Biogeography of avian blood parasites (Leucocytozoon spp.) in two resident hosts across Europe: phylogeographic structuring or the abundance–occupancy relationship?

Relationships between hosts and parasites represent complex co-evolving systems that can vary both temporally and spatially. This variation may result in different phylogeographic outcomes, ranging from highly geographically structured parasite populations comprised of specialist lineages that are locally abundant but have restricted global occupancy to geographically unstructured parasite populations consisting of widespread parasites. Here, we present results from a large biogeographic study of the Leucocytozoon blood parasites of two nonmigrant bird species, conducted at nine sites across Europe. The aim was to determine whether the parasite lineages of the two hosts were phylogeographically structured across Europe. Employing molecular methods, we found a large diversity of parasites, and although overall prevalence varied greatly, the parasites were not genetically structured. Several measures of local parasite abundance were associated with the number of sites that the lineage occurred in, which is consistent with the macroecological phenomenon of the abundance-occupancy relationship. Taken together, our results show that parasite dispersal is somewhat uncoupled to that of the host in this system: we suggest that broad host and/or vector preference may play an important role in determining the distribution of these parasites and in affecting host-parasite coevolution in this system.     

Does the disturbance hypothesis explain the biomass increase in basin‐wide Amazon forest plot data?

Positive aboveground biomass trends have been reported from old-growth forests across the Amazon basin and hypothesized to reflect a large-scale response to exterior forcing. The result could, however, be an artefact due to a sampling bias induced by the nature of forest growth dynamics. Here, we characterize statistically the disturbance process in Amazon old-growth forests as recorded in 135 forest plots of the RAINFOR network up to 2006, and other independent research programmes, and explore the consequences of sampling artefacts using a data-based stochastic simulator. Over the observed range of annual aboveground biomass losses, standard statistical tests show that the distribution of biomass losses through mortality follow an exponential or near-identical Weibull probability distribution and not a power law as assumed by others. The simulator was parameterized using both an exponential disturbance probability distribution as well as a mixed exponential–power law distribution to account for potential large-scale blowdown events. In both cases, sampling biases turn out to be too small to explain the gains detected by the extended RAINFOR plot network. This result lends further support to the notion that currently observed biomass gains for intact forests across the Amazon are actually occurring over large scales at the current time, presumably as a response to climate change.     

Do large‐seeded herbs have a small range size? The seed mass–distribution range trade‐off hypothesis

We aimed to introduce and test the “seed mass–distribution range trade‐off” hypothesis, that is, that range size is negatively related to seed mass due to the generally better dispersal ability of smaller seeds. Studying the effects of environmental factors on the seed mass and range size of species, we also aimed to identify habitats where species may be at risk and need extra conservation effort to avoid local extinctions. We collected data for seed mass, global range size, and indicators for environmental factors of the habitat for 1,600 species of the Pannonian Ecoregion (Central Europe) from the literature. We tested the relationship between species’ seed mass, range size, and indicator values for soil moisture, light intensity, and nutrient supply. We found that seed mass is negatively correlated with range size; thus, a seed mass–distribution range trade‐off was validated based on the studied large species pool. We found increasing seed mass with decreasing light intensity and increasing nutrient availability, but decreasing seed mass with increasing soil moisture. Range size increased with increasing soil moisture and nutrient supply, but decreased with increasing light intensity. Our results supported the hypothesis that there is a trade‐off between seed mass and distribution range. We found that species of habitats characterized by low soil moisture and nutrient values but high light intensity values have small range size. This emphasizes that species of dry, infertile habitats, such as dry grasslands, could be more vulnerable to habitat fragmentation or degradation than species of wet and fertile habitats. The remarkably high number of species and the use of global distribution range in our study support our understanding of global biogeographic processes and patterns that are essential in defining conservation priorities.     

Positive relationship between regional distribution and local abundance in stream insects: a consequence of niche breadth or niche position?

