Fine-scale Habitat Heterogeneity Research Articles

Eradicating an invasive mammal requires local elimination and reduced reinvasion from an urban source population.

Invasive mammal eradications are increasingly attempted across large, complex landscapes. Sequentially controlled management zones can be at risk of reinvasion from adjacent uncontrolled areas, and managers must weigh the relative benefits of ensuring complete elimination from a zone or minimizing reinvasion risk. This is complicated in urban areas, where habitat heterogeneity and a lack of baseline ecological knowledge increase uncertainty. We applied a spatial agent-based model to predict the reinvasion of a well-studied species, the brushtail possum (Trichosurus vulpecula), across an urban area onto a peninsula that is the site of an elimination campaign in Aotearoa New Zealand. We represented fine-scale urban habitat heterogeneity in a land cover layer and tested management scenarios that varied four factors: the density of possums remaining following an elimination attempt, the maintenance trap density on the peninsula, and effort expended toward preventing reinvasion by means of a high-density trap buffer at the peninsula isthmus or control of the source population adjacent to the peninsula. We found that achieving complete elimination on the peninsula was crucial to avoid rapid repopulation. The urban isthmus was predicted to act as a landscape barrier and restrict immigration onto the peninsula, but reliance on this barrier alone would fail to prevent repopulation. In combination, complete elimination, buffer zone, and source population control could reduce the probability of possum repopulation to near zero. Our findings support urban landscape barriers as one tool for sequential invasive mammal elimination but reaffirm that novel methods to expose residual individuals to control will be necessary to secure elimination in management zones. Work to characterize the urban ecology of many invasive mammals is still needed.

Habitat detection, habitat choice copying or mating benefits: What drives conspecific attraction in a nomadic songbird?

Conspecific attraction during habitat selection is common among animals, but the ultimate (i.e. fitness-related) reasons for this behaviour often remain enigmatic. We aimed to evaluate the following three hypotheses for conspecific attraction during the breeding season in male Wood Warblers (Phylloscopus sibilatrix): the habitat detection hypothesis, the habitat choice copying hypothesis and the female preference hypothesis. These hypotheses make different predictions with respect to the relative importance of social and nonsocial information during habitat assessment, and whether benefits accrue as a consequence of aggregation. We tested the above hypotheses using a combination of a 2-year playback experiment, spatial statistics and mate choice models. The habitat detection hypothesis was the most likely explanation for conspecific attraction and aggregation in male Wood Warblers, based on the following results: (1) males were attracted to conspecific song playbacks, but fine-scale habitat heterogeneity was the better predictor of spatial patterns in the density of settling males; (2) male pairing success did not increase, but instead slightly decreased, as connectivity with other males (i.e. the number and proximity of neighbouring males) increased. Our study highlights how consideration of the process by which animals detect and assess habitat, together with the potential fitness consequences of resulting aggregations, are important for understanding conspecific attraction and spatially clustered distributions.

Fine-Scale Heterogeneity of a Cold-Water Coral Reef and Its Influence on the Distribution of Associated Taxa

Benthic fauna form spatial patterns which are the result of both biotic and abiotic processes, which can be quantified with a range of landscape ecology descriptors. Fine- to medium-scale spatial patterns (<1–10 m) have seldom been quantified in deep-sea habitats, but can provide fundamental ecological insights into species’ niches and interactions. Cold-water coral reefs formed by Desmophyllum pertusum (syn. Lophelia pertusa) and Madrepora oculata are traditionally mapped and surveyed with multibeam echosounders and video transects, which limit the ability to achieve the resolution and/or coverage to undertake fine-scale, centimetric quantification of spatial patterns. However, photomosaics constructed from imagery collected with remotely operated vehicles (ROVs) are becoming a prevalent research tool and can reveal novel information at the scale of individual coral colonies. A survey using a downward facing camera mounted on a ROV traversed the Piddington Mound (Belgica Mound Province, NE Atlantic) in a lawnmower pattern in order to create 3D reconstructions of the reef with Structure-from-Motion techniques. Three high resolution orthorectified photomosaics and digital elevation models (DEM) >200 m2 were created and all organisms were geotagged in order to illustrate their point pattern. The pair correlation function was used to establish whether organisms demonstrated a clustered pattern (CP) at various scales. We further applied a point pattern modelling approach to identify four potential point patterns: complete spatial randomness (CSR), an inhomogeneous pattern influenced by environmental drivers, random clustered point pattern indicating biologically driven clustering and an inhomogeneous clustered point pattern driven by a combination of environmental drivers and biological effects. Reef framework presence and structural complexity determined inhabitant distribution with most organisms showing a departure from CSR. These CPs are likely caused by an affinity to local environmental drivers, growth patterns and restricted dispersion reproductive strategies within the habitat across a range of fine to medium scales. These data provide novel and detailed insights into fine-scale habitat heterogeneity, showing that non-random distributions are apparent and detectable at these fine scales in deep-sea habitats.

