Species responses of colonising beetles to variation in patch quality,number, and context in experimental aquatic landscapes

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Spatial and temporal dynamics of habitat selection across canopy gradients generates patterns of species richness and composition in aquatic beetles

Abstract 1. Colonisation is a critical ecological process influencing both population and community level dynamics by connecting spatially discrete habitat patches. How communities respond to both natural and anthropogenic disturbances, furthermore, requires a basic understanding of how any environmental change modifies colonisation rates. For example, disturbance‐induced shifts in the quantity of forest cover surrounding aquatic habitats have been associated with the distribution and abundance of numerous aquatic taxa. However, the mechanisms generating these broad and repeatable field patterns are unclear. 2. Such patterns of diversity could result from differential spatial mortality post colonisation, or from colonisation alone if species select sites non‐randomly along canopy coverage gradients. We examined the colonisation/oviposition dynamics of aquatic beetles in experimental ponds placed under both open and closed forest canopies. 3. Canopy coverage imposed a substantial behavioural filter on the colonisation and reproduction of aquatic beetles representing multiple trophic levels, and resulted in significantly higher abundance, richness, and oviposition activity in open canopy ponds. These patterns strengthened overtime; although early in the experiment, the most abundant beetle had similar abundance in open and closed ponds. However, its abundance subsequently declined and then most other species heavily colonised open canopy ponds. 4. The primary response of many aquatic species to disturbances that generate canopy coverage gradients surrounding aquatic ecosystems is behavioural. The magnitude of the colonisation responses reported here rivals, if not exceeds, those produced by predators, suggesting that aquatic landscapes are behaviourally assessed and partitioned across multiple environmental gradients. The community level structure produced solely by selective colonisation, is predicted to strongly modify how patch area and isolation affect colonisation rates and the degree to which communities are linked by the flux of individuals and species.     

Both local presence and regional distribution of predator cues modulate prey colonisation in pond landscapes

Recent work on habitat selection has shown that the perceived quality of habitat patches may depend on the quality of adjacent patches. However, it is still unclear how local habitat selection cues can alter distribution patterns in metacommunities at a larger (regional) scale. We studied mosquito oviposition in pond landscapes that differed in the proportion of bad patches with fish predation risk. Our experiment provided conclusive evidence for two local and two regional types of habitat selection. Good patches near bad patches were avoided (local risk contagion) while more distant good patches experienced increased oviposition (regional compression). Oviposition in bad patches increased when located next to good patches (reward contagion) or when there were no good patches regionally present (regional compromise). This complex colonisation behaviour involving compromises at different spatial scales forces experimenters to reconsider the independence of spatial replicates and challenges available theories to predict species distribution patterns.     

Habitat patch size alters the importance of dispersal for species diversity in an experimental freshwater community

Increased dispersal of individuals among discrete habitat patches should increase the average number of species present in each local habitat patch. However, experimental studies have found variable effects of dispersal on local species richness. Priority effects, predators, and habitat heterogeneity have been proposed as mechanisms that limit the effect of dispersal on species richness. However, the size of a habitat patch could affect how dispersal regulates the number of species able to persist. We investigated whether habitat size interacted with dispersal rate to affect the number of species present in local habitats. We hypothesized that increased dispersal rates would positively affect local species richness more in small habitats than in large habitats, because rare species would be protected from demographic extinction. To test the interaction between dispersal rate and habitat size, we factorially manipulated the size of experimental ponds and dispersal rates, using a model community of freshwater zooplankton. We found that high‐dispersal rates enhanced local species richness in small experimental ponds, but had no effect in large experimental ponds. Our results suggest that there is a trade‐off between patch connectivity (a mediator of dispersal rates) and patch size, providing context for understanding the variability observed in dispersal effects among natural communities, as well as for developing conservation and management plans in an increasingly fragmented world.     

