three common grazing invertebrates (Hydrobia ulvae, Corophium volutator and Hediste diversicolor) in monoculture and in all possible multispecies combinations. This experimental design specifically addresses interactions between the selected climate change variables and any ecological consequences caused by changes in species composition or richness. Results: The effects of elevated CO2 concentration, temperature and invertebrate diversity were not additive, rather they interacted to determine MPB biomass, and overall this effect was negative. Diversity effects were underpinned by strong species composition effects, illustrating the importance of individual species identity. Conclusions: Overall, our findings suggest that in natural systems, the complex interactions between changing environmental conditions and any associated changes in invertebrate assemblage structure are likely to reduce MPB biomass. Furthermore, these effects would be sufficient to affect the net metabolic balance of the coastal ecosystem, with important implications for system ecology and sustainable exploitation.

Description: Background: Patterns in the association of individuals can shed light on the underlying conditions and processes that shape societies. Here we characterize patterns of association in a population of wild Asian Elephants at Uda Walawe National Park in Sri Lanka. We observed 286 individually-identified adult female elephants over 20 months and examined their social dynamics at three levels of organization: pairs of individuals (dyads), small sets of direct companions (ego-networks), and the population level (complete networks). Results: Corroborating previous studies of this and other Asian elephant populations, we find that the sizes of elephant groups observed in the field on any particular day are typically small and that rates of association are low. In contrast to earlier studies, our longitudinal observations reveal that individuals form larger social units that can be remarkably stable across years while associations among such units change across seasons. Association rates tend to peak in dry seasons as opposed to wet seasons, with some cyclicity at the level of dyads. In addition, we find that individuals vary substantially in their fidelity to companions. At the ego-network level, we find that despite these fluctuations, individuals associate with a pool of long-term companions. At the population level, social networks do not exhibit any clear seasonal structure or hierarchical stratification. Conclusions: This detailed longitudinal study reveals different social dynamics at different levels of organization. Taken together, these results demonstrate that low association rates, seemingly small group sizes, and fission-fusion grouping behavior mask hidden stability in the extensive and fluid social affiliations in this population of Asian elephants.

Description: Background: A territory as a prerequisite for breeding limits the maximum number of breeders in a given area, and thus lowers the proportion of breeders if population size increases. Meanwhile, some territorially breeding animals can have dramatic density fluctuations and little is known about the change from density-dependent processes to density-independendence of breeding during a population increase or an outbreak. We suggest that territoriality, breeding suppression and its break-down can be understood with an incomplete-control model, developed for social breeders and social suppression. Results: We studied density dependence in a microtine species, the bank vole, known as a territorial breeder with cyclic and non-cyclic density fluctuations and periodically high densities in different parts of its range. Our long-term data base from 38 experimental populations in large enclosures in boreal grassland confirms that breeding rates are density-regulated at moderate densities, probably by social suppression of subordinate potential breeders. We conducted an experiment, were we doubled and tripled this moderate density under otherwise the same conditions and measured space use, mortality, reproduction and faecal stress hormone levels (FGM) of adult females. We found that mortality did not differ among the densities, but the regulation of the breeding rate broke down: at double and triple densities all females were breeding, while at the low density the breeding rate was regulated as observed before. Spatial overlap among females increased with density, while a minimum territory size was maintained. Mean stress hormone levels were higher in double and triple densities than at moderate density. Conclusions: At low and moderate densities, breeding suppression was possible by the dominant breeders. But above a density-threshold (similar to a 'competition point'), the dominance of breeders could not be sustained because of the sheer number of non-breeders to be suppressed ('incomplete control'). In our experiment, this point was reached after territories could not shrink any further, but the number of intruders continued to increase. In wild populations, crossing this threshold, possibly by migration or additional cohorts an early onset of breeding in spring, would allow territorial species to escape density-dependency.

Description: Background: Regular seasonal changes in prevalence of infectious diseases are often observed in nature, but the mechanisms are rarely understood. Empirical tests aiming at a better understanding of seasonal prevalence patterns are not feasible for most diseases and thus are widely lacking. Here, we set out to study experimentally the seasonal prevalence in an aquatic host-parasite system. The microsporidian parasite Hamiltosporidium tvarminnensis exhibits pronounced seasonality in natural rock pool populations of its host, Daphnia magna with a regular increase of prevalence during summer and a decrease during winter. An earlier study was, however, unable to test if different starting conditions (initial prevalence) influence the dynamics of the disease in the long term. Here, we aim at testing how the starting prevalence affects the regular prevalence changes over a 4-year period in experimental populations. Results: In an outdoor experiment, populations were set up to include the extremes of the prevalence spectrum observed in natural populations: 5% initial prevalence mimicking a newly invading parasite, 100% mimicking a rock pool population founded by infected hosts only, and 50% prevalence which is commonly observed in natural populations in spring. The parasite exhibited similar prevalence changes in all treatments, but seasonal patterns in the 100% treatment differed significantly from those in the 5% and 50% treatments. Populations started with 5 and 50% prevalence exhibited strong and regular seasonality already in the first year. In contrast, the amplitude of changes in the 100% treatment was low throughout the experiment demonstrating the long-lasting effect of initial conditions on prevalence dynamics. Conclusions: Our study shows that the time needed to approach the seasonal changes in prevalence depends strongly on the initial prevalence. Because individual D. magna populations in this rock pool metapopulation are mostly short lived, only few populations might ever reach a point where the initial conditions are not visible anymore.

Description: Background: Oxygen availability in aquatic habitats is a major environmental factor influencing the ecology, behaviour, and physiology of fishes. This study evaluates the contribution of source population and hypoxic acclimatization of the African fish, Barbus neumayeri, in determining growth and tissue metabolic enzyme activities. Individuals were collected from two sites differing dramatically in concentration of dissolved oxygen (DO), Rwembaita Swamp (annual average DO 1.35 mgO2 L-1) and Inlet Stream West (annual average DO 5.58 mgO2 L-1) in Kibale National Park, Uganda, and reciprocally transplanted using a cage experiment in the field, allowing us to maintain individuals under natural conditions of oxygen, food availability, and flow. Fish were maintained under these conditions for four weeks and sampled for growth rate and the activities of phosphofructokinase (PFK), lactate dehydrogenase (LDH), citrate synthase (CS), and cytochrome c oxidase (CCO) in four tissues, liver, heart, brain, and skeletal muscle. Results: Acclimatization to the low DO site resulted in lower growth rates, lower activities of the aerobic enzyme CCO in heart, and higher activities of the glycolytic enzyme PFK in heart and skeletal muscle. The activity of LDH in liver tissue was correlated with site of origin, being higher in fish collected from a hypoxic habitat, regardless of acclimatization treatment. Conclusions: Our results suggest that the influence of site of origin and hypoxic acclimatization in determining enzyme activity differs among enzymes and tissues, but both factors contribute to higher glycolytic capacity and lower aerobic capacity in B. neumayeri under naturally-occurring conditions of oxygen limitation.

Description: Background: Interference competition occurs when access to resources is negatively affected by the presence of other individuals. Within a species or population, this is known as mutual interference, and it is often modelled with a scaling exponent, m, on the number of predators. Originally, mutual interference was thought to vary along a continuum from prey dependence (no interference; m = 0) to ratio dependence (m = -1), but a debate in the 1990's and early 2000's focused on whether prey or ratio dependence was the better simplification. Some have argued more recently that mutual interference is likely to be mostly intermediate (that is, between prey and ratio dependence), but this possibility has not been evaluated empirically. Results: We gathered estimates of mutual interference from the literature, analyzed additional data, and created the largest compilation of unbiased estimates of mutual interference yet produced. In this data set, both the alternatives of prey dependence and ratio dependence were observed, but only one data set was consistent with prey dependence. There was a tendency toward ratio dependence reflected by a median m of -0.7 and a mean m of -0.8. Conclusions: Overall, the data support the hypothesis that interference is mostly intermediate in magnitude. The data also indicate that interference competition is common, at least in the systems studied to date. Significant questions remain regarding how different factors influence interference, and whether interference can be viewed as a characteristic of a particular population or whether it generally shifts from low to high levels as populations increase in density.

Description: Background: Science-based wildlife management relies on genetic information to infer population connectivity and identify conservation units. The most commonly used genetic marker for characterizing animal biodiversity and identifying maternal lineages is the mitochondrial genome. Mitochondrial genotyping figures prominently in conservation and management plans, with much of the attention focused on the non-coding displacement ("D") loop. We used massively parallel multiplexed sequencing to sequence complete mitochondrial genomes from 40 fishers, a threatened carnivore that possesses low mitogenomic diversity. This allowed us to test a key assumption of conservation genetics, specifically, that the D-loop accurately reflects genealogical relationships and variation of the larger mitochondrial genome. Results: Overall mitogenomic divergence in fishers is exceedingly low, with 66 segregating sites and an average pairwise distance between genomes of 0.00088 across their aligned length (16,290 bp). Estimates of variation and genealogical relationships from the D-loop region (299 bp) are contradicted by the complete mitochondrial genome, as well as the protein coding fraction of the mitochondrial genome. The sources of this contradiction trace primarily to the near-absence of mutations marking the D-loop region of one of the most divergent lineages, and secondarily to independent (recurrent) mutations at two nucleotide position in the D-loop amplicon. Conclusions: Our study has two important implications. First, inferred genealogical reconstructions based on the fisher D-loop region contradict inferences based on the entire mitogenome to the point that the populations of greatest conservation concern cannot be accurately resolved. Whole-genome analysis identifies Californian haplotypes from the northern-most populations as highly distinctive, with a significant excess of amino acid changes that may be indicative of molecular adaptation; D-loop sequences fail to identify this unique mitochondrial lineage. Second, the impact of recurrent mutation appears most acute in closely related haplotypes, due to the low level of evolutionary signal (unique mutations that mark lineages) relative to evolutionary noise (recurrent, shared mutation in unrelated haplotypes). For wildlife managers, this means that the populations of greatest conservation concern may be at the highest risk of being misidentified by D-loop haplotyping. This message is timely because it highlights the new opportunities for basing conservation decisions on more accurate genetic information.

Description: Background: Terabyte-scale collections of string-encoded data are expected from consortia efforts such as the Human Microbiome Project (http://nihroadmap.nih.gov/hmp). Intra- and inter-project data similarity searches are enabled by rapid k-mer matching strategies. Software applications for sequence database partitioning, guide tree estimation, molecular classification and alignment acceleration have benefited from embedded k-mer searches as sub-routines. However, a rapid, general-purpose, open-source, flexible, stand-alone k-mer tool has not been available. Results: Here we present a stand-alone utility, Simrank, which allows users to rapidly identify database strings the most similar to query strings. Performance testing of Simrank and related tools against DNA, RNA, protein and human-languages found Simrank 10X to 928X faster depending on the dataset. Conclusions: Simrank provides molecular ecologists with a high-throughput, open source choice for comparing large sequence sets to find similarity.

