Designing Effective Marine Protected Areas in Seaflower Biosphere Reserve, Colombia, Based on Biological and Sociological Information

Abstract:   Ecologists have paid increasing attention to the design of marine protected areas (MPAs), and their design advice consistently recommends representing all habitat types within MPAs or MPA networks as a means to provide protection to all parts of the natural ocean system. Recent developments of new habitat-mapping techniques make this advice more achievable, but the success of such an approach depends largely on our ability to define habitat types in a way that is ecologically relevant. We devised and tested the ecological relevance of a set of habitat-type definitions through our participation in a stakeholder-driven process to design a network of MPAs, focusing on no-take marine reserves in the Seaflower Biosphere Reserve, San Andrés Archipelago, Colombia. A priori definitions of habitat types were ecologically relevant, in that our habitat-type definitions corresponded to identifiable and unique characteristics in the ecological communities found there. The identification of ecological pathways and connectivity among habitats also helped in designing ecologically relevant reserve boundaries. Our findings contributed to the overall design process, along with our summary of other general principles of marine reserve design. Extensive stakeholder input provided information concerning the resources and their patterns of use. These inputs also contributed to the reserve design process. We anticipate success for the Seaflower Biosphere Reserve at achieving conservation and social goals because its zoning process includes detailed yet flexible scientific advice and the participation of stakeholders at every step .     

Habitat classification modeling with incomplete data: pushing the habitat envelope.

Habitat classification models (HCMs) are invaluable tools for species conservation, land-use planning, reserve design, and metapopulation assessments, particularly at broad spatial scales. However, species occurrence data are often lacking and typically limited to presence points at broad scales. This lack of absence data precludes the use of many statistical techniques for HCMs. One option is to generate pseudo-absence points so that the many available statistical modeling tools can bb used. Traditional techniques generate pseudo-absence points at random across broadly defined species ranges, often failing to include biological knowledge concerning the species-habitat relationship. We incorporated biological knowledge of the species-habitat relationship into pseudo-absence points by creating habitat envelopes that constrain the region from which points were randomly selected. We define a habitat envelope as an ecological representation of a species, or species feature's (e.g., nest) observed distribution (i.e., realized niche) based on a single attribute, or the spatial intersection of multiple attributes. We created HCMs for Northern Goshawk (Accipiter gentilis atricapillus) nest habitat during the breeding season across Utah forests with extant nest presence points and ecologically based pseudo-absence points using logistic regression. Predictor variables were derived from 30-m USDA Landfire and 250-m Forest Inventory and Analysis (FIA) map products. These habitat-envelope-based models were then compared to null envelope models which use traditional practices for generating pseudo-absences. Models were assessed for fit and predictive capability using metrics such as kappa, threshold-independent receiver operating characteristic (ROC) plots, adjusted deviance (D(adj)2), and cross-validation, and were also assessed for ecological relevance. For all cases, habitat envelope-based models outperformed null envelope models and were more ecologically relevant, suggesting that incorporating biological knowledge into pseudo-absence point generation is a powerful tool for species habitat assessments. Furthermore, given some a priori knowledge of the species-habitat relationship, ecologically based pseudo-absence points can be applied to any species, ecosystem, data resolution, and spatial extent.     

Oxygen/sulfide regime and nematode fauna associated with Arenicola marina burrows: new insights in the thiobios case