Spatial and temporal variation in patch occupancy and population density in a model system of an arctic Collembola species assemblage

We employed an experimental model system to investigate the mechanisms underlying patterns of patch occupancy and population density in a high arctic assemblage of Collembola species inhabiting a sedge tussock landscape on Svalbard. The replicate model systems consisted of 5 cores of the tussocks (habitat patches) imbedded in a barren matrix. Four of the patches were open so that animals could migrate between them, while there was one closed patch per system to test the effect of migration on extinction rate. There were model systems of two types: one with long and one with short inter‐patch distances to test the effect of patch isolation on colonisation and extinction rates. Each of the four most common collembolan species at the field site were introduced to two open patches per system (source patches), with the other two functioning as colonisation patches for the species. The experiment was run in an ecotrone over three identical, simulated arctic summers separated by winters of 3 weeks. Six replicates of systems with short and long inter‐patch distances were sampled at the end of each summer. The species varied markedly in their performance in both open arenas and closed patches, indicating differential responses to patch humidity, consistent with their differential distribution along the moisture gradient in the field site. The extinction – colonisation dynamics differed markedly between species as predicted from our field studies. This could partly be ascribed to differential dispersal and colonisation ability, but also to different tolerance to spatially variable patch quality and/or tendency for aggregative behaviour. Three of the species exhibited dynamics that superficially resemble what could be expected from classical metapopulation dynamics. However, there was a striking discrepancy between what would be expected from the effect of migration on the extinction rate of isolated patches (in particular closed patches) and the observed rates. Thus, metapopulation processes, such as stochastic colonisation and extinction events due to demographic stochasticity, were relatively unimportant compared to other sources of spatial variability among which subtle differences in patch quality are probably most important. We discuss the value of combining field studies with model system experiments, in particular when habitat quality cannot easily be measured in the field. However, our field and laboratory studies also emphasise the need for a thorough knowledge of species‐specific life history traits for making biologically sound interpretations based on both observational and experimental data.     

Patch size influences perceived patch quality for colonising Culex mosquitoes

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Host patch traits have scale-dependent effects on diversity in a stickleback parasite metacommunity

Many metacommunities are distributed across habitat patches that are themselves aggregated into groups. Perhaps the clearest example of this nested metacommunity structure comes from multi-species parasite assemblages, which occupy individual hosts that are aggregated into host populations. At both spatial scales, we expect parasite community diversity in a given patch (either individual host or population) to depend on patch characteristics that affect colonization rates and species sorting. But, are these patch effects consistent across spatial scales? Or, do different processes govern the distribution of parasite community diversity among individual hosts, versus among host patches? To answer these questions, we document the distribution of parasite richness among host individuals and among populations in a metapopulation of threespine stickleback Gasterosteus aculeatus. We find some host traits (host size, gape width) are associated with increased parasite richness at both spatial scales. Other patch characteristics affect parasite richness only among individuals (sex), or among populations (lake size, lake area, elevation and population mean heterozygosity). These results demonstrate that some rules governing parasite richness in this metacommunity are shared across scales, while others are scale-specific.     

Between‐Patch Bird Movements within a High‐Andean Polylepis Woodland/Matrix Landscape: Implications for Habitat Restoration

The movement ability of species in fragmented landscapes must be considered if habitat restoration strategies are to allow maximum benefit in terms of increased or healthier wildlife populations. We studied movements of a range of bird species between woodland patches within a high‐altitude Polylepis/matrix landscape in the Cordillera Vilcanota, Peru. Movement rates between Polylepis patches differed across guilds, with arboreal omnivores, arboreal sally‐strikers and nectarivores displaying the highest movement rates, and understorey guilds and arboreal sally‐gleaners the lowest movement rates. Birds tend to avoid flights to more distant neighboring patches, especially when moving from patches which were themselves isolated. The decline in bird flight frequencies with increasing patch isolation followed broken‐stick models most closely, and while we suggest that there is evidence for a decline in between‐patch movements over distances of 30–210 m, there was great variability in movement rates across individual patches. This variability is presumably a result of complex interactions between patch size, quality and configuration, and flight movement patterns of individual bird species. Our study does, however, highlight the contribution small woodland patches make toward fragmented Polylepis ecosystem functioning, and we suggest that, where financial resources permit, small patch restoration would be an important compliment to the restoration of larger woodland patches. Most important is that replanting takes place within 200 m or so of existing larger patches. This will be especially beneficial in allowing more frequent use of woodland elements within the landscape and in improving the total area of woodland patches that are functionally connected.     