Description: Background: Protected areas are the most common and important instrument for the conservation of biological diversity and are called for under the United Nations' Convention on Biological Diversity. Growing human population densities, intensified land-use, invasive species and increasing habitat fragmentation threaten ecosystems worldwide and protected areas are often the only refuge for endangered species. Climate change is posing an additional threat that may also impact ecosystems currently under protection. Therefore, it is of crucial importance to include the potential impact of climate change when designing future nature conservation strategies and implementing protected area management. This approach would go beyond reactive crisis management and, by necessity, would include anticipatory risk assessments. One avenue for doing so is being provided by simulation models that take advantage of the increase in computing capacity and performance that has occurred over the last two decades.Here we review the literature to determine the state-of-the-art in modeling terrestrial protected areas under climate change, with the aim of evaluating and detecting trends and gaps in the current approaches being employed, as well as to provide a useful overview and guidelines for future research. Results: Most studies apply statistical, bioclimatic envelope models and focus primarily on plant species as compared to other taxa. Very few studies utilize a mechanistic, process-based approach and none examine biotic interactions like predation and competition. Important factors like land-use, habitat fragmentation, invasion and dispersal are rarely incorporated, restricting the informative value of the resulting predictions considerably. Conclusion: The general impression that emerges is that biodiversity conservation in protected areas could benefit from the application of modern modeling approaches to a greater extent than is currently reflected in the scientific literature. It is particularly true that existing models have been underutilized in testing different management options under climate change. Based on these findings we suggest a strategic framework for more effectively incorporating the impact of climate change in models exploring the effectiveness of protected areas.

Description: Background: Natural communities are structured by intra-guild competition, predation or parasitism and the abiotic environment. We studied the relative importance of these factors in two host-social parasite ecosystems in three ant communities in Europe (Bavaria) and North America (New York, West Virginia). We tested how these factors affect colony demography, life-history and the spatial pattern of colonies, using a large sample size of more than 1000 colonies. The strength of competition was measured by the distance to the nearest competitor. Distance to the closest social parasite colony was used as a measure of parasitism risk. Nest sites (i.e., sticks or acorns) are limited in these forest ecosystems and we therefore included nest site quality as an abiotic factor in the analysis. In contrast to previous studies based on local densities, we focus here on the positioning and spatial patterns and we use models to compare our predictions to random expectations. Results: Colony demography was universally affected by the size of the nest site with larger and more productive colonies residing in larger nest sites of higher quality. Distance to the nearest competitor negatively influenced host demography and brood production in the Bavarian community, pointing to an important role of competition, while social parasitism was less influential in this community. The New York community was characterized by the highest habitat variability, and productive colonies were clustered in sites of higher quality. Colonies were clumped on finer spatial scales, when we considered only the nearest neighbors, but more regularly distributed on coarser scales. The analysis of spatial positioning within plots often produced different results compared to those based on colony densities. For example, while host and slavemaker densities are often positively correlated, slavemakers do not nest closer to potential host colonies than expected by random. Conclusions: The three communities are differently affected by biotic and abiotic factors. Some of the differences can be attributed to habitat differences and some to differences between the two slavemaking-host ecosystems. The strong effect of competition in the Bavarian community points to the scarcity of resources in this uniform habitat compared to the other more diverse sites. The decrease in colony aggregation with scale indicates fine-scale resource hotspots: colonies are locally aggregated in small groups. Our study demonstrates that species relationships vary across scales and spatial patterns can provide important insights into species interactions. These results could not have been obtained with analyses based on local densities alone. Previous studies focused on social parasitism and its effect on host colonies. The broader approach taken here, considering several possible factors affecting colony demography and not testing each one in isolation, shows that competition and environmental variability can have a similar strong impact on demography and life-history of hosts. We conclude that the effects of parasites or predators should be studied in parallel to other ecological influences.

Description: Background: Plant biodiversity can affect trophic interactions in many ways, including direct bottom-up effects on insects, but is negatively affected by agricultural intensification. Grassland intensification promotes plant productivity, resulting in changes in plant community composition, and impacts on higher trophic levels. Here, we use a novel grassland management experiment combining manipulations of cutting and fertilization with a experimental changes in plant functional group composition (independent of management effects) to disentangle the direct and indirect effects of agricultural management on insect herbivore diversity and abundance. We used leafhoppers as model organisms as they are a key insect taxon in grasslands and react rapidly to management changes. Leafhoppers were sampled between May and September 2010 using standardized sweep netting and pan traps. Results: Plant diversity, functional group composition and management regime in grasslands affect leafhopper species richness and abundance. Higher cutting frequencies directly led to decreasing leafhopper species richness, presumably due to the higher disturbance frequency and the reduction in food-resource heterogeneity. In contrast, fertilizer application had only a small indirect negative effect via enhanced aboveground plant biomass, reduced plant diversity and changes in functional group composition. The manipulated increase in grass cover had contrasting direct and indirect effects on leafhopper species richness: grass cover directly increased leafhopper species richness, but negatively affected plant diversity, which in turn was positively related to leafhopper species richness. In conclusion, insect diversity is driven in complex direct and indirect ways by grassland management including changes in functional group composition. Conclusions: The availability of preferred food sources and the frequency of disturbance are important direct and indirect drivers of leafhopper species richness, interacting in complex ways with plant diversity and food resource heterogeneity.

Description: Background: Animals show consistent individual behavioural patterns over time and over situations. This phenomenon has been referred to as animal personality or behavioural syndromes. Little is known about consistency of animal personalities over entire life times. We investigated the repeatability of behaviour in common voles (Microtus arvalis) at different life stages, with different time intervals, and in different situations. Animals were tested using four behavioural tests in three experimental groups: 1. before and after maturation over three months, 2. twice as adults during one week, and 3. twice as adult animals over three months, which resembles a substantial part of their entire adult life span of several months. Results: Different behaviours were correlated within and between tests and a cluster analysis showed three possible behavioural syndrome-axes, which we name boldness, exploration and activity. Activity and exploration behaviour in all tests was highly repeatable in adult animals tested over one week. In animals tested over maturation, exploration behaviour was consistent whereas activity was not. Voles that were tested as adults with a three-month interval showed the opposite pattern with stable activity but unstable exploration behaviour. Conclusions: The consistency in behaviour over time suggests that common voles do express stable personality over short time. Over longer periods however, behaviour is more flexible and depending on life stage (i.e. tested before/after maturation or as adults) of the tested individual. Level of boldness or activity does not differ between tested groups and maintenance of variation in behavioural traits can therefore not be explained by expected future assets as reported in other studies.

Description: Background: Specialized interactions help structure communities, but persistence of specialized organisms is puzzling because a generalist can occupy more environments and partake in more beneficial interactions. The "Jack-of-all-trades is a master of none" hypothesis asserts that specialists persist because the fitness of a generalist utilizing a particular habitat is lower than that of a specialist adapted to that habitat. Yet, there are many reasons to expect that mutualists will generalize on partners.Plant-soil feedbacks help to structure plant and microbial communities, but how frequently are soil-based symbiotic mutualistic interactions sufficiently specialized to influence species distributions and community composition? To address this question, we quantified realized partner richness and phylogenetic breadth of four wild-grown native legumes (Lupinus bicolor, L. arboreus, Acmispon strigosus and A. heermannii) and performed inoculation trials to test the ability of two hosts (L. bicolor and A. strigosus) to nodulate (fundamental partner richness), benefit from (response specificity), and provide benefit to (effect specificity) 31 Bradyrhizobium genotypes. Results: In the wild, each Lupinus species hosted a broader genetic range of Bradyrhizobium than did either Acmispon species, suggesting that Acmispon species are more specialized. In the greenhouse, however, L. bicolor and A. strigosus did not differ in fundamental association specificity: all inoculated genotypes nodulated both hosts. Nevertheless, A. strigosus exhibited more specificity, i.e., greater variation in its response to, and effect on, Bradyrhizobium genotypes. Lupinus bicolor benefited from a broader range of genotypes but averaged less benefit from each. Both hosts obtained more fitness benefit from symbionts isolated from conspecific hosts; those symbionts in turn gained greater fitness benefit from hosts of the same species from which they were isolated. Conclusions: This study affirmed two important tenets of evolutionary theory. First, as predicted by the Jack-of-all-trades is a master of none hypothesis, specialist A. strigosus obtained greater benefit from its beneficial symbionts than did generalist L. bicolor. Second, as predicted by coevolutionary theory, each test species performed better with partner genotypes isolated from conspecifics. Finally, positive fitness feedback between the tested hosts and symbionts suggests that positive plant-soil feedback could contribute to their patchy distributions in this system.

Description: Background: Introduced earthworms are widespread in forests of North America creating significant negative impacts on forest understory communities. However, much of the reported evidence for negative earthworm effects comes from field investigations either comparing invaded and non-invaded forests or across invasion fronts. While important, such work is rarely able to capture the true effect of earthworms on individual plant species because most forests in North America simultaneously face multiple stressors which may confound earthworm impacts.We used a mesocosm experiment to isolate effects of the anecic introduced earthworm, Lumbricus terrestris L. on seedlings of 14 native plant species representing different life form groups (perennial herb, graminoid, and tree). Results: Earthworm presence did not affect survival, fertility or biomass of any of the seedling plant species tested over a 17-week period. However, L. terrestris presence significantly decreased growth of two sedges (Carex retroflexa Muhl. ex Willd. and Carex radiata (Wahlenb.) Small) by decreasing the number of culms. Conclusions: Our mesocosm results with seedlings contrast with field reports indicating extensive and significant negative effects of introduced earthworms on many mature native forbs, and positive effects on sedges. We suggest that earthworm impacts are context- and age-specific and that generalizations about their impacts are potentially misleading without considering and manipulating other associated factors.

Description: Background: Movements of animals have important consequences, at both the individual and population levels. Due to its important implications in the evolutionary dynamics of populations, dispersal is one of the most studied types of movement. In contrast, non-permanent extra home-range movements are often paid less attention. However, these movements may occur in response to important biological processes such as mating or predation avoidance. In addition, these forays are often preludes to permanent dispersal, because they may help individuals gain cues about their surroundings prior to settlement in a new place.In the European hare, exploration forays occur predominantly in juveniles, the time at which most hares disperse. In France, the timing of dispersal also overlaps with the hare hunting period. However, the determinants of such behaviour have not yet been studied. Herein, we investigate whether these non-permanent explorations are dispersal attempts/preludes or, in contrast, whether they are triggered by other factors such as disturbances related to hunting. Results: Contrary to natal dispersal, we did not find strong male-bias in the propensity to engage in explorations. Exploration forays occurred less in juveniles than in adults and later in the season than natal dispersal. This was the case both for philopatric movements and for movements occurring after dispersal and settlement. These movements were also more likely to occur during the hare hunting period and the mating season. Conclusions: We suggest that explorations in hares are triggered by factors other than dispersal and that hares may respond to hunting disturbances. Overall, we emphasize the need to account for human-related predation risk as a factor driving space-use in harvested species.