Fine-scale habitat heterogeneity favours the coexistence of supergene-controlled social forms in Formica selysi

BackgroundSocial insects vary widely in social organization, yet the genetical and ecological factors influencing this variation remain poorly known. In particular, whether spatially varying selection influences the maintenance of social polymorphisms in ants has been rarely investigated. To fill this gap, we examined whether fine-scale habitat heterogeneity contributes to the co-existence of alternative forms of social organization within populations. Single-queen colonies (monogyne social form) are generally associated with better colonization abilities, whereas multiple-queen colonies (polygyne social form) are predicted to be better competitors and monopolize saturated habitats. We hypothesize that each social form colonizes and thrives in distinct local habitats, as a result of their alternative dispersal and colony founding strategies. Here, we test this hypothesis in the Alpine silver ant, in which a supergene controls polymorphic social organization.ResultsMonogyne and polygyne colonies predominate in distinct habitats of the same population. The analysis of 59 sampling plots distributed across six habitats revealed that single-queen colonies mostly occupy unconnected habitats that were most likely reached by flight. This includes young habitats isolated by water and old habitats isolated by vegetation. In contrast, multiple-queen colonies were abundant in young, continuous and saturated habitats. Hence, alternative social forms colonize and monopolize distinct niches at a very local scale.ConclusionsAlternative social forms colonized and monopolized different local habitats, in accordance with differences in colonization and competition abilities. The monogyne social form displays a colonizer phenotype, by efficiently occupying empty habitats, while the polygyne social form exhibits a competitor phenotype, thriving in saturated habitats. The combination of the two phenotypes, coupled with fine-scale habitat heterogeneity, may allow the coexistence of alternative social forms within populations. Overall, these results suggest that spatially varying selection may be one of the mechanisms contributing to the maintenance of genetic polymorphisms in social organization.

Fine-scale habitat differentiation shapes the composition, structure and aboveground biomass but not species richness of a tropical Atlantic forest

Evaluating the influences of fine-scale habitat heterogeneity on the composition, diversity, structure and functioning of forests is critical to understand how tropical forests will respond to climate change and devise forest management strategies that will enhance biodiversity conservation and aboveground biomass stock. Here, we hypothesized that topographic and soil factors determine fine-scale habitat differentiation, which in turn shape community composition, species richness, structure and aboveground biomass at the local scale in tropical forests. To test this hypothesis, we selected two areas (each 100 × 100 m) with contrasting fine-scale topographic conditions where all trees, palms and lianas with a diameter at breast height ≥ 10 cm were tagged and identified to species. In each selected area, 100 subplots of 10 × 10 m were established. We mainly found that higher topographic variability caused higher habitat differentiation with changes in species composition and community structure, but did not change species richness. Our habitat-scale analyses indicated that, in the less heterogeneous area, the distribution of species was more uniform along a fine-scale topographical gradient with no variation in convexity, which induced changes in structure and aboveground biomass, but not in species richness. The nonsignificant relationship between species richness and aboveground biomass may be attributable to species redundancy or functional dominance. This study suggests that environmental filtering is a fundamental process for shaping community assembly and forest functioning along a local topographical gradient in tropical forests.