A positive relationship between regional distribution and local abundance of species is almost ubiquitous macroecological pattern, yet the mechanisms behind this pattern remain poorly understood. I tested for the relationship between regional distribution and local abundance of stream insect species in a boreal drainage system, with a specific aim to examine if this relationship follows the mechanistic basis of either the niche-based (niche breadth and niche position) or metapopulation models. There was a positive relationship between regional distribution and local abundance of stream insects, and there also were significant relationships between distribution/abundance and niche breadth or niche position. These results thus suggest that widely distributed species tend to be, on average, locally more abundant, have wider niches and lower marginality of niche position with regard to environmental factors than species that have more restricted distributions. However, although significant, there was much unexplained variability around these relationships, suggesting that other mechanisms (e.g. metapopulation dynamics) besides differences in species' niches are likely to affect the distribution and abundance of stream insects, at least within a drainage system. The results thus showed that 1) although niche position was more consistently related to the positive distribution-abundance relationship, ecologists should not abandon niche breadth as a potential mechanism behind this relationship, and 2) that several mechanisms are likely to act in concert in determining the relationship between distribution and abundance of species.     

Ditch the niche – is the niche a useful concept in ecology or species distribution modelling?

In this first of three papers we examine the use of niche concepts in ecology and especially in species distribution modelling (SDM). This paper deliberately focuses on the lack of clarity found in the term ‘niche’. Because its meanings are so diverse, the term niche tends to create confusion and requires constant qualification. The literature houses many idiosyncratic ideas of what the niche is, but few examples where niche is more explanatory than the terminologies of population and community ecology or the statistical methods used to implement SDM analyses. In many cases the original (and inspirational) concepts are not directly applicable to our modern applications (e.g. set theory). There are some conceptual limitations found in individual definitions of niche (e.g. the fundamental niche concept), so it is perhaps understandable why more neutral terminology is becoming popular in SDM. An examination of the literature reveals a wide range of uncritical use of niche terminology. Our findings in this paper do not necessarily support the position of niche as a universally useful concept.     

Combined effects of shade and drought on tulip poplar seedlings: trade‐off in tolerance or facilitation?

It has been hypothesised that many natural plant distribution patterns can be explained by a trade-off between shade and drought tolerance which would make plants more sensitive to shade under dry conditions. On the other hand, shading by nurse plants is often observed to enhance growth and survival of smaller plants in dry areas. This paper describes an experiment designed to address the interactive effects of drought and shade on growth and physiology of tulip poplar seedlings ( Liriodendron tulipifera ) grown in shade houses under different combinations of irradiance (1%, 5%, 12%, 17%, and 27% of ambient PAR) and soil water content (5–9%, 11–15%, and>20%). The results show no evidence that higher PAR levels compensate for the negative effects of drought on photosynthesis and growth. Rather, rates of estimated daily shoot carbon gain decreased with PAR under dry conditions. Daily xylem sapflow patterns indicated that this was associated with a strong reduction in stomatal conductance in plants growing in dry soil and high PAR conditions. Whole-shoot light compensation points were higher for plants raised under higher PAR conditions, but were not significantly influenced by the water treatments. Shoot dark respiration rates decreased under drier conditions. These results do not support the hypothesis of a trade-off between shade and drought tolerance. Instead, they indicate mechanisms that can lead to positive effects of shading by neighbouring plants under dry conditions. Indeed, such facilitation is thought to be important in many plant communities.     

Morphology dependent foraging efficiency in perch: a trade‐off for ecological specialization?