Profiles of oxygen and sulfide around the burrows of the lugworm, Arenicola marina, from a North Sea tidal flat were examined with microelectrodes, and the steep gradients were related to the microdistribution of nematodes. Around the tail shaft free oxygen penetrated only 2 mm into the burrow wall, coinciding with a bright zone sharply limited by the ambient black sediment. Contrastingly, in normal bottoms of the tidal flat (controls) only the surface of the bright zone was supplied with free oxygen. Here, the dark colouration coincided with the presence of free hydrogen sulfide. Around the tail shaft the nearest free hydrogen sulfide was detected 6 mm from the burrow wall leaving several millimetres of black sediment without measurable free sulfide. We discuss how these divergencies may relate to the stability of the oxygen/sulfide gradients and the course of time involved in their formation. A total of 54 nematode species were identified. Based on non-metric Multidimensional Scaling Ordination, four nematode assemblages corresponded to four microhabitats of the A. marina burrow: the funnel, the feeding pocket, the tail shaft and the feacal cast. The tail shaft assemblage (oxic plus partly anoxic zones) was similar to that of the anoxic zone of the control sediment. It was dominated by the most abundant nematode in the present study, Metalinhomoeus biformis (mean abundance in tail shaft 202 indx10 cm^-3). Adults of common nematode species from sulfidic microhabitats had a significantly higher length/diameter ratio than those inhabiting the oxic zone of the control sediment (p<0.001). The chemical recordings and metric analysis indicate that these slender nematodes around the A. marina tail shaft and in the reduced horizons of the reference sites represent thiobiotic assemblages, as compared to the shorter and stouter oxybiotic species characterising the assemblages from the surface zone and (partly) the funnel.     

Invasive ecosystem engineer selects for different phenotypes of an associated native species

Invasive habitat-forming ecosystem engineers modify the abiotic environment and thus represent a major perturbation to many ecosystems. Because native species often persist in these invaded habitats but have no shared history with the ecosystem engineer, the engineer may impose novel selective pressure on native species. In this study, we used a phenotypic selection framework to determine whether an invasive habitat-forming ecosystem engineer (the seaweed Caulerpa taxifolia) selects for different phenotypes of a common co-occurring native species (the bivalve Anadara trapezia). Compared to unvegetated habitat, Caulerpa habitat has lower water flow, lower dissolved oxygen, and sediments are more silty and anoxic. We determined the performance consequences of variation in key functional traits that may be affected by these abiotic changes (shell morphology, gill mass, and palp mass) for Anadara transplanted into Caulerpa and unvegetated habitat. Both linear and nonlinear performance gradients in Anadara differed between habitats, and these gradients were stronger in Caulerpa compared to unvegetated sediment. Moreover, in Caulerpa alternate phenotypes performed well, and these phenotypes were different from the dominant phenotype in unvegetated sediment. By demonstrating that phenotype-performance gradients differ between habitats, we have highlighted a role for Caulerpa as an agent of selection on native species.     

Trade‐offs and efficiencies in optimal budget‐constrained multispecies corridor networks

Conservation biologists recognize that a system of isolated protected areas will be necessary but insufficient to meet biodiversity objectives. Current approaches to connecting core conservation areas through corridors consider optimal corridor placement based on a single optimization goal: commonly, maximizing the movement for a target species across a network of protected areas. We show that designing corridors for single species based on purely ecological criteria leads to extremely expensive linkages that are suboptimal for multispecies connectivity objectives. Similarly, acquiring the least‐expensive linkages leads to ecologically poor solutions. We developed algorithms for optimizing corridors for multispecies use given a specific budget. We applied our approach in western Montana to demonstrate how the solutions may be used to evaluate trade‐offs in connectivity for 2 species with different habitat requirements, different core areas, and different conservation values under different budgets. We evaluated corridors that were optimal for each species individually and for both species jointly. Incorporating a budget constraint and jointly optimizing for both species resulted in corridors that were close to the individual species movement‐potential optima but with substantial cost savings. Our approach produced corridors that were within 14% and 11% of the best possible corridor connectivity for grizzly bears (Ursus arctos) and wolverines (Gulo gulo), respectively, and saved 75% of the cost. Similarly, joint optimization under a combined budget resulted in improved connectivity for both species relative to splitting the budget in 2 to optimize for each species individually. Our results demonstrate economies of scale and complementarities conservation planners can achieve by optimizing corridor designs for financial costs and for multiple species connectivity jointly. We believe that our approach will facilitate corridor conservation by reducing acquisition costs and by allowing derived corridors to more closely reflect conservation priorities.     