Description: As part of BioMed Central’s open science mission, we are pleased to announce that two of our journals have integrated with the open data repository Dryad. Authors submitting their research to either BMC Ecology or BMC Evolutionary Biology will now have the opportunity to deposit their data directly into the Dryad archive and will receive a permanent, citable link to their dataset. Although this does not affect any of our current data deposition policies at these journals, we hope to encourage a more widespread adoption of open data sharing in the fields of ecology and evolutionary biology by facilitating this process for our authors. We also take this opportunity to discuss some of the wider issues that may concern researchers when making their data openly available. Although we offer a number of positive examples from different fields of biology, we also recognise that reticence to data sharing still exists, and that change must be driven from within research communities in order to create future science that is fit for purpose in the digital age. This editorial was published jointly in both BMC Ecology and BMC Evolutionary Biology.

Description: Background: Studies of host-parasite interactions have the potential to provide insights into the ecology of both organisms involved. We monitored the movement of sucking lice (Lemurpediculus verruculosus), parasites that require direct host-host contact to be transferred, in their host population of wild mouse lemurs (Microcebus rufus). These lemurs live in the rainforests of Madagascar, are small (40 g), arboreal, nocturnal, solitary foraging primates for which data on population-wide interactions are difficult to obtain. We developed a simple, cost effective method exploiting the intimate relationship between louse and lemur, whereby individual lice were marked, without removal from their host, with an individualized code, and tracked throughout the lemur population. We then tested the hypotheses that 1) the frequency of louse transfers, and thus interactions, would decrease with increasing distance between paired individual lemurs; 2) due to host polygynandry, social interactions and hence louse transfers would increase during the onset of the breeding season; and 3) individual mouse lemurs would vary in their contributions to the spread of lice. Results: We show that louse transfers involved 43.75% of the studied lemur population, exclusively males. Louse transfers peaked during the breeding season, perhaps due to increased social interactions between lemurs. Although trap-based individual lemur ranging patterns are restricted, louse transfer rate does not correlate with the distance between lemur trapping locales, indicating wider host ranging behavior and a greater risk of rapid population-wide pathogen transmission than predicted by standard trapping data alone. Furthermore, relatively few lemur individuals contributed disproportionately to the rapid spread of lice throughout the population. Conclusions: Using a simple method, we were able to visualize exchanges of lice in a population of cryptic wild primates. This method not only provided insight into the previously unseen parasite movement between lemurs, but also allowed us to infer social interactions between them. As lice are known pathogen vectors, our method also allowed us to identify the lemurs most likely to facilitate louse-mediated epidemics. Our approach demonstrates the potential to uncover otherwise inaccessible parasite-host, and host social interaction data in any trappable species parasitized by sucking lice.

Description: Background: Disturbance is an important process structuring ecosystems worldwide and has long been thought to be a significant driver of diversity and dynamics. In forests, most studies of disturbance have focused on large-scale disturbances such as hurricanes or tree-falls. However, smaller sub-canopy disturbances could also have significant impacts on community structure. One such sub-canopy disturbance in tropical forests is abscising leaves of large arborescent palm (Arecaceae) trees. These leaves can weigh up to 15 kg and cause physical damage and mortality to juvenile plants. Previous studies examining this question suffered from the use of static data at small spatial scales. Here we use data from a large permanent forest plot combined with dynamic data on the survival and growth of 〉66,000 individuals over a seven-year period to address whether falling palm fronds do impact neighbouring seedling and sapling communities, or whether there is an interaction between the palms and peccaries rooting for fallen palm fruit in the same area as falling leaves. We tested the wider generalisation of these hypotheses by comparing seedling and sapling survival under fruiting and non-fruiting trees in another common Neotropical family, the Myristicaceae. Results: We found a spatially-restricted but significant effect of large arborescent fruiting palms on the spatial structure, population dynamics and species diversity of neighbouring sapling and seedling communities. However, these effects were not found around slightly smaller non-fruiting palm trees, suggesting it is seed predators such as peccaries rather than falling leaves that impact on the communities around palm trees. Conversely, this hypothesis was not supported in data from other edible species in the family Myristicaceae. Conclusions: Given the abundance of arborescent palm trees in Amazonian forests, it is reasonable to conclude that their presence does have a significant, if spatially-restricted, impact on juvenile plants, most likely on the survival and growth of seedlings and saplings damaged by foraging peccaries. Given the abundance of fruit produced by each palm, the widespread effects of these small-scale disturbances appear, over long time-scales, to cause directional changes in community structure at larger scales.

Description: Background: When a specimen belongs to a species not yet represented in DNA barcode reference libraries there is disagreement over the effectiveness of using sequence comparisons to assign the query accurately to a higher taxon. Library completeness and the assignment criteria used have been proposed as critical factors affecting the accuracy of such assignments but have not been thoroughly investigated. We explored the accuracy of assignments to genus, tribe and subfamily in the Sphingidae, using the almost complete global DNA barcode reference library (1095 species) available for this family. Costa Rican sphingids (118 species), a well-documented, diverse subset of the family, with each of the tribes and subfamilies represented were used as queries. We simulated libraries with different levels of completeness (10-100% of the available species), and recorded assignments (positive or ambiguous) and their accuracy (true or false) under six criteria. Results: A liberal tree-based criterion assigned 83% of queries accurately to genus, 74% to tribe and 90% to subfamily, compared to a strict tree-based criterion, which assigned 75% of queries accurately to genus, 66% to tribe and 84% to subfamily, with a library containing 100% of available species (but excluding the species of the query). The greater number of true positives delivered by more relaxed criteria was negatively balanced by the occurrence of more false positives. This effect was most sharply observed with libraries of the lowest completeness where, for example at the genus level, 32% of assignments were false positives with the liberal criterion versus

Description: Background: Efficient division of reproductive labor is a crucial characteristic of social insects and underlies their ecological and evolutionary success. Despite of the harmonious appearance of insect societies, nestmates may have different interests concerning the partitioning of reproduction among group members. This may lead to conflict about reproductive rights. As yet, few studies have investigated the allocation of reproduction among queens in multi - queen societies ("reproductive skew"). In the ant Leptothorax acervorum, reproductive skew varies considerably among populations. While reproduction is quite equally shared among nestmate queens in most populations from boreal Eurasia (low skew), colonies from populations at the edge of the species' range are characterized by "functional monogyny," i.e., high skew. The proximate mechanisms underlying high skew, in particular how workers influence which queen lays eggs, are not well understood. We investigated the behavior of queens and workers in functionally monogynous colonies of L. acervorum from two mountain ranges in central Spain. Results: We provide evidence for both queen and worker influence on the outcome of conflict over reproduction in colonies of L. acervorum from Spain. The patterns of queen - queen aggression and worker - queen grooming and feeding after hibernation allowed predicting, which queen later began to lay eggs. In contrast, worker aggression towards queens was not clearly associated with a queen's future reproductive success. Queen - queen and worker - queen aggression differed in quality: queens typically engaged in ritualized dominance behavior, such as antennal boxing, while workers also attacked queens by biting and prolonged pulling on their legs and antennae. In several cases, overt worker aggression led to the expulsion of queens from the nest or their death. Conclusion: We conclude that queens of L. acervorum from Spain establish rank orders by ritualized dominance interactions, such as antennal boxing. Workers may reinforce these hierarchies by preferentially feeding and grooming high ranking queens and attacking lower ranking queens. Aggressive worker policing may thus stabilize functional monogyny. Optimal skew models predict that high skew in ants is associated with high dispersal costs. In central Spain, L. acervorum is restricted to small patches at higher elevations, which presumably makes dispersal and colony founding difficult. Because of the ecological requirements of L. acervorum and the predicted large impact of global change on central Spain, the functionally monogynous populations of this ant must be considered as threatened.

Description: Background: Bacillus anthracis, the causative agent of anthrax, is a globally distributed zoonotic pathogen that continues to be a veterinary and human health problem in Central Asia. We used a database of anthrax outbreak locations in Kazakhstan and a subset of genotyped isolates to model the geographic distribution and ecological associations of B. anthracis in Kazakhstan. The aims of the study were to test the influence of soil variables on a previous ecological niche based prediction of B. anthracis in Kazakhstan and to determine if a single sub-lineage of B. anthracis occupies a unique ecological niche. Results: The addition of soil variables to the previously developed ecological niche model did not appreciably alter the limits of the predicted geographic or ecological distribution of B. anthracis in Kazakhstan. The A1.a experiment predicted the sub-lineage to be present over a larger geographic area than did the outbreak based experiment containing multiple lineages. Within the geographic area predicted to be suitable for B. anthracis by all ten best subset models, the A1.a sub-lineage was associated with a wider range of ecological tolerances than the outbreak-soil experiment. Analysis of rule types showed that logit rules predominate in the outbreak-soil experiment and range rules in the A1.a sub-lineage experiment. Random sub-setting of locality points suggests that models of B. anthracis distribution may be sensitive to sample size. Conclusions: Our analysis supports careful consideration of the taxonomic resolution of data used to create ecological niche models. Further investigations into the environmental affinities of individual lineages and sub-lineages of B. anthracis will be useful in understanding the ecology of the disease at large and small scales. With model based predictions serving as approximations of disease risk, these efforts will improve the efficacy of public health interventions for anthrax prevention and control.

Description: Background: The Svalbard endemic aphid Acyrthosiphon svalbardicum (Heikinheimo, 1968) is host specific to Dryas octopetala L. ssp octopetala (Rosaceae). It has been hypothesized that the aphid is present on those areas with a thin winter snow cover and which therefore clear of snow earlier in the season. This early snow clearance results in a longer growing period and allows the aphid to experience at least the minimum number of degree days required to complete its life cycle. However, this hypothesis lacked a detailed field validation. We aimed to test the relationship between the aphid distribution and time of snow clearance at landscape scale, mapping snow depth at peak of snow accumulation for the two succeeding years 2009 and 2010 and examining site occupancy and plant phenology the following summers. Additionally, the distribution range mapped by Strathdee & Bale (1995) was revisited to address possible changes in range along the coast of the fjord. Results: A linear relation between snow depth and timing of snow melt was found but with strong inter-annual and landscape variation. Both snow depth and plant phenology were found to affect patch occupancy. In August, the aphid, at the three life stages scored (viviparae, oviparae/males and eggs), was present most frequently in those D. octopetala patches with the most advanced plant phenology and which showed shallower snow depths in spring. However, many patches predicted to contain aphids were empty. The aphid distribution range has expanded 4.7 km towards the fjord mouth from 1995. Conclusions: Snow depth alone, and hence date of snow clearance, cannot precisely define species distribution at landscape scale, as this cannot explain why are they unoccupied patches under shallow snow depths with advanced plant phenology. We nonetheless present a model Arctic system that could form the basis for long term monitoring for climate- driven species shifts.

