Spatial variability in prey phenology determines predator movement patterns and prey survival

Species have phenological variation among local habitats that are located at relatively small spatial scales. However, less studies have tested how this spatial variability in phenology can mediate intra-/inter-specific interactions. When predators track phenological variation of prey among local habitats, survival of prey within a local habitat strongly influenced by phenological synchrony with their conspecifics in adjacent habitats. Theory predicts that phenological synchrony among local habitats increases prey survival in local habitat within spatially structured environments because the predators have to make a habitat choice for foraging. Consequently, total survival of prey at regional scale should be higher. By using a spatially explicit field experiment, we tested above hypothesis using a prey–predator interaction between tadpole (Rhacophorus arboreus) and newt (Cynops pyrrhogaster). We established enclosures (≈regional scale) consisting of two tanks (≈local habitat scale) with different degree of prey phenological synchrony. We found that phenological synchrony of prey between tanks within each enclosure decreased the mean residence time of the predator in each tank, which resulted in higher survival of prey at a local habitat scale, supporting the theoretical prediction. Furthermore, individual-level variation in predator residence time explained the between-tank variation in prey survival in enclosures with phenological synchrony, implying that movement patterns of the predator can mediate variation in local population dynamics of their prey. However, total survival at each enclosure was not higher under phenological synchrony. These results suggest the importance of relative timing of prey phenology, not absolute timing, among local habitats in determining prey–predator interactions.     

Spatiotemporal food web dynamics along a desert riparian–upland transition

Increased awareness of spatiotemporal variation in species interactions has motivated the study of temporally-resolved food web dynamics at the landscape level. Empiricists have focused attention on cross-habitat flows of materials, nutrients, and prey, largely ignoring the movement of predators between habitats that differ in productivity (and how predators integrate pulses in resource availability over time). We set out to study seasonal variation in food web interactions between mammalian carnivores and their rodent prey along a riparian–upland gradient in semi-arid southeastern Arizona which features both spatial and temporal heterogeneity in resource availability. Specifically, we tested whether mammalian carnivores spill over from productive, near-river habitats into adjacent, desert-scrub habitats; and if they do, to document the effects of this spillover on rodent communities. Furthermore, we examined seasonal variation in top-down effects by measuring changes in carnivore diet and distribution patterns and rodent populations over time. The results indicate that carnivores track seasonally-abundant resources across the landscape, varying both their diet and movement patterns. In turn, desert-scrub rodent population dynamics track seasonal shifts in carnivore habitat use but not resource availability, suggesting that predation plays a role in structuring rodent communities along the San Pedro River. Further evidence comes from data on rodent community composition, which differs between desert-scrub habitats near and far from the river, despite similarities in resource availability. Our data also suggest that seasonal omnivory helps predators survive lean times, increasing their effects on prey populations. Taken together, these results underscore the importance of spatiotemporal variation in species interactions, highlighting the complexity of natural systems and the need for further detailed studies of food web dynamics.     

Spatial variation in antler investment of Apennine red deer

Heterogeneity in resource availability and quality can trigger spatial patterns in the expression of sexually selected traits such as body mass and weaponry. While relationships between habitat features and phenotypic quality are well established at broad geographical scales, information is poor on spatial patterns at finer, intrapopulation scales. We analyzed biometric data collected on 1965 red deer Cervus elaphus males over 20 years from a nonmigratory population living on two sides of a mountainous ridge, with substantial differences in land cover and habitat quality but similar climate and population density. We investigate spatial patterns in (i) body mass, (ii) antler mass, and (iii) antler investment. We also tested for site‐ and age‐specific patterns in allometric relationship between body mass and antler mass. Statistically significant fine‐scale spatial variations in body mass, antler mass, and, to a lesser extent, antler allocation matched spatial differences in land cover. All three traits were greater in the northern slope, characterized by higher habitat heterogeneity and greater availability of open habitats, than in the southern slope. Moreover, the allometric relationship between body mass and antler mass differed among age‐classes, in a pattern that was consistent between the two mountain slopes. Our results support the occurrence of spatial patterns in the expression of individual attributes also at a fine, intrapopulation scale. Our findings emphasize the role of environmental heterogeneity in shaping spatial variations of key life‐history traits, with potential consequences for reproductive success.     