Colonisation rate and adaptive foraging control the emergence of trophic cascades

Ecological communities are assembled and sustained by colonisation. At the same time, predators make foraging decisions based on the local availabilities of potential resources, which reflects colonisation. We combined field and laboratory experiments with mathematical models to demonstrate that a feedback between these two processes determines emergent patterns in community structure. Namely, our results show that prey colonisation rate determines the strength of trophic cascades – a feature of virtually all ecosystems – by prompting behavioural shifts in adaptively foraging omnivorous fish predators. Communities experiencing higher colonisation rates were characterised by higher invertebrate prey and lower producer biomasses. Consequently, fish functioned as predators when colonisation rate was high, but as herbivores when colonisation rate was low. Human land use is changing habitat connectivity worldwide. A deeper quantitative understanding of how spatial processes modify individual behaviour, and how this scales to the community level, will be required to predict ecosystem responses to these changes.     

The effects of habitat manipulation on population distribution and foraging behavior in meadow voles

Individuals, free to choose between different habitat patches, should settle among them such that fitness is equalized. Alternatives to this ideal free distribution result into fitness differences among the patches. The concordance between fitnesses and foraging costs among inhabitants of different quality patches, demonstrated in recent studies, suggests that the mode of habitat selection and the resulting fitness patterns may have important implications to the resource use of a forager and to the survival of its prey. We studied how coarse scale selection between habitat patches of different quality and quitting harvest rate in these patches are related to each other and to fine scale patch use in meadow voles (Microtus pennsylvanicus). To demonstrate these relationships, we manipulated habitat patches within large field enclosures by mowing vegetative cover and adding supplemental food according to a 2×2 factorial design. We tracked vole population densities, collected giving‐up densities (GUDs, a measure of patch quitting harvest rate), and monitored the removal of seeds from lattice grids with 1.5 m intervals (an index of fine‐scale space use) in the manipulated habitat patches. Changes in habitat quality induced changes in habitat use at different spatial scales. In preferred habitats with intact cover, voles were despotic and GUDs were low, but increased with the addition of food. In contrast, voles in less‐preferred mowed habitats settled into an ideal free distribution, GUDs were high and uninfluenced by the addition of food. Seed removal was enhanced by the presence of cover but inhibited by supplemental food. Across all treatments, vole densities and GUDs were strongly correlated making it impossible to separate their effects on seed removal rates. However, this relationship broke down in unmowed habitats, where GUDs rather than vole density primarily influenced seed removal by voles. GUDs and seed removal correlated with predation on tree seedlings formerly planted into the enclosures, demonstrating the mechanisms between coarse‐scale habitat manipulations and community level consequences on a forager's prey.     

Within‐patch oviposition site shifts by spider mites in response to prior predation risks decrease predator patch exploitation

In egg‐laying animals with no post‐oviposition parental care, between‐ or within‐patch oviposition site selection can determine offspring survival. However, despite the accumulation of evidence supporting the substantial impact predators have on oviposition site selection, few studies have examined whether oviposition site shift within patches (“micro‐oviposition shift”) reduces predation risk to offspring. The benefits of prey micro‐oviposition shift are underestimated in environments where predators cannot disperse from prey patches. In this study, we examined micro‐oviposition shift by the herbivorous mite Tetranychus kanzawai in response to the predatory mite, Neoseiulus womersleyi, by testing its effects on predator patch exploitation in situations where predatory mites were free to disperse from prey patches. Adult T. kanzawai females construct three‐dimensional webs on leaf surfaces and usually lay eggs under the webs; however, females that have experienced predation risks, shift oviposition sites onto the webs even in the absence of current predation risks. We compared the predation of eggs on webs deposited by predator‐experienced females with those on leaf surfaces. Predatory mites left prey patches with more eggs unpredated when higher proportions of prey eggs were located on webs, and egg survival on webs was much higher than that on leaf surfaces. These results indicate that a micro‐oviposition shift by predator‐experienced T. kanzawai protects offspring from predation, suggesting adaptive learning and subsociality in this species. Conversely, fecundity and longevity of predator‐experienced T. kanzawai females were not reduced compared to those of predator‐naïve females; we could not detect any costs associated with the learned micro‐oviposition shift. Moreover, the previously experienced predation risks did not promote between‐patch dispersal of T. kanzawai females against subsequently encountered predators. Based on these results, the relationships of between‐patch oviposition site selection and micro‐oviposition shift are discussed.     