Trade-offs in foraging efficiency leading to divergent natural selection between and within populations exploiting different resources are thought to be a primary cause of trophic polymorphism. In this study we focused on the trade-offs in foraging efficiency and growth in a polymorphic perch population. Specifically, we related habitat-specific growth and diet of perch to perch morphology. In a subsequent laboratory study we experimentally tested the trade-off by testing the efficiency of perch with different morphology feeding on pelagic ( Daphnia sp., Chaoborus sp.) and littoral (mayfly larvae) food resources. The feeding performance was tested in different physical environments to see if we could predict growth patterns in the field based on foraging rate and behavior of perch.
In the field study, we found that the perch from the littoral and the pelagic zones differed in both morphology and diet. Within the littoral zone the deeper-bodied individuals grew faster compared to the more streamlined individuals, whereas the opposite pattern was found in the pelagic zone. In the aquarium experiments, perch from the littoral zone had higher capture rates on the pelagic prey types in vegetation trials and on mayfly larvae in both open water and vegetation trials. The pelagic perch had higher capture rates on the pelagic prey types in open water trials. The littoral perch had lower search velocity than the pelagic perch in open water trials whereas the opposite pattern was found in vegetation trials. The attack velocity of the pelagic perch was also higher than that of the littoral perch independent of vegetation structure. Our results suggest that there is a functional trade-off between performance in alternate habitats and general body form in perch. Such trade-offs may promote divergent natural selection and could be the mechanism that give rise to and upholds the pattern in the field.     

Does scale‐dependent feedback explain spatial complexity in salt‐marsh ecosystems?

Complexity theory highlights scale-dependent feedback mechanisms as an explanation for regular spatial patterning in ecosystems. To what extent scale-dependent feedback clarifies spatial structure in more complex, non-regular systems remains unexplored so far. We report on a scale-dependent feedback process generating patchy landscapes at the interface of intertidal flats and salt marshes. Here, vegetation was characterized by Spartina anglica tussocks, surrounded by erosion gullies. To demonstrate the presence of a scale-dependent feedback, we determined if vegetation induced habitat modification resulted in local facilitation and large scale-inhibition of plant growth. Field surveys revealed that larger tussocks have deeper gullies, suggesting that gully erosion is caused by increased water flow around tussocks. This was confirmed by flume experiments, showing that feedback effects vary with current velocity and water depth. Transplantation of small Spartina units inside and just outside present tussocks revealed that the growth of Spartina transplants compared to transplant growth on bare sediment was higher within the raised Spartina tussocks, but lower in the gully just outside Spartina tussocks, providing clear evidence of scale-dependent feedback. Our results emphasize that scale-dependent feedback is a more general explanation for spatial complexity in ecosystems than previously considered.     

Does a trade‐off between current reproductive success and survival affect the honesty of male signalling in species with male parental care?

Recent theory predicted that male advertisement will reliably signal investment in paternal care in species where offspring survival requires paternal care and males allocate resources between advertisement and care. However, the predicted relationship between care and advertisement depended on the marginal gains from investment in current reproductive traits. Life history theory suggests that these fitness gains are also subject to a trade‐off between current and future reproduction. Here, we investigate whether male signalling remains a reliable indicator of parental care when males allocate resources between current advertisement, paternal care and survival to future reproduction. We find that advertisement is predicted to remain a reliable signal of male care but that advertisement may cease to reliably indicate male quality because low‐quality males are predicted to invest in current reproduction, whereas higher‐quality males are able to invest in both current reproduction and survival to future reproduction.     

Is there a trade‐off between horn growth and survival in adult female chamois?

Life‐history theory predicts trade‐offs in energy allocation between different life‐history traits when resources are limited, i.e. certain traits should be negatively correlated. However, individuals differ in their ability to acquire resources, which can lead to positive correlations between traits at the population level. Here, we investigated the consequences of the allocation in horn growth and body mass on survival in a bovid (Rupicapra rupicapra) with capture‐mark re‐sighting data on 161 females. In female ungulates, body mass often covaries positively with demographic performance and the few studies on horn size suggest that this trait could be a signal of individual quality. Thus, we expected to measure positive correlations between the allocation in these traits and female survival. However, body mass was not correlated to female survival and there was only a negative, though marginal, effect of horn growth. Hence, it seems that the allocation in growth is not an indicator of female quality. Future studies could investigate the importance of growth on female reproduction to evaluate its effect on lifetime reproductive success. Moreover, it is important to confirm in other populations our result that suggests a cost of the allocation in horn growth to better understand the presence of horns in female bovids. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 516–521.     