Contrasting bee foraging in response to resource scale and local habitat management

It is hypothesized that two main factors drive the foraging patterns of native and exotic species: food resource availability and habitat composition. These factors are particularly relevant for native bees and exotic honeybees, essential crop pollinators that are sensitive to floral resources and habitat management, and that have recently exhibited alarming population declines. Mechanisms driving native and exotic bee foraging patterns may critically depend on floral resource availability and habitat composition, yet the impacts of these factors on bee foraging have never been simultaneously analyzed. In a coffee producing region in southern Mexico, we investigated the influence of coffee floral resource levels and habitat management on native and exotic bee foraging. We measured the amount of flowering coffee available at multiple spatial scales within two distinct agroforestry habitat types (high-shade and low-shade coffee) and recorded visits to coffee flowers, documenting bee species, visit duration and visit frequency. We observed a significantly greater number of visits in high-shade coffee habitats than in low-shade coffee habitats for both native and exotic bees. In high-shade coffee habitats, native solitary bee and native social bee visitation decreased significantly in response to increasing floral resource availability, exhibiting a 'dilution effect' at the smallest spatial scale. In contrast, in low-shade coffee habitats, Africanized honeybees exhibited a 'concentration effect', increasing visitation significantly in response to increasing floral resource availability at the largest spatial scale. This study is the first to show that foraging patterns of native bees and exotic honeybees contrast in response to floral resource level and scale and that this response is mediated by the vegetation management of the local habitat.     

Genetic differentiation and gene flow among populations of the alpine butterfly, Parnassius smintheus, vary with landscape connectivity

Levels of gene flow among populations vary both inter- and intraspecifically, and understanding the ecological bases of variation in levels of gene flow represents an important link between the ecological and evolutionary dynamics of populations. The effects of habitat spatial structure on gene flow have received considerable attention; however, most studies have been conducted at a single spatial scale and without background data on how individual movement is affected by landscape features. We examined the influence of habitat connectivity on inferred levels of gene flow in a high-altitude, meadow-dwelling butterfly, Parnassius smintheus. For this species, we had background data on the effects of landscape structure on both individual movement and on small-scale population genetic differentiation. We compared genetic differentiation and patterns of isolation by distance, based on variation at seven microsatellite loci, among three regions representing two levels of connectivity of high-altitude, nonforested habitats. We found that reduced connectivity of habitats, resulting from more forest cover at high altitudes, was associated with greater genetic differentiation among populations (higher estimated FST), a breakdown of isolation by distance, and overall lower levels of inferred gene flow. These observed differences were consistent with expectations based on our knowledge of the movement behaviour of this species and on previous population genetic analyses conducted at the smaller spatial scale. Our results indicate that the role of gene flow may vary among groups of populations depending on the interplay between individual movement and the structure of the surrounding landscape.     

Subsidized Island Biogeography Hypothesis: another new twist on an old theory

We present a new hypothesis for predicting and describing patterns of species diversity on small islands and habitat fragments. We have modified the traditional island biogeography equilibrium theory to incorporate the influence of spatial subsidies from the surrounding matrix, which vary among islands and habitat fragments, on species diversities. The modification indicates three possible directions for the effects of spatial subsidies on diversity, which depend on where the focal community falls on the hypothesized unimodal curve of the productivity–diversity relationship. The idea is novel because no recent syntheses of productivity–diversity–area relationships examine the role of allochthonous resources on recipient communities’ diversity patterns.     

Patterns of genetic variability and habitat occupancy in Crepis triasii (Asteraceae) at different spatial scales: insights on evolutionary processes leading to diversification in continental islands

Background and Aims

Archipelagos are unique systems for studying evolutionary processes promoting diversification and speciation. The islands of the Mediterranean basin are major areas of plant richness, including a high proportion of narrow endemics. Many endemic plants are currently found in rocky habitats, showing varying patterns of habitat occupancy at different spatial scales throughout their range. The aim of the present study was to understand the impact of varying patterns of population distribution on genetic diversity and structure to shed light on demographic and evolutionary processes leading to population diversification in Crepis triasii, an endemic plant from the eastern Balearic Islands.