Description: Background: There are a growing number of genomes sequenced with tentative functions assigned to a large proportion of the individual genes. Model organisms in laboratory settings form the basis for the assignment of gene function, and the ecological context of gene function is lacking. This work addresses this shortcoming by investigating expressed genes of sockeye salmon (Oncorhynchus nerka) muscle tissue. We compared morphology and gene expression in natural juvenile sockeye populations related to river and lake habitats. Based on previously documented divergent morphology, feeding strategy, and predation in association with these distinct environments, we expect that burst swimming is favored in riverine population and continuous swimming is favored in lake-type population. In turn we predict that morphology and expressed genes promote burst swimming in riverine sockeye and continuous swimming in lake-type sockeye. Results: We found the riverine sockeye population had deep, robust bodies and lake-type had shallow, streamlined bodies. Gene expression patterns were measured using a 16k microarray, discovering 141 genes with significant differential expression. Overall, the identity and function of these genes was consistent with our hypothesis. In addition, Gene Ontology (GO) enrichment analyses with a larger set of differentially expressed genes found the "biosynthesis" category enriched for the riverine population and the "metabolism" category enriched for the lake-type population. Conclusions: This study provides a framework for understanding sockeye life history from a transcriptomic perspective and a starting point for more extensive, targeted studies determining the ecological context of genes.

Description: Background: The Andes-Amazon basin of Peru and Bolivia is one of the most data-poor, biologically rich, and rapidly changing areas of the world. Conservation scientists agree that this area hosts extremely high endemism, perhaps the highest in the world, yet we know little about the geographic distributions of these species and ecosystems within country boundaries. To address this need, we have developed conservation data on endemic biodiversity (~800 species of birds, mammals, and amphibians) and terrestrial ecological systems (~90; groups of vegetation communities resulting the action of ecological processes, substrates, and/or environmental gradients) with which we conducted a fine scale conservation prioritization across the Amazon watershed of Peru and Bolivia. We modelled the geographic distributions of 435 endemic plants and all 347 endemic vertebrate species, from existing museum and herbaria specimens at a regional conservation practitioner's scale (1:250,000-1:1,000,000), based on the best available tools and geographic data. We mapped ecological systems, endemic species concentrations, and irreplaceable areas with respect to national level protected areas. Results: We found that sizes of endemic species distributions ranged widely ( 〈 20 km2 to 〉 200,000 km2) across the study area. Bird and mammal endemic species richness was greatest within a narrow 2500-3000 m elevation band along the length of the Andes Mountains. Endemic amphibian richness was highest at 1000-1500 m elevation and concentrated in the southern half of the study area. Geographical distribution of plant endemism was highly taxon-dependent. Irreplaceable areas, defined as locations with the highest number of species with narrow ranges, overlapped slightly with areas of high endemism, yet generally exhibited unique patterns across the study area by species group. We found that many endemic species and ecological systems are lacking national-level protection; a third of endemic species have distributions completely outside of national protected areas. Protected areas cover only 20% of areas of high endemism and 20% of irreplaceable areas. Almost 40% of the 91 ecological systems are in serious need of protection (= 〈 2% of their ranges protected). Conclusions: We identify for the first time, areas of high endemic species concentrations and high irreplaceability that have only been roughly indicated in the past at the continental scale. We conclude that new complementary protected areas are needed to safeguard these endemics and ecosystems. An expansion in protected areas will be challenged by geographically isolated micro-endemics, varied endemic patterns among taxa, increasing deforestation, resource extraction, and changes in climate. Relying on pre-existing collections, publically accessible datasets and tools, this working framework is exportable to other regions plagued by incomplete conservation data.

Description: Background: Interactions between aboveground and belowground terrestrial communities are often mediated by plants, with soil organisms interacting via the roots and aboveground organisms via the shoots and leaves. Many studies now show that plant genetics can drive changes in the structure of both above and belowground communities; however, the role of plant genetic variation in mediating aboveground-belowground interactions is still unclear. We used an earthworm-plant-aphid model system with two aphid species (Aphis fabae and Acyrthosiphon pisum) to test the effect of host-plant (Vicia faba) genetic variation on the indirect interaction between the belowground earthworms (Eisenia veneta) on the aboveground aphid populations. Results: Our data shows that host-plant variety mediated an indirect ecological effect of earthworms on generalist black bean aphids (A. fabae), with earthworms increasing aphid growth rate in three plant varieties but decreasing it in another variety. We found no effect of earthworms on the second aphid species, the pea aphid (A. pisum), and no effect of competition between the aphid species. Plant biomass was increased when earthworms were present, and decreased when A. pisum was feeding on the plant (mediated by plant variety). Although A. fabae aphids were influenced by the plants and worms, they did not, in turn, alter plant biomass. Conclusions: Previous work has shown inconsistent effects of earthworms on aphids, but we suggest these differences could be explained by plant genetic variation and variation among aphid species. This study demonstrates that the outcome of belowground-aboveground interactions can be mediated by genetic variation in the host-plant, but depends on the identity of the species involved.

Description: Background: A remarkable range of environmental conditions is present in the Hawaiian Islands due to their gradients of elevation, rainfall and island age. Despite being well known as a location for the study of evolutionary processes and island biogeography, little is known about the composition of the non-marine algal flora of the archipelago, its degree of endemism, or affinities with other floras. We conducted a biodiversity survey of the non-marine macroalgae of the six largest main Hawaiian Islands using molecular and microscopic assessment techniques. We aimed to evaluate whether endemism or cosmopolitanism better explain freshwater algal distribution patterns, and provide a baseline data set for monitoring future biodiversity changes in the Hawaiian Islands. Results: 1,786 aquatic and terrestrial habitats and 1,407 distinct collections of non-marine macroalgae were collected from the islands of Kauai, Oahu, Molokai, Maui, Lanai and Hawaii from the years 2009?2014. Targeted habitats included streams, wet walls, high elevation bogs, taro fields, ditches and flumes, lakes/reservoirs, cave walls and terrestrial areas. Sites that lacked freshwater macroalgae were typically terrestrial or wet wall habitats that were sampled for diatoms and other microalgae. Approximately 50% of the identifications were of green algae, with lesser proportions of diatoms, red algae, cyanobacteria, xanthophytes and euglenoids. 898 DNA sequences were generated representing eight different markers, which enabled an assessment of the number of taxonomic entities for genera collected as part of the survey. Forty-four well-characterized taxa were assessed for global distribution patterns. This analysis revealed no clear biogeographic affinities of the flora, with 27.3% characterized as ?cosmopolitan?, 11.4% ?endemic?, and 61.3% as intermediate. Conclusions: The Hawaiian freshwater algal biodiversity survey represents the first comprehensive effort to characterize the non-marine algae of a tropical region in the world using both morphological and molecular tools. Survey data were entered in the Hawaiian Freshwater Algal Database, which and serves as a digital repository of photographs and micrographs, georeferenced localities and DNA sequence data. These analyses yielded an updated checklist of the non-marine macroalgae of the Hawaiian Islands, and revealed varied biogeographic affinities of the flora that are likely a product of both natural and anthropogenic dispersal.

Description: Background: Nutrient deficiency affects the growth and population dynamics of consumers. Endoparasites can be seen as consumers that drain carbon (C) or energy from their host while simultaneously competing for limiting resources such as phosphorus (P). Depending on the relative demands of the host and the parasite for the limiting nutrient, intensified resource competition under nutrient limitation can either reduce the parasite?s effect on the host or further reduce the fitness of the nutrient-limited host. So far, knowledge of how nutrient limitation affects parasite performance at the host population level and how this affects the host populations is limited. Results: We followed the population growth of Daphnia magna that were uninfected or experimentally infected with a microsporidian, Glugoides intestinalis. The Daphnia were fed either P-sufficient or P-limited algae. The P-limited diet decreased the population density and biomass compared with the populations fed with the P-sufficient algae. In the P-sufficient populations, infection with the parasite reduced the population density but not the biomass of Daphnia, while in the P-limited populations, both the density and biomass of Daphnia decreased toward the end of the 32?day experiment compared with the uninfected controls. The infected animals from the P-limited populations had higher parasite spore cluster counts, while, in a separate experiment, host diet quality did not affect the number of parasites in individually kept Daphnia. Conclusions: Because host diet quality did not affect parasite numbers at the individual level, we suggest that the higher parasite load in the P-limited populations is a result of feedback effects arising at the population level. Because of the density-dependent transmission of the parasite and the time lag between exposure and transmission, the lower host population density in the P-limited populations led to a higher spore:host ratio. This effect may have been further reinforced by decreases in filtration rates caused by crowding in the P-sufficient populations and/or increases in filtration rates as a response to poor food quality in the P-limited populations. The increases in exposure led to a higher parasite load and aggravated the negative effects of parasite infection at the population level.

Description: Background: Most vertebrates experience coinfections, and many pathogen-pathogen interactions occur indirectly through the host immune system. These interactions are particularly strong in mixed micro-macroparasite infections because of immunomodulatory effects of helminth parasites. While these trade-offs have been examined extensively in laboratory animals, few studies have examined them in natural systems. Additionally, many wildlife pathogens fluctuate seasonally, at least partly due to seasonal host immune changes. We therefore examined seasonality of immune resource allocation, pathogen abundance and exposure, and interactions between infections and immunity in plains zebra (Equus quagga) in Etosha National Park (ENP), Namibia, a system with strongly seasonal patterns of gastrointestinal (GI) helminth infection intensity and concurrent anthrax outbreaks. Both pathogens are environmentally transmitted, and helminth seasonality is driven by environmental pressures on free living life stages. The reasons behind anthrax seasonality are currently not understood, though anthrax is less likely directly driven by environmental factors. Results: We measured a complex, interacting set of variables and found evidence that GI helminth infection intensities, eosinophil counts, IgE and IgGb antibody titers, and possibly IL-4 cytokine signaling were increased in wetter seasons, and that ectoparasite infestations and possibly IFN-γ cytokine signaling were increased in drier seasons. Monocyte counts and anti-anthrax antibody titers were negatively associated with wet season eosinophilia, and monocytes were negatively correlated with IgGb and IgE titers. Taken together, this supports the hypothesis that ENP wet seasons are characterized by immune resource allocation toward Th-2 type responses, while Th1-type immunity may prevail in drier seasons, and that hosts may experience Th1-Th2 trade-offs. We found evidence that this Th2-type resource allocation is likely driven by GI parasite infections, and that these trade-offs may render hosts less capable of concurrently mounting effective Th1-type immune responses against anthrax. Conclusions: This study is one of the first to examine laboratory-demonstrated Th1-Th2 trade-offs in a natural system. It provides evidence that seasonally bound pathogens may affect, through immunology, transmission dynamics of pathogens that might otherwise not be seasonally distributed. It suggests that, by manipulating the internal host ecosystem, GI parasites may influence the external ecosystem by affecting the dynamics of another environmentally transmitted pathogen.