An ecological model of the habitat mosaic in estuarine nursery areas: Part I—Interaction of dispersal theory and habitat variability in describing juvenile fish distributions

Identification of critical habitat in estuarine nursery areas is an important conservation and management objective. Habitat can be viewed as a mosaic of both temporally variable environmental features and spatially variable structural features that combine to define optimal habitat. Effective models of juvenile distributions should account for individual movement, as well as the full suite of habitat variability including both spatial and temporal components. We have extended a terrestrial model of small-scale movement patterns to describe habitat choices of an index juvenile fish in an estuarine nursery system. Movement of small juvenile fishes was found to be influenced by both spatial and temporal patterns in habitat quality, and it was a balanced mix of both that resulted in an optimal distribution. Fishes that perceive habitat on a scale much smaller than the scale of spatial heterogeneity may respond to temporal change as a movement cue allowing for more deterministic outcomes at larger scales despite perceptual limitations. These model outcomes suggest a hierarchical approach is best for describing habitat choice in juvenile fishes and this approach will be used in the future to explore individual and population responses to predictable habitat change.     

Behavioral response of fish larvae to low dissolved oxygen concentrations in a stratified water column

Density stratification and respiration lead to vertical gradients in dissolved oxygen in many aquatic habitats. The behavioral responses of fish larvae to low dissolved oxygen in a stratified water column were examined during 1990–1991 with the goal of understanding how vertical gradients in dissolved oxygen may directly affect the distribution and survival of fish larvae in Chesapeake Bay, USA. In addition, the effects of low oxygen on 24-h survival rates were tested so that results of behavior experiments could be interpreted in the context of risk to the larve. Naked goby [Gobiosoma bosc (Lacépède)] and bay anchovy [Anchoa mitchilli (Valenciennes)] larvae strongly avoided dissolved oxygen concentrations <1 mg 1^-1, which were lethal within 24 h at 25 to 27°C. In addition, naked goby larvae, whose behavior was tested at a wider range of dissolved oxygen concentrations, also showed a reduced preference for an oxygen concentration of 2 mg 1^-1, which leads to reduced survival during long-term exposures and to reduced feeding rates. There were no major differences in behavior or survival between the two species, or between the two age classes of naked gobies tested. Results suggest that behavioral responses to oxygen gradients will play a large role in producing marked vertical changes in abundance of feeding-stage larvae in Chesapeake Bay; mortality from direct exposure to low oxygen will likely be much less important in producing vertical patterns of larval abundance.     

Synergistic interactions between edge and area effects in a heavily fragmented landscape

Both area and edge effects have a strong influence on ecological processes in fragmented landscapes, but there is little understanding of how these two factors might interact to exacerbate local species declines. To test for synergistic interactions between area and edge effects, we sampled a diverse beetle community in a heavily fragmented landscape in New Zealand. More than 35,000 beetles of approximately 900 species were sampled over large gradients in habitat area (10(-2) 10(6) ha) and distance from patch edge (2(0)-2(10) m from the forest edge into both the forest and adjacent matrix). Using a new approach to partition variance following an ordination analysis, we found that a synergistic interaction between habitat area and distance to edge was a more important determinant of patterns in beetle community composition than direct edge or area effects alone. The strength of edge effects in beetle-species composition increased nonlinearly with increasing fragment area. One important consequence of the synergy is that the slopes of species area (SA) curves constructed from habitat islands depend sensitively on the distance from edge at which sampling is conducted. Surprisingly, we found negative SA curves for communities sampled at intermediate distances from habitat edges, caused by differential edge responses of matrix- vs. forest-specialist species in fragments of increasing area. Our data indicate that distance to habitat edge has a consistently greater impact on beetle community composition than habitat area and that variation in the strength of edge effects may underlie many patterns that are superficially related to habitat area.     