Methods

Using allozyme and chloroplast markers, we related patterns of genetic structure and diversity to those of habitat occupancy at a regional (between islands and among populations within islands) and landscape (population size and connectivity) scale.

Key Results

Genetic diversity was highly structured both at the regional and at the landscape level, and was positively correlated with population connectivity in the landscape. Populations located in small isolated mountains and coastal areas, with restricted patterns of regional occupancy, were genetically less diverse and much more differentiated. In addition, more isolated populations had stronger fine-scale genetic structure than well-connected ones. Changes in habitat availability and quality arising from marine transgressions during the Quaternary, as well as progressive fragmentation associated with the aridification of the climate since the last glaciation, are the most plausible factors leading to the observed patterns of genetic diversity and structure.

Conclusions

Our results emphasize the importance of gene flow in preventing genetic erosion and maintaining the evolutionary potential of populations. They also agree with recent studies highlighting the importance of restricted gene flow and genetic drift as drivers of plant evolution in Mediterranean continental islands.     

The effect of mean and variance in resource supply on survival of annuals from Mediterranean and desert environments

Resource availability is often characterized by mean annual amounts, while ignoring the spatial variation within habitats and the temporal variation within a year. Yet, temporal and spatial variation may be especially important for identifying the source of stress in low productivity environments such as deserts where resources are often pulsed and resource renewal events are separated by long periods of low resource availability. Therefore, the degree of stress will be determined in part by the length of time between recharge events. Here, we investigated the effect of timing and total amount of water application on two congeneric pairs, each with a population from a low (desert) and a high (Mediterranean) productivity habitat. As expected, highest survival and greatest growth were found at low or intermediate recharge intervals, and the magnitude of response to increases in total seasonal amounts was greater for Mediterranean species than desert species. The species that had greater survival switched in the hierarchy under high total water depending on interval length. These results demonstrate that temporal variation in resource availability can be as important as annual total amounts for plant performance and that response to temporal dynamics can vary between species. This has implications for community-level processes, as competitive hierarchies may switch based on resource dynamics rather than only total availability.     

Consumer movement through differentially subsidized habitats creates a spatial food web with unexpected results

Energy and nutrient flow between habitats, or allochthonous input, can have a significant impact on food web dynamics. Previous theory demonstrated that resource abundance decreases in habitats where consumers are subsidized. Here we examine the effect of subsidies that are available in localized parts of a habitat (such as near the shore in a marine‐subsidized terrestrial ecosystem) with a two‐patch model in which consumers move between patches, resources are stationary, and consumers receive the subsidy in only one of the two patches. In contrast to previous theory, our results show that subsidized consumers can increase resource abundance, though only in the subsidized patch. Furthermore, the total resource population responds positively to increasing consumer movement. These results demonstrate the importance of spatial heterogeneity in food web dynamics and the need for further examination of the role of space in multispecies trophic webs.     

Spatial patterns in abundance of a damselfish reflect availability of suitable habitat

For species with metapopulation structures, variation in abundance among patches can arise from variation in the input rate of colonists. For reef fishes, variability in larval supply frequently is invoked as a major determinant of spatial patterns. We examined the extent to which spatial variation in the amount of suitable habitat predicted variation in the abundance of the damselfish Dascyllus aruanus, an abundant planktivore that occupies live, branched coral throughout the Indo-Pacific. Reef surveys established that size, branching structure and location (proximity to sand) of the coral colonies together determined the ”suitability” of microhabitats for different ontogenetic stages of D. aruanus. Once these criteria were known, patterns of habitat use were quantified within lagoons of five Pacific islands. Availability of suitable habitat generally was an excellent predictor of density, and patterns were qualitatively consistent at several spatial scales, including among different lagoons on the same island, among different islands and between the central (French Polynesia and Rarotonga) and western (Great Barrier Reef, Australia) South Pacific. A field experiment that varied the amount of suitable coral among local plots indicated that habitat for settlers accounted for almost all of the spatial variation in the number of D. aruanus that settled at that location, suggesting that spatial patterns of abundance can be established at settlement without spatial variation in larval supply. Surveys of four other species of reef-associated fish revealed that a substantial fraction of their spatial variation in density also was explained by availability of suitable reef habitat, suggesting that habitat may be a prevalent determinant of spatial patterns. The results underscore the critical need to identify accurately the resource requirements of different species and life stages when evaluating causes of spatial variation in abundance of reef fishes. Received: 18 May 1999 / Accepted: 9 January 1999     