Digest: Does sexual conflict complicate a trade‐off between fecundity and survival?*

Why do some species exhibit apparently suboptimal combinations of life history traits? In a comparative study of lizard species, Reedy et al. test the idea that variation in the trade‐off between fecundity and survival can be explained by sexual conflict. Their results show that degree of sexual conflict alone cannot explain this variation, and they found a positive correlation between trade‐off optimality and the size of males relative to females. These findings suggest a more complicated picture of the fecundity‐survival trade‐off, possibly involving a third, unknown source of selective pressure.     

Costly carotenoids: a trade‐off between predation and infection risk?

Carotenoid reserves in copepods seem costly in terms of predation risk because they make individuals conspicuous. However, carotenoids also seem to play an important role in immune defence as free radical scavengers. To test whether predation risk influences carotenoid levels and whether changes in carotenoid levels are related to changes in immune defence, I examined individual changes in large carotenoid and other lipid droplets upon exposure to predation risk and subsequent exposure to parasites in the copepod Macrocyclops albidus. Copepods reduced carotenoid reserves upon exposure to predators, through which they potentially avoided the costs of being conspicuous under predation risk. Thus, the size of carotenoid reserves is a plastic trait. Such a decrease in carotenoid reserves may also have a negative impact on the copepods' immune system as individuals that decreased their reserves suffered higher parasite prevalence upon exposure to the cestode Schistocephalus solidus. These results suggest that carotenoid reserves may be individually optimized to trade-off each individual's unique costs (predation risk) and benefits (immune defence) of having these reserves.     

Does the seed size/number trade‐off model determine plant community structure? An assessment of the model mechanisms and their generality

This paper examines four key mechanisms of the seed size/number trade-off (SSNT) models to assess their relevance to a general understanding of plant community structure. Mechanism 1 is that large seeds have a greater probability of winning in competition against smaller seeds. I provide interspecific experimental evidence that there is a competitive hierarchy among seedlings based on seed size. Mechanism 2 is that a trade-off exists between the number and size of seeds produced for a given reproductive allocation. Negative correlations between seed size and number were found consistently across a range of species from a range of habitats, from published literature. Mechanism 3, that seedling-seedling competition is an important influence on species composition, was found to exist potentially in a range of environments, including annual-dominated, post-fire and gap-dynamic communities. However, there is little quantitative evidence available and this is likely to be a restrictive mechanism. Mechanism 4, that small seeds are superior colonists due to their greater number, was tested in a field experiment in a calcareous grassland community. No supporting evidence was found, suggesting that the SSNT is not an important determinant of structure in this community. Thus two of the four mechanisms can be considered to hold true generally, while the third mechanism may be valid in particular environments. The fourth mechanism did not apply in the community tested, but could be tested in a wider range of communities.     

Are seed germination and ecological breadth associated? Testing the regeneration niche hypothesis with bromeliads in a heterogeneous neotropical montane vegetation

The role of seed germination in contributing to species ecological breadth and geographic distribution is still a matter of debate. Here, we attempted to relate seed germination requirements with ecological breadth in 12 bromeliad species from heterogeneous montane vegetation in southeastern Brazil. Seeds were set to germinate under both light and dark conditions at a broad range of temperatures to determine the breadth of the germination niche. We ran a RLQ analysis based on the matrices of species occurrence, environmental parameters and germination traits and found a significant association between germination traits, and the characteristics of sites where adult plants occur. The variation of germination responses to environmental factors was not random with habitat-generalist plants having broader germination niches and habitat-specialist plants having narrower germination niches. The RLQ analysis showed that substrate moisture and light environment were the most important factors correlated with germination traits. Phylogenetic niche conservatism appears to play a role in the patterns found here, especially in the Tillandsioideae. There is an association between the regeneration niche of a species and its ecological range, and this also provides support for the idea that the regeneration niche may help assemble plant species into heterogeneous, species-rich communities.     

Does the temporal mismatch hypothesis match in boreal populations?