Optimal movement strategies for social foragers in unpredictable environments

Spatial movement models often base movement decision rules on traditional optimal foraging theories, including ideal free distribution (IFD) theory, more recently generalized as density-dependent habitat selection (DDHS) theory, and the marginal value theorem (MVT). Thus optimal patch departure times are predicted on the basis of the density-dependent resource level in the patch. Recently, alternatives to density as a habitat selection criterion, such as individual knowledge of the resource distribution, conspecific attraction, and site fidelity, have been recognized as important influences on movement behavior in environments with an uncertain resource distribution. For foraging processes incorporating these influences, it is not clear whether simple optimal foraging theories provide a reasonable approximation to animal behavior or whether they may be misleading. This study compares patch departure strategies predicted by DDHS theory and the MVT with evolutionarily optimal patch departure strategies for a wide range of foraging scenarios. The level of accuracy with which individuals can navigate toward local food sources is varied, and individual tendency for conspecific attraction or repulsion is optimized over a continuous spectrum. We find that DDHS theory and the MVT accurately predict the evolutionarily optimal patch departure strategy for foragers with high navigational accuracy for a wide range of resource distributions. As navigational accuracy is reduced, the patch departure strategy cannot be accurately predicted by these theories for environments with a heterogeneous resource distribution. In these situations, social forces improve foraging success and have a strong influence on optimal patch departure strategies, causing individuals to stay longer in patches than the optimal foraging theories predict.     

Estimating the influence of overhead transmission power lines and landscape context on the density of little bustard Tetrax tetrax breeding populations

Collision with conductors and earth cables is a known impact generated by transmission power lines, however there is virtually no information on how these infrastructures might affect bird distribution in a landscape context. With this work we specifically hypothesise that transmission power lines may affect the occurrence of a threatened bird, the little bustard (Tetrax tetrax). To test this hypothesis we used a Stochastic Dynamic Methodology (StDM), analysing the effects of power lines in a landscape perspective and simulating population trends as a response to power line installation and habitat changes induced by agricultural shifts in southern Portugal. The data used in the dynamic model construction included relevant gradients of environmental conditions and was sampled during the breeding seasons of 2003-2006. Transmission power lines were significantly avoided by the little bustard and the developed StDM model showed that the distance to these utility structures is the most important factor determining breeding densities in sites with suitable habitat for the species, which possibly leads to displacement of populations and habitat fragmentation. The model simulations also provided the base to analyse the cumulative effects caused by the habitat degradation that can ultimately lead to the extinction of local populations. Within priority conservation sites, the dismantling of existing transmission lines should be considered whenever possible, in order to ensure adequate breeding habitat. The model is considered useful as an auxiliary tool to be used in environmental impact assessments, management and conservation studies.     

The Prairie Dog and Biotic Diversity

Since the turn of this century, prairie dog populations have declined as much as 98% throughout North America, largely as a result of prairie dog eradication programs. The prairie dog is a keystone species that plays an important role in maintaining the biotic integrity of the western grasslands that stretch from southern Canada to northern Mexico. The fragmentation of prairie dog distribution has degraded diversity on those prairies, and several species depending on prairie dogs have achieved listing status under the Endangered Species Act. We propose that managing the praire dog would provide an effective avenue from single-species management to management of a system. Because prairie dogs have declined so profoundly, some form of legal protection will be required. In addition, protected areas can preserve habitat and integrate ecologically sound agricultural opportunities. Positive incentives for ranchers to manage in the interests of both livestock and wildlife will enhance the attitude change necessary for grassland conservation. These management options hinge critically on an end to U.S. government subsidies for prairie dog eradication programs. The subsidies are financially and ecologically unsound, and they only contribute to the prevailing misconceptions about the role of the prairie dog on the grasslands.     