Density dependence in foraging habitat preference of eastern grey kangaroos

For a free‐ranging forager, the suitability of a patch is dependent on population density, resource supply, resource quality, and the costs of foraging or dispersal. We quantified differences among three foraging habitats and compared this variation to temporal patterns of habitat preference by free‐ranging eastern grey kangaroos, Macropus giganteus. We investigated selection on a fine‐grained spatial scale, and asked whether habitat preference is constrained by density‐dependent mechanisms. Variation in the quantity and quality of resources among habitats was greatest during spring, when biomass and quality were highest, and differences among habitats were most pronounced. However, consistent and discernable differences among habitats were not obtained, indicating that the system fluctuated around an equilibrium state. Using isodar regressions to examine the consumer‐density relationships among habitats, open‐woodland habitat was favoured over the two open‐forest habitats for foraging. Seasonal isodars indicated that density dependence regulated preference between the three foraging habitats during autumn, spring and summer, but not during winter, when variability in resources among habitats was lowest.     

Anthropogenic subsidies mitigate environmental variability for insular rodents

The exogenous input of nutrients and energy into island systems fuels a large array of consumers and drives bottom-up trophic cascades in island communities. The input of anthropogenic resources has increased on islands and particularly supplemented non-native consumers with extra resources. We test the hypothesis that the anthropogenic establishments of super-abundant gulls and invasive iceplants Carpobrotus spp. have both altered the dynamics of an introduced black rat Rattus rattus population. On Bagaud Island, two habitats have been substantially modified by the anthropogenic subsidies of gulls and iceplants, in contrast to the native Mediterranean scrubland with no anthropogenic inputs. Rats were trapped in all three habitats over two contrasting years of rainfall patterns to investigate: (1) the effect of anthropogenic subsidies on rat density, age-ratio and growth rates, and (2) the role of rainfall variability in modulating the effects of subsidies between years. We found that the growth rates of rats dwelling in the non-subsidized habitat varied with environmental fluctuation, whereas rats dwelling in the gull colony maintained high growth rates during both dry and rainy years. The presence of anthropogenic subsidies apparently mitigated environmental stress. Age ratio and rat density varied significantly and predictably among years, seasons, and habitats. While rat densities always peaked higher in the gull colony, especially after rat breeding in spring, higher captures of immature rats were recorded during the second year in all habitats, associated with higher rainfall. The potential for non-native rats to benefit from anthropogenic resources has important implications for the management of similar species on islands.     

Direct and indirect effects of area,energy and habitat heterogeneity on breeding bird communities

Aim To compare the ability of island biogeography theory, niche theory and species–energy theory to explain patterns of species richness and density for breeding bird communities across islands with contrasting characteristics. Location Thirty forested islands in two freshwater lakes in the boreal forest zone of northern Sweden (65°55′ N to 66°09′ N; 17°43′ E to 17°55′ E). Methods We performed bird censuses on 30 lake islands that have each previously been well characterized in terms of size, isolation, habitat heterogeneity (plant diversity and forest age), net primary productivity (NPP), and invertebrate prey abundance. To test the relative abilities of island biogeography theory, niche theory and species–energy theory to describe bird community patterns, we used both traditional statistical approaches (linear and multiple regressions) and structural equation modelling (SEM; in which both direct and indirect influences can be quantified). Results Using regression‐based approaches, area and bird abundance were the two most important predictors of bird species richness. However, when the data were analysed by SEM, area was not found to exert a direct effect on bird species richness. Instead, terrestrial prey abundance was the strongest predictor of bird abundance, and bird abundance in combination with NPP was the best predictor of bird species richness. Area was only of indirect importance through its positive effect on terrestrial prey abundance, but habitat heterogeneity and spatial subsidies (emerging aquatic insects) also showed important indirect influences. Thus, our results provided the strongest support for species–energy theory. Main conclusions Our results suggest that, by using statistical approaches that allow for analyses of both direct and indirect influences, a seemingly direct influence of area on species richness can be explained by greater energy availability on larger islands. As such, animal community patterns that seem to be in line with island biogeography theory may be primarily driven by energy availability. Our results also point to the need to consider several aspects of habitat quality (e.g. heterogeneity, NPP, prey availability and spatial subsidies) for successful management of breeding bird diversity at local spatial scales and in fragmented or insular habitats.     