The temporal mismatch hypothesis suggests that fitness is related to the degree of temporal synchrony between the energetic needs of the offspring and their food supply. The hypothesis has been a basis in studying the influence of climate warming on nature. This study enhances the knowledge on prevalence of temporal mismatches and their consequences in boreal populations, and questions the role of the temporal mismatch hypothesis as the principal explanation for the evolution of timing of breeding. To test this, we examined if synchrony with caterpillar prey or timing of breeding per se better explains reproductive output in North European parid populations. We compared responses of temperate-origin species, the great tit (Parus major) and the blue tit (Cyanistes caeruleus), and a boreal species, the willow tit (Poecile montanus). We found that phenologies of caterpillars and great tits, but not of blue tits, have advanced during the past decades. Phenologies correlated with spring temperatures that may function as cues about the timing of the food peak for great and blue tits. The breeding of great and blue tits and their caterpillar food remained synchronous. Synchrony explained breeding success better than timing of breeding alone. However, the synchrony effect arose only in certain conditions, such as with high caterpillar abundances or high breeding densities. Breeding before good synchrony seems advantageous at high latitudes, especially in the willow tit. Thus, the temporal mismatch hypothesis appears insufficient in explaining the evolution of timing of breeding.     

Does attraction to conspecifics explain the patch‐size effect? An experimental test

Recent theory suggests that attraction to conspecifics during habitat selection can be one potential, yet untested, mechanism for animal sensitivity to habitat fragmentation. The least flycatcher Empidonax minimus , a highly territorial migratory bird, has previously been shown to be attracted to conspecifics and sensitive to patch size by avoiding small patches of riparian forest in Montana, USA. I used a large-scale field experiment in this region to test the conspecific attraction hypothesis for explaining sensitivity to patch size, and I supplemented this experiment by estimating whether vegetation structure, nest predation, or nest parasitism rates could better explain patterns of sensitivity to patch size. Vegetation structure did not vary consistently with patch size, based on a random sample of patches across 150  km of the Madison and Missouri Rivers, Montana. Nest predation and parasitism rates by brown-headed cowbirds Molothrus ater also did not vary with patch size during the experiment. However, when conspecific cues were simulated across a gradient of patch sizes, flycatchers settled in all patches – and their sensitivity to patch size vanished – providing strong support for the conspecific attraction hypothesis. These results provide the first experimental evidence that attraction to conspecifics can indeed help explain area sensitivity in nature and highlight how understanding the role of animal behavior in heterogeneous landscapes can aid in interpreting pressing conservation issues.     

Extending the stress‐gradient hypothesis – is competition among animals less common in harsh environments?

The role of positive interactions has become widely accepted as a mechanism shaping community dynamics. Most empirical evidence comes from plant communities and sessile marine organisms. However, evidence for the relative role of positive interactions in organizing terrestrial animal communities is more limited, and a general framework that includes positive interactions among animals is lacking. The ‘stress gradient hypothesis’ (SGH) developed by plant ecologists predicts that the balance between positive and negative interactions will vary along gradients of biotic and abiotic stress, with positive interactions being more important in stressful environments. Paralleling the SGH, stress gradients for terrestrial herbivores could be equated to inverse primary productivity gradients, so we would expect positive interactions to prevail in more stressful, low productivity environments. However, this contradicts the typical view of terrestrial animal ecology that low primary productivity systems will foster intense competition for resources among consumers. Here we use alpine herbivores as a case study to test one of the predictions of the SGH in animal communities, namely the prevalence of positive interactions in low productivity environments. We identify potential mechanisms of facilitation and review the limited number of examples of interspecific interactions among alpine herbivores to assess the role of positive and negative interactions in structuring their communities. A meta‐analysis showed no clear trend in the strength and direction of interactions among alpine herbivores. Although studies were biased towards reporting significant negative inter actions, we found no evidence of competition dominating in harsh environments. Thus, our results only partially support the SGH, but directly challenge the dominant view among animal ecologists. Clearly, a sound theoretical framework is needed to include competition, positive and neutral interactions as potential mechanisms determining the structure of animal communities under differing environmental conditions, and the stress‐gradient hypothesis can provide a solid starting point.