Evaluating Effects of Habitat Loss and Land-Use Continuity on Ant Species Richness in Seminatural Grassland Remnants

Abstract:  Seminatural grasslands in Europe are susceptible to habitat destruction and fragmentation that result in negative effects on biodiversity because of increased isolation and area effects on extinction rate. However, even small habitat patches of seminatural grasslands might be of value for conservation and restoration of species richness in a landscape with a long history of management, which has been argued to lead to high species richness. We tested whether ant communities have been negatively affected by habitat loss and increased isolation of seminatural grasslands during the twentieth century. We examined species richness and community composition in seminatural grasslands of different size in a mosaic landscape in Central Sweden. Grasslands managed continuously over centuries harbored species-rich and ecologically diverse ant communities. Grassland remnant size had no effect on ant species richness. Small grassland remnants did not harbor a nested subset of the ant species of larger habitats. Community composition of ants was mainly affected by habitat conditions. Our results suggest that the abandonment of traditional land use and the encroachment of trees, rather than the effects of fragmentation, are important for species composition in seminatural grasslands. Our results highlight the importance of considering land-use continuity and dispersal ability of the focal organisms when examining the effects of habitat loss and fragmentation on biodiversity. Landscape history should be considered in conservation programs focusing on effects of land-use change.     

How temperature and habitat quality affect parasitoid lifetime reproductive success—A simulation study

Adult parasitoid females lay their eggs in or on host insects. Most species are incapable of de novo lipogenesis as adults, and lipids accumulated during the larval stage are allocated either to egg production or to adult survival. Lipid consumption increases with distance covered by the parasitoids and thus with the distance between available hosts within a habitat. Temperature should affect parasitoid fitness because it changes the constraint imposed by a limited reserve of lipids and because it influences behaviour. Climate change involves both an increase in average temperature and an increased frequency of extreme weather such as heat waves. We investigated how the predicted increase of temperature will affect parasitoid fitness and how this depends on habitat parameters (spatial distribution of hosts and lipid cost of habitat exploitation). We studied optimal behaviour and calculated fitness at different temperatures and in different habitats using a stochastic dynamic programming model for pro-ovigenic parasitoids (which mature all their eggs before becoming adult). We show that an increase in temperature decreases fitness of parasitoids adapted to lower temperatures. This decrease in fitness depends on habitat quality. In field conditions (assuming small costs of intra-patch foraging), the loss of fitness should be larger in habitats with high inter-patch distance and in habitats with a more aggregated distribution of hosts. The foraging behaviour of parasitoids is also affected; at higher temperature we show that intra-patch foraging becomes less efficient, and patch residence times are longer.     

Evolution of within-colony distribution patterns of birds in response to habitat structure

It has long been suggested that habitat structure affects how colonial birds are distributed within their nesting aggregations, but this hypothesis has never been formally tested. The aim of this study was to test for a correlated evolution between habitat heterogeneity and within-colony distributions of Ciconiiformes by using Pagel’s general method of comparative analysis for discrete variables. The analysis indicated that central-periphery gradients of distribution (high-quality individuals occupying central nesting locations) prevail in species breeding in homogeneous habitats. These were mainly ground-nesting larids and spheniscids, where clear central-periphery patterns were recorded in ca. 80 % of the taxa. Since homogeneous habitats provide little variation in the physical quality of nest sites, central nesting locations should be largely preferred because they give better protection against predators by means of more efficient predator detection and deterrence. By contrast, central-periphery gradients tended to be disrupted in heterogeneous habitats, where 75 % of colonial Ciconiiform species showed uniform patterns of distribution. Under this model of distribution, edge nest sites of high physical quality confer higher fitness benefits in comparison to low-quality central sites, and thus, high-quality pairs are likely to choose nest sites irrespectively of their within-colony location. Breeding in homogeneous habitats and uniform distribution patterns were identified as probable ancestral states in Ciconiiformes, but there was a significant transition rate from uniform to central-periphery distributions in species occupying homogeneous habitats.     

Using Presence-Absence Data to Build and Test Spatial Habitat Models for the Fisher in the Klamath Region, U.S.A.