Habitat effects on resource tracking ability: do wintering Blackcaps Sylvia atricapilla track fruit availability?

If resource availability shapes population distribution, changes in resource abundance should cause parallel changes in population numbers. However, tracking ability may be disrupted by different environmental and behavioural factors that act at different spatial and temporal scales. Here we analyse the ability of wintering Blackcap Sylvia atricapilla populations to track spatio-temporal variation in fruit availability in southern Spain in two habitats (forests and shrublands) with different population structure. Former studies had shown that forests are equally used by both adult migrant and local Blackcaps, whereas shrublands are nearly monopolized by juvenile migrants. These differences might affect resource tracking: it should be disrupted in forests, as local birds remain over winter in their breeding territories, but not in shrublands where similarly competitive juvenile migrants can freely track the spatial distribution of fruits. We analysed the fruit-tracking ability of Blackcap populations among sites and years in both habitat types using a habitat-matching model, which predicts spatio-temporal changes in population abundance proportional to changes in resource availability. We counted Blackcaps and fruiting shrubs (dominated by Lentiscs Pistacia lentiscus and Wild Olives Olea europaea sylvestris ) during four winters in forest and shrubland patches. The abundance of fruits was always higher in shrublands than in forests. In shrublands, Blackcaps seemed to move freely across fruit-rich habitat patches, tracking changes in fruiting-shrub abundance among sites and years. However, such tracking was not observed in forests. This supports the view that fruit-tracking ability may be constrained by local factors, such as the social structure of populations occurring in different habitat types, which introduces spatio-temporal variation in the way fruit availability shapes the abundance distribution of these birds in their Mediterranean wintering grounds.     

Trophic disruption: a meta‐analysis of how habitat fragmentation affects resource consumption in terrestrial arthropod systems

Habitat fragmentation is a complex process that affects ecological systems in diverse ways, altering everything from population persistence to ecosystem function. Despite widespread recognition that habitat fragmentation can influence food web interactions, consensus on the factors underlying variation in the impacts of fragmentation across systems remains elusive. In this study, we conduct a systematic review and meta‐analysis to quantify the effects of habitat fragmentation and spatial habitat structure on resource consumption in terrestrial arthropod food webs. Across 419 studies, we found a negative overall effect of fragmentation on resource consumption. Variation in effect size was extensive but predictable. Specifically, resource consumption was reduced on small, isolated habitat fragments, higher at patch edges, and neutral with respect to landscape‐scale spatial variables. In general, resource consumption increased in fragmented settings for habitat generalist consumers but decreased for specialist consumers. Our study demonstrates widespread disruption of trophic interactions in fragmented habitats and describes variation among studies that is largely predictable based on the ecological traits of the interacting species. We highlight future prospects for understanding how changes in spatial habitat structure may influence trophic modules and food webs.     

Spatio-Temporal Factors Shaping Diurnal Space Use by Pygmy Rabbits

Abstract: Factors influencing patterns of space use by pygmy rabbits (Brachylagus idahoensis) are poorly understood. We studied diurnal space use by adult pygmy rabbits during multiple breeding and nonbreeding seasons at 3 sites in the Lemhi Valley, Idaho, USA, during 2004–2005. Pygmy rabbits used larger areas than predicted by allometric models and documented by some previous investigations. Sex and season strongly influenced space use by rabbits. Males used larger home ranges and core areas, more burrow systems, and more widely dispersed burrow systems than did female rabbits. We also documented significant differences among study sites in many movement parameters, which suggested that local resource distribution also might influence how pygmy rabbits use space. Our results indicated that pygmy rabbits use large areas and exhibit seasonal, sex, and site-specific variation in patterns of movement and space use. Therefore, larger areas of habitat may be needed to conserve pygmy rabbits to accommodate seasonal, regional, and potentially annual variation in resource availability and to maintain linkages among populations.     