Does variation in patch use among butterfly species contribute to nestedness at fine spatial scales?

A nested pattern occurs whenever the species observed in depauperate habitat patches are a subset of those found in more species‐rich patches. Ecologists have documented many instances of nestedness caused by population‐level processes such as colonization and extinction at biogeographic scales. However, few researchers have examined whether nestedness may exist at fine scales due to the ways in which individual organisms discriminate among potential habitat patches. In 1999, we experimentally fragmented an old‐field habitat into patches of varying size to test whether nestedness could exist on a fine spatial scale. Five treatments of differing patch size were replicated five times in a Latin square design by selectively mowing 15×15 m² plots within an old‐field (patch areas: 225, 180, 135, 90, and 45 m²). Specifically, we tested whether butterflies foraging within a network of patches differing in area conformed to a nested subset structure. We also classified species according to (1) their flight height while foraging (high or low), and (2) their adult habitat breadth (ubiquitous, general, or restricted) to determine whether nestedness could be explained by difference in species’ tendency to discriminate among patches differing in area.
We found significant evidence that a community of foraging Lepidoptera conformed to a nested subset structure based on the difference between the observed nestedness within the butterfly community and the nestedness obtained from randomly generated species presence/absence matrices. Poisson regression analyses demonstrated that high‐flying, habitat‐restricted species avoided the smallest patches (90 and 45 m²) in favor of larger remnants, whereas low‐flying, habitat generalists used all patch sizes. Thus, our study is one of the first to demonstrate that nestedness among species subsets can be observed at fine spatial scales (within a single 1.5 hectare field) and may be maintained by species behavioral differences: discriminating species (i.e. high‐flying, habitat restricted) avoided the smallest patches, and less discriminating species (i.e. low‐flying, ubiquitous) were distributed throughout the field without regard to patch size. Our results also suggest that nestedness should be viewed as yet another scalar pattern in ecology, generated by variation in patch use by individuals at fine‐scales as well as the more traditionally invoked processes of extinction and colonization of species at broad‐scales.     

Rethinking patch size and isolation effects: the habitat amount hypothesis

I challenge (1) the assumption that habitat patches are natural units of measurement for species richness, and (2) the assumption of distinct effects of habitat patch size and isolation on species richness. I propose a simpler view of the relationship between habitat distribution and species richness, the ‘habitat amount hypothesis’, and I suggest ways of testing it. The habitat amount hypothesis posits that, for habitat patches in a matrix of non‐habitat, the patch size effect and the patch isolation effect are driven mainly by a single underlying process, the sample area effect. The hypothesis predicts that species richness in equal‐sized sample sites should increase with the total amount of habitat in the ‘local landscape’ of the sample site, where the local landscape is the area within an appropriate distance of the sample site. It also predicts that species richness in a sample site is independent of the area of the particular patch in which the sample site is located (its ‘local patch’), except insofar as the area of that patch contributes to the amount of habitat in the local landscape of the sample site. The habitat amount hypothesis replaces two predictor variables, patch size and isolation, with a single predictor variable, habitat amount, when species richness is analysed for equal‐sized sample sites rather than for unequal‐sized habitat patches. Studies to test the hypothesis should ensure that ‘habitat’ is correctly defined, and the spatial extent of the local landscape is appropriate, for the species group under consideration. If supported, the habitat amount hypothesis would mean that to predict the relationship between habitat distribution and species richness: (1) distinguishing between patch‐scale and landscape‐scale habitat effects is unnecessary; (2) distinguishing between patch size effects and patch isolation effects is unnecessary; (3) considering habitat configuration independent of habitat amount is unnecessary; and (4) delineating discrete habitat patches is unnecessary.     