Abstract: Forest carnivores such as the fisher ( Martes pennanti ) have frequently been the target of conservation concern because of their association in some regions with older forests and sensitivity to landscape-level habitat alteration. Although the fisher has been extirpated from most of its former range in the western United States, it is still found in northwestern California. Fisher distribution, however, is still poorly known in most of this region where surveys have not been conducted. To predict fisher distribution across the region, we created a multiple logistic regression model using data from 682 previously surveyed locations and a vegetation layer created from satellite imagery. A moving-window function in a geographic information system was used to derive landscape-level indices of canopy closure, tree size class, and percent conifer. The model was validated with new data from 468 survey locations. The correct classification rate of 78.6% with the new data was similar to that achieved with the original data set (80.4%). Whereas several fine-scale habitat attributes were significantly correlated with fisher presence, the multivariate model containing only landscape- and regional-scale variables performed as well as one incorporating fine-scale data, suggesting that habitat selection by fishers may be dominated by factors operating at the home-range scale and above. Fisher distribution was strongly associated with landscapes with high levels of tree canopy closure. Regional gradients such as annual precipitation were also significant. At the plot level, the diameter of hardwoods was greater at sites with fisher detections. A comparison of regional fisher distribution with land-management categories suggests that increased emphasis on the protection of biologically productive, low- to mid-elevation forests is important to ensuring the long-term viability of fisher populations.     

Behaviour and distribution of the sand-beach amphipod Orchestoidea corniculata

The field distribution of Orchestoidea corniculata Stont a talitrid amphipod inhabiting the upper intertidal zone of sandy beaches, was measured by pitfall trap and subsurface core transects. Variations in the distribution of juveniles and adults were recorded. At Coal Oil Point (Goleta, California, USA), the population was concentrated on the leeward side of the Point and, with increasing distance from the Point, both the number and size of beachhoppers decreased. The amphipods burrowed in a 5 to 7 m wide band during the day, and were active over a 15 to 25 m area at night. When they emerged from their burrows, a net seaward movement of 3 m was observed for the population. Behavioural tests suggested that sand penetrability is an important factor determining the location of burrow sites. Diurnal orientation was investigated, and it was found that the amphipods oriented in a landward direction despite overcast sky conditions, beach slope, moisture gradients, or displacement to a new beach.     

Functional mechanisms underlying cetacean distribution patterns: hotspots for bottlenose dolphins are linked to foraging

Many studies have shown that the distribution of cetaceans can be closely linked to habitat, but the underlying function of the preferred habitats often remains unclear. Only when behavioural observations are made in relation to habitat types can functional mechanisms behind the habitat use be revealed. Within the range of a bottlenose dolphin (Tursiops truncatus) population off NE Scotland, dolphins show clear preferences for several discrete areas. If the observed patterns of distribution are related to foraging, we predict that behaviour patterns shown by dolphins would reflect this relationship. In this study we identify behaviours of dolphins at the water surface that were related to feeding events, evaluate whether the patterns of distribution were related to foraging and whether they were related the local submarine habitat characteristics. To investigate whether visible surface evidence of foraging behaviour varied spatially, we analysed data collected from 104 regular boat-based surveys made within the Moray Firth, NE Scotland, between 1990 and 2000. To determine whether underlying bathymetry had any influence on the surface behaviour of dolphins, a land-based observation study was carried out in the populations core region of use. The results of this study show that feeding behaviour by dolphins was significantly higher in areas used intensively by dolphins. Furthermore, there were clear relationships between feeding events and the submarine habitat characteristics; certain forms of feeding occur primarily over steep seabed gradients, and in deeper waters during June and July. These results quantitatively support the hypothesis that the distinctive patterns of distribution shown by these dolphins are related to foraging behaviour or opportunities, and that submarine habitat characteristics may be a significant factor in the foraging efficiency of dolphins. Future work should focus on collecting detailed information on the distribution patterns of prey within the study area to allow direct comparisons between predator and prey distributions.Communicated by J.P. Thorpe, Port Erin     