Description: Background: The proto-periodical cicada Okanagana rimosa is subject to infection by the acoustically orientating parasitoid fly Emblemasoma auditrix. Furthermore, it is also the only known host of E. auditrix. Here we test the question, whether the highly adapted parasitoid can also infect other cicadas, like the periodical cicada (Magicicada cassinii) and which steps of the parasitization process can be completed. The experiments might also reveal whether such a parasitoid could hypothetically have been involved in the evolution of periodicity. Results: The hearing threshold of E. auditrix matches with the spectrum of the calling song of M. cassinii, indicating potential host localization. Behaviourally, host localization is possible by the parasitoid as it approaches a loudspeaker broadcasting the buzz part of the calling song of M. cassinii. Magicicada cassinii is readily accepted as host and for host infection the parasitoid uses the same behavioural sequence as for its host O. rimosa. A larva is deposited into the timbal of the cicada. By contrast to O. rimosa the development of the fly larva is delayed and eventually suppressed in M. cassinii. Conclusions: The host range of E. auditrix is mainly determined by acoustic parameters. This filter is important, as other sensory cues seem not to be involved in the host selection process and larva will not develop in unsuited host. Although the recent parasitoid-host system seems to be stable in terms of coexistence of both species, an acoustically hunting parasitoid could have been a selective force during evolution of prime numbered periodicity in cicadas.

Description: Background: Studying the drivers and determinants of species, population and community spatial patterns is central to ecology. The observed structure of community assemblages is the result of deterministic abiotic (environmental constraints) and biotic factors (positive and negative species interactions), as well as stochastic colonization events (historical contingency). We analyzed the role of multi-scale spatial component of soil environmental variability in structuring earthworm assemblages in a gallery forest from the Colombian ?Llanos.? We aimed to disentangle the spatial scales at which species assemblages are structured and determine whether these scales matched those expressed by soil environmental variables. We also tested the hypothesis of the ?single tree effect? by exploring the spatial relationships between root-related variables and soil nutrient and physical variables in structuring earthworm assemblages. Multivariate ordination techniques and spatially explicit tools were used, namely cross-correlograms, Principal Coordinates of Neighbor Matrices (PCNM) and variation partitioning analyses. Results: The relationship between the spatial organization of earthworm assemblages and soil environmental parameters revealed explicitly multi-scale responses. The soil environmental variables that explained nested population structures across the multi-spatial scale gradient differed for earthworms and assemblages at the very-fine- ( 30?m), fine (10?20?m) and very fine scales (

Description: Background: The division of labor in social insect colonies involves transitions by workers from one task to another and is critical to the organization and ecological success of colonies. The differential regulation of genetic pathways is likely to be a key mechanism involved in plasticity of social insect task behavior. One of the few pathways implicated in social organization involves the cGMP-activated protein kinase gene, foraging, a gene associated with foraging behavior in social insect species. The association of the foraging gene with behavior is conserved across diverse species, but the observed expression patterns and proposed functions of this gene vary across taxa. We compared the protein sequence of foraging across social insects and explored whether the differential regulation of this gene is associated with task behaviors in the harvester ant, Pogonomyrmex occidentalis. Results: Phylogenetic analysis of the coding region of the foraging gene reveals considerable conservation in protein sequence across insects, particularly among hymenopteran species. The absence of amino acid variation in key active and binding sites suggests that differences in behaviors associated with this gene among species may be the result of changes in gene expression rather than gene divergence. Our results also indicate that differential gene expression of foraging may regulate behavioral plasticity within species. Using real time qPCR analyses with a harvester ant ortholog to foraging (Pofor), we found that the brains of harvester ant foragers have a daily fluctuation in expression of foraging with mRNA levels peaking at midday. In contrast, young workers inside the nest have low levels of Pofor mRNA with no evidence of daily fluctuations in expression. As a result, the association of foraging expression with task behavior within a species changes depending on the time of day the individuals are sampled. Conclusions: The amino acid protein sequence of foraging is highly conserved across social insects. Differences in foraging behaviors associated with this gene among social insect species are likely due to differences in gene regulation rather than evolutionary changes in the encoded protein. Within the harvester ant species, P. occidentalis, foraging behavior is associated with foraging gene expression and the daily regulation of this gene is task-dependent. The task-specific expression patterns of foraging appear to be consistent with the task-specific circadian rhythms observed in harvester ants. Whether the molecular clock plays a role in regulating foraging gene expression (or vice versa) remains to be determined. Our results represent the first time series analysis of foraging gene expression and underscore the importance of assaying time-related expression differences in behavioral studies. Understanding how this gene is regulated within species is critical to explaining the mechanism by which foraging influences behavior.

Description: Background: Biodiversity databases serve the important role of highlighting species-level diversity from defined geographical regions. Databases that are specially designed to accommodate the types of data gathered during regional surveys are valuable in allowing full data access and display to researchers not directly involved with the project, while serving as a Laboratory Information Management System (LIMS). The Hawaiian Freshwater Algal Database, or HfwADB, was modified from the Hawaiian Algal Database to showcase non-marine algal specimens collected from the Hawaiian Archipelago by accommodating the additional level of organization required for samples including multiple species.DescriptionThe Hawaiian Freshwater Algal Database is a comprehensive and searchable database containing photographs and micrographs of samples and collection sites, geo-referenced collecting information, taxonomic data and standardized DNA sequence data. All data for individual samples are linked through unique 10-digit accession numbers ("Isolate Accession"), the first five of which correspond to the collection site ("Environmental Accession"). Users can search online for sample information by accession number, various levels of taxonomy, habitat or collection site. HfwADB is hosted at the University of Hawaii, and was made publicly accessible in October 2011. At the present time the database houses data for over 2,825 samples of non-marine algae from 1,786 collection sites from the Hawaiian Archipelago. These samples include cyanobacteria, red and green algae and diatoms, as well as lesser representation from some other algal lineages. Conclusions: HfwADB is a digital repository that acts as a Laboratory Information Management System for Hawaiian non-marine algal data. Users can interact with the repository through the web to view relevant habitat data (including geo-referenced collection locations) and download images of collection sites, specimen photographs and micrographs, and DNA sequences. It is publicly available at http://algae.manoa.hawaii.edu/hfwadb/.

Description: Background: Morphological data suggest that, unlike most other groups of marine organisms, scleractinian corals of the genus Stylophora are more diverse in the western Indian Ocean and in the Red Sea than in the central Indo-Pacific. However, the morphology of corals is often a poor predictor of their actual biodiversity: hence, we conducted a genetic survey of Stylophora corals collected in Madagascar, Okinawa, the Philippines and New Caledonia in an attempt to find out the true number of species in these various locations. Results: A molecular phylogenetic analysis of the mitochondrial ORF and putative control region concurs with a haploweb analysis of nuclear ITS2 sequences in delimiting three species among our dataset: species A and B are found in Madagascar whereas species C occurs in Okinawa, the Philippines and New Caledonia. Comparison of ITS1 sequences from these three species with data available online suggests that species C is also found on the Great Barrier Reef, in Malaysia, in the South China Sea and in Taiwan, and that a distinct species D occurs in the Red Sea. Shallow-water morphs of species A correspond to the morphological description of Stylophora madagascarensis, species B presents the morphology of Stylophora mordax, whereas species C comprises various morphotypes including Stylophora pistillata and Stylophora mordax. Conclusions: Genetic analysis of the coral genus Stylophora reveals species boundaries that are not congruent with morphological traits. Of the four hypotheses that may explain such discrepancy (phenotypic plasticity, morphological stasis, morphological convergence, and interspecific hybridization), the first two appear likely to play a role but the fourth one is rejected since mitochondrial and nuclear markers yield congruent species delimitations. The position of the root in our molecular phylogenies suggests that the center of origin of Stylophora is located in the western Indian Ocean, which probably explains why this genus presents a higher biodiversity in the westernmost part of its area of distribution than in the "Coral Triangle".

Description: Background: Development rates of Aedes aegypti are known to vary with respect to many abiotic and biotic factors including temperature, resource availability, and intraspecific competition. The relative importance of these factors and their interactions are not well established across populations. We performed meta-analysis on a dataset of development rate estimates from 49 studies. Results: Meta-analytic results indicated that the environmental factor of temperature is sufficient to explain development rate variability in Ae. aegypti. While diet and density may greatly impact other developmental phenotypes, these results suggest that for development rate these factors should never be considered to the exclusion of temperature. The effect of temperature on development rate is not homogenous or constant. The sources of heterogeneity of the effect of temperature are difficult to analyze due to lack of consistent reporting of larval rearing methods. Conclusions: Temperature is the most important ecological determinant of development rate in Ae. aegypti, but its effect is heterogeneous. Ignoring this heterogeneity is problematic for models of vector population and vector-borne disease transmission.

Description: Background: Although the nematode Caenorhabditis elegans is a major model organism in diverse biological areas and well studied under laboratory conditions, little is known about its ecology. Therefore, characterization of the species' natural habitats should provide a new perspective on its otherwise well-studied biology. The currently best characterized populations are in France, demonstrating that C. elegans prefers nutrient- and microorganism-rich substrates such as rotting fruits and decomposing plant matter. In order to extend these findings, we sampled C. elegans continuously across 1.5 years from rotting apples and compost heaps in three North German locations. Results: C. elegans was found throughout summer and autumn in both years. It shares its habitat with the related nematode species C. remanei, which could thus represent an important competitor for a similar ecological niche. The two species were isolated from the same site, but rarely the same substrate sample. In fact, C. elegans was mainly found on compost and C. remanei on rotten apples, possibly suggesting niche separation. The occurrence of C. elegans itself was related to environmental humidity and rain, although the correlation was significant for only one sampling site each. Additional associations between nematode prevalence and abiotic parameters could not be established. Conclusions: Taken together, our findings vary from the previous results for French C. elegans populations in that the considered German populations always coexisted with the congeneric species C. remanei (rather than C. briggsae as in France) and that C. elegans prevalence can associate with humidity and rain (rather than temperature, as suggested for French populations). Consideration of additional locations and time points is thus essential for full appreciation of the nematode's natural ecology.

Description: Background: Non-essential facultative endosymbionts can provide their hosts with protection from parasites, pathogens, and predators. For example, two facultative bacterial symbionts of the pea aphid (Acyrthosiphon pisum), Serratia symbiotica and Hamiltonella defensa, protect their hosts from parasitism by two species of parasitoid wasp. Previous studies have not explored whether facultative symbionts also play a defensive role against predation in this system. We tested whether feeding on aphids harboring different facultative symbionts affected the fitness of an aphid predator, the lady beetle Hippodamia convergens. Results: While these aphid faculative symbionts did not deter lady beetle feeding, they did decrease survival of lady beetle larvae. Lady beetle larvae fed a diet of aphids with facultative symbionts had significantly reduced survival from egg hatching to pupation and therefore had reduced survival to adult emergence. Additionally, lady beetle adults fed aphids with facultative symbionts were significantly heavier than those fed facultative symbiont-free aphids, though development time was not significantly different. Conclusions: Aphids reproduce clonally and are often found in large groups. Thus, aphid symbionts, by reducing the fitness of the aphid predator H. convergens, may indirectly defend their hosts' clonal descendants against predation. These findings highlight the often far-reaching effects that symbionts can have in ecological systems.