The nest predator community of grassland birds responds to agroecosystem habitat at multiple scales

Nest predation is the leading cause of reproductive failure for grassland birds of conservation concern. Understanding variation in nest predation rates is complicated by the diverse assemblage of species known to prey on nests. As part of a long‐term study of grassland bird ecology, we monitored populations of predators known to prey on grassland bird nests. We used information theoretic approach to examine the predator community's association with habitat at multiple scales, including local vegetation structure of grassland patches, spatial attributes of grassland patches (size and shape), and landscape composition surrounding grassland patches (land cover within 400 and 1600 m). Our results confirmed that nest predators respond to habitat at multiple scales and different predator species respond to habitat in different ways. The most informative habitat models we selected included variability in local vegetation (CV in the density of forbs), local patch (area and edge‐to‐interior ratio), and landscape within a 1600 m buffer around grasslands (percent of land covered by human structures and development). As a separate question, we asked if models that incorporated information from multiple scales simultaneously might improve the ability to explain variation in the predator community. Multi‐ scale models were not consistently superior to models derived from variables focused at a single spatial scale. Our results suggest that minimizing human development on and surrounding conservation land and the management of the vegetation structure on grassland fragments both may benefit grassland birds by decreasing the risk of nest predation.     

Foraging behavior of an estuarine predator, the blue crab Callinectes sapidus in a patchy environment

To define general principles of predator‐prey dynamics in an estuarine subtidal environment, we manipulated predator density (the blue crab, Callinectes sapidus) and prey (the clam, Macoma balthica) patch distribution in large field enclosures in the Rhode River subestuary of the central Chesapeake Bay. The primary objectives were to determine whether predators forage in a way that maximizes prey consumption and to assess how their foraging success is affected by density of conspecifics. We developed a novel ultrasonic telemetry system to observe behavior of individual predators with unprecedented detail. Behavior of predators was more indicative of optimal than of opportunistic foraging. Predators appeared responsive to the overall quality of prey in their habitat. Rather than remaining on a prey patch until depletion, predators appeared to vary their patch use with quality of the surrounding environment. When multiple (two) prey patches were available, residence time of predators on a prey patch was shorter than when only a single prey patch was available. Predators seemed to move among the prey patches fairly regularly, dividing their foraging time between the patches and consuming prey from each of them at a similar rate. That predators more than doubled their consumption of prey when we doubled the number of prey (by adding the second patch) is consistent with optimizing behaviors ‐ rather than with an opportunistic increase in prey consumption brought about simply by the addition of more prey. Predators at high density, however, appeared to interfere with each other's foraging success, reflected by their lower rates of prey consumption. Blue crabs appear to forage more successfully (and their prey to experience higher mortality) in prey patches located within 15–20 meters of neighboring patch, than in isolated patches. Our results are likely to apply, at least qualitatively, to other crustacean‐bivalve interactions, including those of commercial interest; their quantitative applicability will depend on the mobility of other predators and the scale of patchiness they perceive.     

Spatio‐temporal dynamics of density‐dependent dispersal during a population colonisation

Predicting population colonisations requires understanding how spatio‐temporal changes in density affect dispersal. Density can inform on fitness prospects, acting as a cue for either habitat quality, or competition over resources. However, when escaping competition, high local density should only increase emigration if lower‐density patches are available elsewhere. Few empirical studies on dispersal have considered the effects of density at the local and landscape scale simultaneously. To explore this, we analyze 5 years of individual‐based data from an experimental introduction of wild guppies Poecilia reticulata. Natal dispersal showed a decrease in local density dependence as density at the landscape level increased. Landscape density did not affect dispersal among adults, but local density‐dependent dispersal switched from negative (conspecific attraction) to positive (conspecific avoidance), as the colonisation progressed. This study demonstrates that densities at various scales interact to determine dispersal, and suggests that dispersal trade‐offs differ across life stages.     