Where do species' geographic ranges stop and why? Landscape impermeability and the Afrotropical avifauna

Although understanding large-scale spatial variation in species'' distributions is a major goal in macroecology, relatively little attention has been paid to the factors limiting species'' ranges. An understanding of these factors may improve predictions of species'' movements in response to global change. We present a measure of landscape impermeability, defined as the proportion of resident species whose ranges end in an area. We quantify and map impermeability for Afrotropical birds and use multi-model inference to assess support for a wide suite of hypotheses about its potential environmental correlates. Non-spatial analyses emphasize the importance of broad-scale environmental patterns of energy availability and habitat heterogeneity in limiting species'' distributions. Conversely, spatial analyses focus attention on small-scale factors of habitat and topographic complexity. These results hold even when only species from the top quartile of range sizes are assessed. All our analyses highlight that range edges are concentrated in heterogeneous habitats. Global change is expected to alter the nature and distribution of such habitats, necessitating range movement by many resident species. Therefore, impermeability provides a simple measure for identifying regions, where continuing global change and human encroachment are likely to cause profound changes in regional diversity patterns.     

Factors shaping the range-size frequency distribution of the endemic fish fauna of the Tropical Eastern Pacific

Aim To assess the effect of habitat fragmentation and isolation in determining the range‐size frequency distribution (RFD) of the shorefish fauna endemic to a discrete biogeographical region. Location The Tropical Eastern Pacific (TEP). Methods Habitat isolation represents the separation between oceanic islands and the continental shore of the TEP and habitat fragmentation the degree of spatial continuity of habitats (i.e. reefs, soft bottom, nearshore waters) along the continental coast of the TEP. The effects of habitat isolation and fragmentation were quantified by comparing the RFDs of (1) the species found on oceanic islands vs. the continental shore, and (2) species on the continental shore that use different habitat types. Results The RFD of the entire TEP fauna was bimodal, with peaks at both small‐ and large‐range ends of the spectrum. The small‐range peak was due almost entirely to island species and the large‐range peak due mainly to species found in both the continental shore and oceanic islands. RFDs varied among species using different habitats on the continental shore: reef‐fishes had a right‐skewed RFD, soft‐bottom species a flat RFD, and coastal‐pelagic fishes a left‐skewed RFD. Main conclusions Variation in dispersal capabilities associated with habitat isolation and fragmentation in the TEP appears to be the main mechanism contributing to differences among RFD structure, although variation in tolerances arising from the dynamic regional environment may contribute to some patterns. Because diversity patterns are strongly affected by RFD structure, it is now evident that the insular and continental components of a fauna should be treated separately when analysing such patterns. Furthermore, contrasts in RFD structure among species using different habitats demonstrate that a full understanding of the causes of diversity patterns requires analyses of complete regional faunas in relation to regional geography.     

Population substructure and space use of Foxe Basin polar bears

Climate change has been identified as a major driver of habitat change, particularly for sea ice-dependent species such as the polar bear (Ursus maritimus). Population structure and space use of polar bears have been challenging to quantify because of their circumpolar distribution and tendency to range over large areas. Knowledge of movement patterns, home range, and habitat is needed for conservation and management. This is the first study to examine the spatial ecology of polar bears in the Foxe Basin management unit of Nunavut, Canada. Foxe Basin is in the mid-Arctic, part of the seasonal sea ice ecoregion and it is being negatively affected by climate change. Our objectives were to examine intrapopulation spatial structure, to determine movement patterns, and to consider how polar bear movements may respond to changing sea ice habitat conditions. Hierarchical and fuzzy cluster analyses were used to assess intrapopulation spatial structure of geographic position system satellite-collared female polar bears. Seasonal and annual movement metrics (home range, movement rates, time on ice) and home-range fidelity (static and dynamic overlap) were compared to examine the influence of regional sea ice on movements. The polar bears were distributed in three spatial clusters, and there were differences in the movement metrics between clusters that may reflect sea ice habitat conditions. Within the clusters, bears moved independently of each other. Annual and seasonal home-range fidelity was observed, and the bears used two movement patterns: on-ice range residency and annual migration. We predict that home-range fidelity may decline as the spatial and temporal predictability of sea ice changes. These new findings also provide baseline information for managing and monitoring this polar bear population.