Description: Background: The timing of migration substantially influences individual fitness. To match peak requirements with peak resource availability, we hypothesized that individual migrants schedule spring migration in close relation to seasonal changes in environmental conditions along the route and particularly, at the breeding destination.To test this hypothesis, we investigated the timing of spring migration in male common nightingales Luscinia megarhynchos, a small Palearctic-African long-distance migrant, by linking spring migration timing to the phenology of local environmental conditions at wintering, migratory stopover and breeding sites. In particular, we related individual migration decisions (i.e. departure and arrival) of nine males to site-specific vegetation phenology (based on remotely sensed vegetation index) and a proxy of food availability (based on insects' thermal requirements). Results: We found weak relation of departures from wintering and no relation of stopover timing with local phenology. However, our results showed that individuals, which departed early from their non-breeding sites and arrived early at the breeding site closely matched spring green-up there. Early arrival at the breeding site meant also a close match with peak food availability for adults and in a time-lagged manner, for offspring. Conclusion: Our findings suggest that male nightingale used cues other than local phenology for their departure decisions from wintering grounds and that there is some evidence for equalizing late departures during the course of migration.

Description: Background: Bromelia pinguin (Bromeliaceae) is a terrestrial bromeliad commonly found under forest stands throughout the Neotropics that has been shown to have antifungal activity in vitro. We have hypothesized that this bromeliad would also have an effect on the fungal populations in nearby soil by decreasing fungaldiversity and negatively impacting C and N cycle-related activities. A previous study in the lowland forest of Costa Rica showed the soil beneath these bromeliads had decreased fungal ITS DNA and differences in C and N levels compared to adjacent primary forest soils. Results: In this follow-up study, we found that the bromeliad soils had lower rates of C and N biomass development and lower phenol oxidase activity (suggesting less decreased fungal decomposition activity). The results of T-RFLP and cloning-based taxonomic analyses showed the community level diversity and abundance of fungal ITS DNA was less in bromeliad soils. Sequence analysis of fungal ITS DNA clones showed marked differences in fungal community structure between habitats of Basidiomycota (Tremellales, Agricales, Thelephorales), Ascomycota (Helotiales), and Zycomycota populations. Conclusions: The data show there to be differences in the soil nutrient dynamics and fungal community structure and activity associated with these bromeliads, as compared to the adjacent primary forest. This suggests the possibility that the anti-fungal activity of the bromeliad extends into the soil. The bromeliad-dense regions of these primary forest habitats provide a unique natural micro-habitat within the forests and the opportunity to better identify the role of fungal communities in the C and N cycles in tropical soils.

Description: Background: Windthrow, the uprooting of trees during storms associated with strong winds, is a well-established cause of mortality in temperate regions of the world, often with large ecological consequences. However, this phenomenon has received little attention within arid regions and is not well documented in southern Africa. Slow rates of post-disturbance recovery and projected increases in extreme weather events in arid areas mean that windthrow could be more common and have bigger impacts on these ecosystems in the future. This is of concern due to slow rates of post-disturbance recovery in arid systems and projected increases in extreme weather events in these areas. This study investigated the spatial pattern, magnitude and likely causes of windthrown mortality in relation to other forms of mortality in Aloe dichotoma, an iconic arid-adapted arborescent succulent and southern Africa climate change indicator species. Results: We found that windthrown mortality was greatest within the equatorward summer rainfall zone (SRZ) of its distribution (mean = 31%, n = 11), and was derived almost exclusively from the larger adult age class. A logistic modelling exercise indicated that windthrown mortality was strongly associated with greater amounts of warm season (summer) rainfall in the SRZ, higher wind speeds, and leptosols. A statistically significant interaction term between higher summer rainfall and wind speeds further increased the odds of being windthrown. While these results would benefit from improvements in the resolution of wind and substrate data, they do support the hypothesised mechanism for windthrow in A. dichotoma. This involves powerful storm gusts associated with either the current or subsequent rainfall event, heavy convective rainfall, and an associated increase in soil malleability. Shallow rooting depths in gravel-rich soils and an inflexible, top-heavy canopy structure make individuals especially prone to windthrown mortality during storms. Conclusions: Results highlight the importance of this previously unrecognised form of mortality in A. dichotoma, especially since it seems to disproportionately affect reproductively mature adult individuals in an infrequently recruiting species. Smaller, more geographically isolated and adult dominated populations in the summer rainfall zone are likely to be more vulnerable to localised extinction due to windthrow events.

Description: Background: Species coexistence in mosquito assemblages may depend on mechanisms related to interspecific resource partitioning occurring at multiple scales. In the present work we investigated co-occurrence or spatial segregation in mosquito assemblages sharing resources at micro-habitat, habitat and landscape scales. Environmental characteristics, mosquito fauna as adults and larvae were assessed along vegetation gradient in a natural landscape of tropical rainforest. Huisman-Olff-Fresco (HOF) and Generalized Additive (GAM) models were employed to explore relationships between abundances of potential competitors in mosquito assemblages and vegetation gradient (e.g., scrublands, mixed arboreal vegetation and dense ombrophilous forest). We tested hypotheses concerning mosquito species co-occurrence or spatial segregation employing binomial logistic regression models. Results: Co-occurrences and spatial segregation of mosquito species showed evidences of three scales of coexistence mechanisms: 1) micro-habitat ? scale 1: different behaviors in response to food availability in specific vertical strata within larval container; 2) habitat ? scale 2: specialized strategies related to heterogeneity of resource availability among larval containers and 3) landscape ? scale 3: asymmetrical competition dependent upon the context of abiotic and biotic variables. Conclusion: Results of the present work suggest that coexistence mechanisms can concomitantly work at multiple scales.

Description: Background: Most vertebrates experience coinfections, and many pathogen-pathogen interactions occur indirectly through the host immune system. These interactions are particularly strong in mixed micro-macroparasite infections because of immunomodulatory effects of helminth parasites. While these trade-offs have been examined extensively in laboratory animals, few studies have examined them in natural systems. Additionally, many wildlife pathogens fluctuate seasonally, at least partly due to seasonal host immune changes. We therefore examined seasonality of immune resource allocation, pathogen abundance and exposure, and interactions between infections and immunity in plains zebra (Equus quagga) in Etosha National Park (ENP), Namibia, a system with strongly seasonal patterns of gastrointestinal (GI) helminth infection intensity and concurrent anthrax outbreaks. Both pathogens are environmentally transmitted, and helminth seasonality is driven by environmental pressures on free living life stages. The reasons behind anthrax seasonality are currently not understood, though anthrax is less likely directly driven by environmental factors. Results: We measured a complex, interacting set of variables and found evidence that GI helminth infection intensities, eosinophil counts, IgE and IgGb antibody titers, and possibly IL-4 cytokine signaling were increased in wetter seasons, and that ectoparasite infestations and possibly IFN-? cytokine signaling were increased in drier seasons. Monocyte counts and anti-anthrax antibody titers were negatively associated with wet season eosinophilia, and monocytes were negatively correlated with IgGb and IgE titers. Taken together, this supports the hypothesis that ENP wet seasons are characterized by immune resource allocation toward Th-2 type responses, while Th1-type immunity may prevail in drier seasons, and that hosts may experience Th1-Th2 trade-offs. We found evidence that this Th2-type resource allocation is likely driven by GI parasite infections, and that these trade-offs may render hosts less capable of concurrently mounting effective Th1-type immune responses against anthrax. Conclusions: This study is one of the first to examine laboratory-demonstrated Th1-Th2 trade-offs in a natural system. It provides evidence that seasonally bound pathogens may affect, through immunology, transmission dynamics of pathogens that might otherwise not be seasonally distributed. It suggests that, by manipulating the internal host ecosystem, GI parasites may influence the external ecosystem by affecting the dynamics of another environmentally transmitted pathogen.

Description: Background: Nearly 40 years ago, Freeland and Janzen predicted that liver biotransformation enzymes dictated diet selection by herbivores. Despite decades of research on model species and humans, little is known about the biotransformation mechanisms used by mammalian herbivores to metabolize plant secondary compounds (PSCs). We investigated the independent evolution of PSC biotransformation mechanisms by capitalizing on a dramatic diet change event--the dietary inclusion of creosote bush (Larrea tridentata)--that occurred in the recent evolutionary history of two species of woodrats (Neotoma lepida and N. bryanti). Results: By comparing gene expression profiles of two populations of woodrats with evolutionary experience to creosote and one population naive to creosote, we identified genes either induced by a diet containing creosote PSCs or constitutively higher in populations with evolutionary experience of creosote. Although only one detoxification gene (an aldo-keto reductase) was induced by both experienced populations, these populations converged upon functionally equivalent strategies to biotransform the PSCs of creosote bush by constitutively expressing aldehyde and alcohol dehydrogenases, Cytochromes P450s, methyltransferases, glutathione S-transferases and sulfotransferases. The response of the naive woodrat population to creosote bush was indicative of extreme physiological stress. Conclusions: The hepatic detoxification system of mammals is notoriously complex, with hundreds of known biotransformation enzymes. The comparison herein of woodrat taxa that differ in evolutionary and ecological experience with toxins in creosote bush reveals convergence in the overall strategies used by independent species after a historical shift in diet. In addition, remarkably few genes seemed to be important in this dietary shift. The research lays the requisite groundwork for future studies of specific biotransformation pathways used by woodrats to metabolize the toxins in creosote and the evolution of diet switching in woodrats. On a larger level, this work advances our understanding of the mechanisms used by mammalian herbivores to process toxic diets and illustrates the importance of the selective relationship of PSCs in shaping herbivore diversity.

Description: BMC Ecology showcases the winning entries from its second Ecology Image Competition. More than 300 individual images were submitted from an international array of research scientists, depicting life on every continent on earth. The journal’s Editorial Board and guest judge Caspar Henderson outline why their winning selections demonstrated high levels of technical skill and aesthetic sense in depicting the science of ecology, and we also highlight a small selection of highly commended images that we simply couldn’t let you miss out on.

Description: Background: Ecologists recognize that plants capture nitrogen in many chemical forms that include amino acids. Access to multiple nitrogen types in plant communities has been argued to enhance plant performance, access to nitrogen and alter ecological interactions in ways that may promote species coexistence. However, data supporting these arguments have been limited. While it is known that plants uptake amino acids from soil, long term studies that link amino acid uptake to measures of plant performance and potential reproductive effort are not typically performed. Here, a series of experiments that link uptake of nitrate, glutamine or asparagine with lifetime reproductive effort in Arabidopsis thaliana are reported. Nitrogen was offered either singly or in mixture and at a variety of combinations. Traits related to reproductive output were measured, as was the preference for each type of nitrogen. Results: When plants were supplied with a single nitrogen type at concentrations from 0.1-0.9 mM, the ranking of nitrogen types was nitrate 〉 glutamine 〉 asparagine in terms of the relative performance of plants. When plants were supplied with two types of nitrogen in mixture at ratios between 0.1:0.9-0.9:0.1 mM, again plants performed best when nitrate was present, and poorly when amino acids were mixed. Additionally, stable isotopes revealed that plants preferentially captured nitrogen types matching the hierarchy of nitrate 〉 glutamine 〉 asparagine. Comparing between the two experiments revealed that mixed nitrogen nutrition was a net cost to the plants. Conclusions: Plant performance on mixed nitrogen was less than half the performance on equal amounts of any single nitrogen type. We asked: why did A. thaliana capture amino acids when doing so resulted in a net cost? We argue that available data cannot yet answer this question, but hypothesize that access to lower quality forms of nitrogen may become important when plants compete.