Metacommunity,mainland‐island system or island communities? Assessing the regional dynamics of plant communities in a fragmented landscape

Understanding the regional dynamics of plant communities is crucial for predicting the response of plant diversity to habitat fragmentation. However, for fragmented landscapes the importance of regional processes, such as seed dispersal among isolated habitat patches, has been controversially debated. Due to the stochasticity and rarity of among‐patch dispersal and colonization events, we still lack a quantitative understanding of the consequences of these processes at the landscape‐scale. In this study, we used extensive field data from a fragmented, semi‐arid landscape in Israel to parameterize a multi‐species incidence‐function model. This model simulates species occupancy pattern based on patch areas and habitat configuration and explicitly considers the locations and the shapes of habitat patches for the derivation of patch connectivity. We implemented an approximate Bayesian computation approach for parameter inference and uncertainty assessment. We tested which of the three types of regional dynamics – the metacommunity, the mainland‐island, or the island communities type – best represents the community dynamics in the study area and applied the simulation model to estimate the extinction debt in the investigated landscape. We found that the regional dynamics in the patch‐matrix study landscape is best represented as a system of highly isolated ‘island’ communities with low rates of propagule exchange among habitat patches and consequently low colonization rates in local communities. Accordingly, the extinction rates in the local communities are the main drivers of community dynamics. Our findings indicate that the landscape carries a significant extinction debt and in model projections 33–60% of all species went extinct within 1000 yr. Our study demonstrates that the combination of dynamic simulation models with field data provides a promising approach for understanding regional community dynamics and for projecting community responses to habitat fragmentation. The approach bears the potential for efficient tests of conservation activities aimed at mitigating future losses of biodiversity.     

Migration of the clouded Apollo butterfly Parnassius mnemosyne in a network of suitable habitats – effects of patch characteristics

The clouded Apollo Parnassius mnemosyne is a food plant specialist with short but frequent movements between habitat patches. The short average dispersal distances suggest that the probability of colonisation of vacant patches decreases rapidly as the distance between the source and target patches increases, which means that a dense habitat network is needed for the conservation of the species. Both emigration rate and the number of immigrants varied among patches and were not affected only by isolation but also by several other patch characteristics. The model that explained most of the variation in emigration rates among patches included patch area and the number of conspecifics. The area and the population density of the target patch had significant effects on the number of arriving immigrants. Thus, the colonisation of vacant patches is dependent on these patch characteristics. Generally, emigration rates were lower and residence times longer in large patches with many conspecifics. Butterfly density was the most important single factor explaining the variation in the number of immigrants among patches, although the positive effect of the area of the target patch was also significant. As a consequence of the marked positive density dependence caused by conspecific attraction, small patches with higher than average butterfly density, receive more immigrants than could be expected based on the patch area only. Due to conspecific attraction, per capita immigration rates are higher in small than large patches. Thus, immigration may have a more significant effect on the local dynamics of small than large populations.     

Disease hotspots or hot species? Infection dynamics in multi‐host metacommunities controlled by species identity,not source location

Pathogen persistence in host communities is influenced by processes operating at the individual host to landscape‐level scale, but isolating the relative contributions of these processes is challenging. We developed theory to partition the influence of host species, habitat patches and landscape connectivity on pathogen persistence within metacommunities of hosts and pathogens. We used this framework to quantify the contributions of host species composition and habitat patch identity on the persistence of an amphibian pathogen across the landscape. By sampling over 11 000 hosts of six amphibian species, we found that a single host species could maintain the pathogen in 91% of observed metacommunities. Moreover, this dominant maintenance species contributed, on average, twice as much to landscape‐level pathogen persistence compared to the most influential source patch in a metacommunity. Our analysis demonstrates substantial inequality in how species and patches contribute to pathogen persistence, with important implications for targeted disease management.