Contrasting ecological information content in whaling archives with modern cetacean surveys for conservation planning and identification of historical distribution changes

Many species are restricted to a marginal or suboptimal fraction of their historical range due to anthropogenic impacts, making it hard to interpret their ecological preferences from modern-day data alone. However, inferring past ecological states is limited by the availability of robust data and biases in historical archives, posing a challenge for policy makers . To highlight how historical records can be used to understand the ecological requirements of threatened species and inform conservation, we investigated sperm whale (Physeter macrocephalus) distribution in the Western Indian Ocean. We assessed differences in information content and habitat suitability predictions based on whale occurrence data from Yankee whaling logs (1792–1912) and from modern cetacean surveys (1995–2020). We built maximum entropy habitat suitability models containing static (bathymetry-derived) variables to compare models comprising historical-only and modern-only data. Using both historical and modern habitat suitability predictions  we assessed marine protected area (MPA) placement by contrasting suitability in- and outside MPAs. The historical model predicted high habitat suitability in shelf and coastal regions near continents and islands, whereas the modern model predicted a less coastal distribution with high habitat suitability more restricted to areas of steep topography. The proportion of high habitat suitability inside versus outside MPAs was higher when applying the historical predictions than the modern predictions, suggesting that different marine spatial planning optimums can be reached from either data sources. Moreover, differences in relative habitat suitability predictions between eras were consistent with the historical depletion of sperm whales from coastal regions, which were easily accessed and targeted by whalers, resulting in a modern distribution limited more to steep continental margins and remote oceanic ridges. The use of historical data can provide important new insights and, through cautious interpretation, inform conservation planning and policy, for example, by identifying refugee species and regions of anticipated population recovery.     

Males in the shade: habitat use and sexual segregation in the Galápagos sea lion (Zalophus californianus wollebaeki)

In many sexually dimorphic species adult sexes tend to segregate socially, spatially, or in habitat use. Several hypotheses have been formulated regarding underlying mechanisms. We investigated terrestrial habitat use and sexual segregation in a tropical otariid, the Galápagos sea lion (Zalophus californianus wollebaeki), where most of the hypotheses can be ruled out a priori. Factors relating to thermoregulation and costs of locomotion were of prime importance for habitat use. Habitats directly adjacent to the sea, with simple structured flat surfaces, shade, and tide pools were most frequented, but sexes and age classes differed in their usage patterns. Sexual segregation, both spatial and by habitat was pronounced in the reproductive period (RP), but remained high during the nonreproductive period (NRP). A GLM model of habitat use showed that in both seasons adult males frequented habitat types that adult females and other age classes used much less. Males were most abundant in suboptimal inland habitats, which offered only shade for cooling. Females with newborns differed in habitat use from females with older offspring and lone females. Spatial and habitat segregation are explained most parsimoniously as by-products of social processes, primarily intrasexual competition and female avoidance of male harassment, linked to the polygynous mating system.Electronic Supplementary Material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Slope,aspect and climate: Spatially explicit and implicit models of topographic microclimate in chalk grassland

The slope and aspect of a vegetated surface strongly affects the amount of solar radiation intercepted by that surface. Solar radiation is the dominant component of the surface energy balance and influences ecologically critical factors of microclimate, including near-surface temperatures, evaporative demand and soil moisture content. It also determines the exposure of vegetation to photosynthetically active and ultra-violet wavelengths. Spatial variation in slope and aspect is therefore a key determinant of vegetation pattern, species distribution and ecosystem processes in many environments. Slope and aspect angle may vary considerably over distances of a few metres, and fine-scale species’ distribution patterns frequently follow these topographic patterns. The availability of suitable microclimate at such scales may be critical for the response of species distributions to climatic change at much larger spatial scales. However, quantifying the relevant microclimatic gradients is not straightforward, as the potential variation in solar radiation flux under clear-sky conditions is modified by local and regional variations in cloud cover, and interacts with long-wave radiation exchange, local meteorology and surface characteristics.