Description: Background: A multi-faceted approach was used to investigate the wintertime ecophysiology and behavioral patterns of the raccoon dog, Nyctereutes procyonoides, a suitable model for winter sleep studies. By utilizing GPS tracking, activity sensors, body temperature (Tb) recordings, change-point analysis (CPA), home range, habitat and dietary analyses, as well as fatty acid signatures (FAS), the impact of the species on wintertime food webs was assessed. The timing of passive bouts was determined with multiple methods and compared to Tb data analyzed by CPA. Results: Raccoon dogs displayed wintertime mobility, and the home range sizes determined by GPS were similar or larger than previous estimates by radio tracking. The preferred habitats were gardens, shores, deciduous forests, and sparsely forested areas. Fields had close to neutral preference; roads and railroads were utilized as travel routes. Raccoon dogs participated actively in the food web and gained benefit from human activity. Mammals, plants, birds, and discarded fish comprised the most important dietary classes, and the consumption of fish could be detected in FAS. Ambient temperature was an important external factor influencing Tb and activity. The timing of passive periods approximated by behavioral data and by CPA shared 91% similarity. Conclusions: Passive periods can be determined with CPA from Tb recordings without the previously used time-consuming and expensive methods. It would be possible to recruit more animals by using the simple methods of data loggers and ear tags. Hunting could be used as a tool to return the ear-tagged individuals allowing the economical extension of follow-up studies. The Tb and CPA methods could be applied to other northern carnivores.

Description: The recent declines in managed honey bee populations are of scientific, ecological and economic concern, and are partially attributed to honey bee parasites and related disease. McDonnell et al. investigate behavioral, chemical and neurogenomic effects of parasitization by the ectoparasite Varroa destructor and the endoparasite Nosema ceranae. The study reveals important links between underlying mechanisms of immunity and parasitization in social insects by demonstrating that chemical signals and neurogenomic states are significantly different between parasitized and non-parasitized honey bees, and that neurogenomic states are partially conserved between bees infected with distinct parasites. However the study does not reveal whether differences measured are primarily the result of adaptive host responses or of manipulation of the honey bee host by the parasites and/or confounding viral loads of parasitized individuals. Questions answered and raised by McDonnell et al. will lead to an improved understanding of honey bee health and, more generally, host-parasite interactions.

Description: Background: Exclusion from a social group is an effective way to avoid parasite transmission. This type of social removal has also been proposed as a form of collective defense, or social immunity, in eusocial insect groups. If parasitic modification of host behavior is widespread in social insects, the underlying physiological and neuronal mechanisms remain to be investigated. We studied this phenomenon in honey bees parasitized by the mite Varroa destructor or microsporidia Nosema ceranae, which make bees leave the hive precociously. We characterized the chemical, behavioral and neurogenomic changes in parasitized bees, and compared the effects of both parasites. Results: Analysis of cuticular hydrocarbon (CHC) profiles by gas chromatography coupled with mass spectrophotometry (GC-MS) showed changes in honey bees parasitized by either Nosema ceranae or Varroa destructor after 5 days of infestation. Levels of 10-HDA, an antiseptic important for social immunity, did not change in response to parasitism. Behavioral analysis of N. ceranae- or V. destructor- parasitized bees revealed no significant differences in their behavioral acts or social interactions with nestmates. Digital gene expression (DGE) analysis of parasitized honey bee brains demonstrated that, despite the difference in developmental stage at which the bee is parasitized, Nosema and Varroa-infested bees shared more gene changes with each other than with honey bee brain expression gene sets for forager or nurse castes. Conclusions: Parasitism by Nosema or Varroa induces changes to both the CHC profiles on the surface of the bee and transcriptomic profiles in the brain, but within the social context of the hive, does not result in observable effects on her behavior or behavior towards her. While parasitized bees are reported to leave the hive as foragers, their brain transcription profiles suggest that their behavior is not driven by the same molecular pathways that induce foraging behavior.

Description: Background: The life history strategy of a species can influence how populations of that species respond to environmental variation. In this study, we used a matrix modeling approach to examine how life history differences among sympatric rock and white-tailed ptarmigan affect the influence of demographic rates on population growth (lambda) and the potential response to a changing climate. Rock ptarmigan have a slower life history strategy than white-tailed ptarmigan in the study region with lower annual reproductive effort but higher adult survival. Results: Based on data from a 5-year field study, deterministic estimates of lambda indicated that populations were stable for rock ptarmigan (lambda = 1.01), but declining for white-tailed ptarmigan (lambda = 0.96). The demographic rates with the highest elasticity for rock ptarmigan were the survival of after-second year females, followed by juvenile survival and success of the first nest. For white-tailed ptarmigan, juvenile survival had the highest elasticity followed by success of the first nest and survival of second-year females. Incorporating stochasticity into the demographic rates led to a 2 and 4% drop in lambda for rock and white-tailed ptarmigan respectively. Using data from the first three years we also found that population growth rates of both species were depressed following an increased frequency of severe years, but less so for rock ptarmigan which showed greater resilience under these conditions. Conclusions: Our results provide evidence that populations of closely related species can vary in their response to environmental change as a consequence of life history differences. Rock ptarmigan, with a slower life history, are more responsive to demographic rates that influence survival and older life stages but this response is tempered by the extent of variability in each of the rates. Thus, predictions need to consider both aspects in modeling population response to a varying climate. Juvenile survival was a highly influential rate for both species, but the period from independence to first breeding is a poorly understood stage for many bird species. Additional study on juvenile survival, the influence of density dependence and the effects of predators as the mechanism driving survival-reproduction tradeoffs are all areas requiring further study.

Description: Background: Intraguild predation (IGP) is common in nature, but its ecological role is still illusive. A number of studies have investigated a three species IGP module that consists of an intraguild predator, intraguild prey, and resource species in which the intraguild predator and the intraguild prey exploitatively compete for the resource while the intraguild predator also consumes the intraguild prey. A common prediction of models of the IGP module is that the coexistence of the species is difficult, which is considered inconsistent to the ubiquity of IGP in nature. This study revisits the IGP module and provides an alternative coexistence mechanism by focusing on a commonly used analysis method (i.e., invasion analysis) in light of individual variation in adaptive behavior. Results: Invasion analysis underestimates the possibility of coexistence regardless of the presence or absence of adaptive behavior. Coexistence is possible even when invasion analysis predicts otherwise. The underestimation by invasion analysis is pronounced when the intraguild predator forages adaptively, which is even further pronounced when the expression of foraging behavior is variable among intraguild predators. Conclusions: The possibility of coexistence in the IGP module is greater than previously thought, which may have been partly due to how models were analyzed. Inconsistent conclusions may result from the same model depending on how the model is analyzed. Individual variation in adaptive behavior can be an important factor promoting the coexistence of species in IGP modules.

Description: Background: Deployment of cultivars with different resistance in mixtures is one means to manage plant diseases and prolong the life of resistance genes. One major concern in adopting mixtures is the development of 'super-races' that can overcome many resistance genes present in the mixture. A stochastic simulation model was developed to study the dynamics of virulence alleles in two-cultivar mixtures of perennial crops, focusing on the effects of cost of virulence and pathogen reproduction mechanism. The simulated mechanism of virulence has characteristics of both major and minor genes. Results: Random genetic drift due to repeated population crashes during the overwintering phase led to fixation of a single fungal genotype (in terms of its virulence), often within 100 seasons. Overall, cost of virulence is most important in determining the virulence dynamics under the present model formulation. With cost of virulence incorporated, nearly all simulation runs ended up with a single fungal genotype that can infect only one of the two cultivars. In absence of cost of virulence, most of the simulation runs ended up with fungal genotypes that can infect both host cultivars but in many cases do not contain the maximum possible number of virulence alleles due to random drift. A minimum of 20% sexual reproduction between strains from different cultivars is necessary to ensure that the final fixed strains are able to infect both cultivars. Although the number of virulence alleles in the final genotype and the time to fixation are affected by simulation factors, most of the variability was among replicate simulation runs (i.e. stochastic in nature). The time to fixation is generally long relative to cropping cycles. Conclusions: A single fungal genotype will dominate a population due to the bottleneck in overwintering with cost of virulence primarily determining whether the dominant genotype can infect both cultivars. However, the dominate genotype is unlikely to accumulate all the virulence alleles due to genetic drift. The risk of emergence and spread of super-races is insufficiently great to prevent the use of cultivar mixtures of perennial crops as a means to reduce disease development provided that host resistance structure in mixtures is altered every cropping cycle.

Description: Background: Avian malaria (Plasmodium sp.) is globally widespread, but considerable variation exists in infection (presence/absence) patterns at small spatial scales. This variation can be driven by variation in ecology, demography, and phenotypic characters, in particular those that influence the host's resistance. Generation of reactive oxygen species (ROS) is one of the host's initial immune responses to combat parasitic invasion. However, long-term ROS exposure can harm the host and the redox response therefore needs to be adjusted according to infection stage and host phenotype. Here we use experimental and correlational approaches to assess the relative importance of host density, habitat composition, individual level variation and redox physiology for Plasmodium infection in a wild population of great tits, Parus major. Results: We found that 36% of the great tit population was infected with Plasmodium (22% P. relictum and 15% P. circumflexum prevalence) and that patterns of infection were Plasmodium species-specific. First, the infection of P. circumflexum was significantly higher in areas with experimental increased host density, whereas variation in P. relictum infection was mainly attributed to age, sex and reproduction. Second, great tit antioxidant responses -- total and oxidizied glutathione - showed age- , sex- and Plasmodium species-specific patterns between infected and uninfected individuals, but reactive oxygen metabolites (ROM) showed only a weak explanatory power for patterns of P. relictum infection. Instead ROM significantly increased with Plasmodium parasitaemia. Conclusions: These results identify some key factors that influence Plasmodium infection in wild birds, and provide a potential explanation for the underlying physiological basis of recently documented negative effects of chronic avian malaria on survival and reproductive success.

Description: Background: Sampling expeditions to Churchill in the Canadian subarctic were completed with the aim of compiling a molecular-assisted survey of the macroalgal flora (seaweeds) for comparison to published accounts for this area, which are based on morphological identifications. Further, because the Churchill region was covered by ice until recently (~10,000 before present), the current algal flora has had to migrate from adjacent waters into that region. We used our DNA barcode data to predict the relative contribution of the North Atlantic and North Pacific floras (Likely Source Region) in repopulating the Churchill region following the most recent glacial retreat. Results: We processed 422 collections representing ~50 morpho-species, which is the approximate number reported for this region, and generated DNA barcode data for 346 of these. In contrast to the morpho-species count, we recovered 57 genetic groups indicating overlooked species (this despite failing to generate barcode data for six of the ~50 morpho-species). However, we additionally uncovered numerous inconsistencies between the species that are currently listed in the Churchill flora (again as a result of overlooked species diversity, but combined with taxonomic confusion) and those identified following our molecular analyses including eight new records and another 17 genetic complexes in need of further study. Based on a comparison of DNA barcode data from the Churchill flora to collections from the contiguous Atlantic and Pacific floras we estimate that minimally 21% (possibly as much as 44%) of the Churchill flora was established by migration from the Pacific region with the balance of species arriving from the Atlantic (predominantly North American populations) following the last glacial retreat. Conclusions: Owing to difficulties associated with the morphological identification of macroalgae, our results indicate that current comprehension of the Canadian Arctic flora is weak. We consider that morphology-based field-identifications are ill-advised in carrying out floristic and ecological surveys for macroalgae and that much of the current data, at least for the Canadian Arctic, should be used with caution. Our efforts to use DNA barcode data to identify the most Likely Source Regions for macroalgal species currently found in Churchill suggests that migration from both the Atlantic and the Pacific have played important roles in establishing the Canadian Arctic flora. This result has significance for understanding both the current and future biodiversity and biogeography of macroalgae in these waters.

Description: Background: Biodiversity surveys have long depended on traditional methods of taxonomy to inform sampling protocols and to determine when a representative sample of a given species pool of interest has been obtained. Questions remain as to how to design appropriate sampling efforts to accurately estimate total biodiversity. Here we consider the biodiversity of freshwater ostracods (crustacean class Ostracoda) from the region of Churchill, Manitoba, Canada. Through an analysis of observed species richness and complementarity, accumulation curves, and richness estimators, we conduct an a posteriori analysis of five bioblitz-style collection strategies that differed in terms of total duration, number of sites, protocol flexibility to heterogeneous habitats, sorting of specimens for analysis, and primary purpose of collection. We used DNA barcoding to group specimens into molecular operational taxonomic units for comparison. Results: Forty-eight provisional species were identified through genetic divergences, up from the 30 species previously known and documented in literature from the Churchill region. We found differential sampling efficiency among the five strategies, with liberal sorting of specimens for molecular analysis, protocol flexibility (and particularly a focus on covering diverse microhabitats), and a taxon-specific focus to collection having strong influences on garnering more accurate species richness estimates. Conclusions: Our findings have implications for the successful design of future biodiversity surveys and citizen-science collection projects, which are becoming increasingly popular and have been shown to produce reliable results for a variety of taxa despite relying on largely untrained collectors. We propose that efficiency of biodiversity surveys can be increased by non-experts deliberately selecting diverse microhabitats; by conducting two rounds of molecular analysis, with the numbers of samples processed during round two informed by the singleton prevalence during round one; and by having sub-teams (even if all non-experts) focus on select taxa. Our study also provides new insights into subarctic diversity of freshwater Ostracoda and contributes to the broader "Barcoding Biotas" campaign at Churchill. Finally, we comment on the associated implications and future research directions for community ecology analyses and biodiversity surveys through DNA barcoding, which we show here to be an efficient technique enabling rapid biodiversity quantification in understudied taxa.

Description: Background: The algal endosymbionts (genus Symbiodinium) associated with scleractinian corals (and other reef invertebrates) have received a lot of research attention in the past decade, particularly as certain host-symbiont associations appear more affected by increasing seawater temperatures than others. With the rapid accumulation of information on the diversity of Symbiodinium, it is becoming increasingly difficult to compare newly acquired Symbiodinium data with existing data to detect patterns of host-symbiont specificity on broader spatial scales. The lack of a general consensus on the classification of Symbiodinium species coupled with the variety of different markers used to identify the genus Symbiodinium (ITS1, ITS2, LSU D1/D2, chloroplast 23S rDNA and psbA minicircle) further complicate direct comparison.Description: The SymbioGBR database compiles all currently available Symbiodinium sequences and associated host information of data collected from the Great Barrier Reef into a single relational database that is accessible via a user-friendly, searchable web-based application (http://www.SymbioGBR.org). SymbioGBR allows users to query Symbiodinium types or sequences sourced from various genetic markers (e.g. ITS1, ITS2, LSU D1/D2 and chloroplast 23S) and invertebrate host species to explore their reported associations. In addition, as the database includes sequence information of multiple genetic markers, it allows cross-referencing between conventional (e.g. ITS2 region) and novel markers that exhibit low intragenomic variability (e.g. psbA region). Finally, the database is based on the collection details of individual specimens. Such host-symbiont associations can be assessed quantitatively and viewed in relation to their environmental and geographic context. Conclusions: The SymbioGBR database provides a comprehensive overview of Symbiodinium diversity and host-associations on the Great Barrier Reef. It provides a quick, user-friendly means to compare newly acquired data on Symbiodinium (e.g. raw sequences or characterized Symbiodinium types) with previous data on the diversity of invertebrate host-symbiont associations on the GBR. The inclusion of psbAncr sequence information allows for validation of widely used ITS1/ITS2 markers and their ability to accurately identify relevant sequences. Most importantly, centralization of sequence information from multiple genetic markers will aid the classification of Symbiodinium species diversity and allow researchers to easily compare patterns of host-Symbiodinium associations.

Description: Background: Animal colour patterns offer good model systems for studies of biodiversity and evolution of local adaptations. An increasingly popular approach to study the role of selection for camouflage for evolutionary trajectories of animal colour patterns is to present images of prey on paper or computer screens to human 'predators'. Yet, few attempts have been made to confirm that rates of detection by humans can predict patterns of selection and evolutionary modifications of prey colour patterns in nature. In this study, we first analyzed encounters between human 'predators' and images of natural black, grey and striped colour morphs of the polymorphic Tetrix subulata pygmy grasshoppers presented on background images of unburnt, intermediate or completely burnt natural habitats. Next, we compared detection rates with estimates of capture probabilities and survival of free-ranging grasshoppers, and with estimates of relative morph frequencies in natural populations. Results: The proportion of grasshoppers that were detected and time to detection depended on both the colour pattern of the prey and on the type of visual background. Grasshoppers were detected more often and faster on unburnt backgrounds than on 50% and 100% burnt backgrounds. Striped prey were detected less often than grey or black prey on unburnt backgrounds; grey prey were detected more often than black or striped prey on 50% burnt backgrounds; and black prey were detected less often than grey prey on 100% burnt backgrounds. Rates of detection mirrored previously reported rates of capture by humans of free-ranging grasshoppers, as well as morph specific survival in the wild. Rates of detection were also correlated with frequencies of striped, black and grey morphs in samples of T. subulata from natural populations that occupied the three habitat types used for the detection experiment. Conclusions: Our findings demonstrate that crypsis is background-dependent, and implicate visual predation as an important driver of evolutionary modifications of colour polymorphism in pygmy grasshoppers. Our study provides the clearest evidence to date that using humans as 'predators' in detection experiments may provide reliable information on the protective values of prey colour patterns and of natural selection and microevolution of camouflage in the wild.

Description: Background: Rising temperatures and other environmental factors influenced by global climate change can cause increased physiological stress for many species and lead to range shifts or regional population extinctions. To advance the understanding of species' response to change and establish links between individual and ecosystem adaptations, physiological reactions have to be compared between populations living in different environments. Although changes in expression of stress genes are relatively easy to quantify, methods for reliable comparison of the data remain a contentious issue. Using normalization algorithms and further methodological considerations, we compare cellular stress response gene expression levels measured by RT-qPCR after air exposure experiments among different subpopulations of three species of the intertidal limpet Nacella. Results: Reference gene assessment algorithms reveal that stable reference genes can differ among investigated populations and / or treatment groups. Normalized expression values point to differential defense strategies to air exposure in the investigated populations, which either employ a pronounced cellular stress response in the inducible Hsp70 forms, or exhibit a comparatively high constitutive expression of Hsps (heat shock proteins) while showing only little response in terms of Hsp induction. Conclusions: This study serves as a case study to explore the methodological prerequisites of physiological stress response comparisons among ecologically and phylogenetically different organisms. To improve the reliability of gene expression data and compare the stress responses of subpopulations under potential genetic divergence, reference gene stability algorithms are valuable and necessary tools. As the Hsp70 isoforms have been shown to play different roles in the acute stress responses and increased constitutive defenses of populations in their different habitats, these comparative studies can yield insight into physiological strategies of adaptation to environmental stress and provide hints for the prudent use of the cellular stress response as a biomarker to study environmental stress and stress adaptation of populations under changing environmental conditions.

Description: Background: Conservation of biological diversity and economical utilization of natural resources form an almost inevitable confrontation between the two. In practice, however, a balance between the two ought to be found, and in managed boreal forests, preservation of woodland key habitats is increasingly used strategy to safeguard biological diversity. According to the Finnish Forests Act, certain Forest Act habitat (FAH) types must be safeguarded, provided they are clearly distinguishable from their surroundings. Furthermore, once the habitat has been identified as a FAH, its special characteristics must not be altered. Both of these aspects contain ambiguities that potentially undermine the practical application of the Act. We designed a replicated sampling study to address these ambiguities at the most common FAH type, riparian habitat of small boreal streams. As response variables we used vascular plants and mosses. We asked i) how wide is the FAH around small streams that is distinguishable from its surrounding and ii) how wide buffer strip around the FAH is sufficient for long term to preserve the natural species community composition of the FAH. Results: We found that an average three meters wide strip around the stream constitutes the distinguishable FAH and that a minimum of 45 meters wide buffers on both sides of the stream are needed for the species community composition to remain unaltered. Conclusions: We conclude that 45 meters wide buffers appear sufficient to safeguard vascular plant and moss species communities within the FAH, prevent local populations from extinctions and thus pre-empt extinction debt that would be realised with more narrow buffers. While 45 meters may seem intolerable from the commercial forestry point of view, anything less than that may be intolerable from the point of view of conservation, and thus against the idea of sustainable use of natural resources.

Description: Background: Arctic ecosystems, especially those near transition zones, are expected to be strongly impacted by climate change. Because it is positioned on the ecotone between tundra and boreal forest, the Churchill area is a strategic locality for the analysis of shifts in faunal composition. This fact has motivated the effort to develop a comprehensive biodiversity inventory for the Churchill region by coupling DNA barcoding with morphological studies. The present study represents one element of this effort; it focuses on analysis of the spider fauna at Churchill. Results: 198 species were detected among 2704 spiders analyzed, tripling the count for the Churchill region. Estimates of overall diversity suggest that another 10--20 species await detection. Most species displayed little intraspecific sequence variation (maximum