Habitat heterogeneity of hadal trenches: Considerations and implications for future studies

The hadal zone largely comprises a series of subduction trenches that do not form part of the continental shelf-slope rise to abyssal plain continuum. Instead they form geographically isolated clusters of deep-sea (6000–11,000 m water depth) environments. There is a growing realization in hadal science that ecological patterns and processes are not driven solely by responses to hydrostatic pressure, with comparable levels of habitat heterogeneity as observed in other marine biozones. Furthermore, this heterogeneity can be expressed at multiple scales from inter-trench levels (degrees of geographical isolation, and biochemical province), to intra-trench levels (variation between trench flanks and axis), topographical features within the trench interior (sedimentary basins, ridges, escarpments, ‘deeps’, seamounts) to the substrate of the trench floor (seabed-sediment composition, mass movement deposits, bedrock outcrop). Using best available bathymetry data combined with the largest lander-derived imaging dataset that spans the full depth range of three hadal trenches (including adjacent slopes); the Mariana, Kermadec and New Hebrides trenches, the topographic variability, fine-scale habitat heterogeneity and distribution of seabed sediments of these three trenches have been assessed for the first time. As well as serving as the first descriptive study of habitat heterogeneity at hadal depths, this study also provides guidance for future hadal sampling campaigns taking into account geographic isolation, total trench particulate organic matter flux, maximum water depth and area.

The potential of military training areas for bird conservation in a central European landscape

European biodiversity has suffered from serious declines during the past few decades, with alterations of land use practices resulting in a loss of fine-scale habitat heterogeneity being a dominant driver. This heterogeneity was maintained by extensive landscape management, which has gradually been replaced by either intensive exploitation or land abandonment. It has been suggested that military training can generate habitat heterogeneity that may support the existence of species of conservation concern, but studies rigorously testing the real importance of military training areas for biodiversity are lacking. Here we address this issue by analyses of two datasets. First, we compared land cover classes between all large military training areas (MTAs) and surrounding control areas (CAs) of the same size in the Czech Republic using multivariate redundancy analysis. We found that the difference in land cover between MTAs and CAs was significant and represented the strongest gradient in land cover classes: from various farmland and artificial habitats typical for CAs to forest and scrubland-grassland mosaic typical for MTAs. Second, we selected one of these areas and compared bird species richness between the MTA and the nearby CA using generalized linear mixed effects models. We found that the number of species of conservation concern was significantly higher in the MTA than in the CA. With respect to habitats, bird species richness was significantly higher in the MTA than in the CA for open habitats, but not for forest habitats. Our results are thus consistent with the view that military training creates areas that are different from the surrounding landscape in terms of land cover, and that this difference translates to a suitability for species of conservation concern. It is remarkable that the positive influence of military training is confined to open habitats, which are subject to the most intensive military activities and also suffer the highest degree of deterioration in European landscapes.

Environmental drivers of benthic communities and habitat heterogeneity on an East Antarctic shelf

AbstractThis study presents the first analysis of benthic megafauna and habitats from the Sabrina Coast shelf, encompassing a proposed Marine Protected Area. Sea bed imagery indicated an abundant benthic fauna compared to other parts of the Antarctic shelf, dominated by brittle stars, polychaete tubeworms, and a range of other sessile and mobile taxa. The distribution of taxa was related (ρ=0.592, P<0.001) to variations in water depth, latitude, substrate type and phytodetritus. High phytodetritus cover was associated with muddy/sandy sediments and abundant holothurians and amphipods, while harder substrates hosted abundant brachiopods, hard bryozoans, polychaete tubeworms, massive and encrusting sponges, and sea whips. Brittle stars, irregular urchins and anemones were ubiquitous. Variations in substrate largely reflected the distribution of dropstones, creating fine-scale habitat heterogeneity. Several taxa were found only on hard substrates, and their broad regional distribution indicated that the density of dropstones was sufficient for most sessile invertebrates to disperse across the region. The hexactinellid sponge Anoxycalyx joubini and branching hydrocorals exhibited a more restricted distribution, probably related to water depth and limited dispersal capability, respectively. Dropstones were associated with significant increases in taxa diversity, abundance and biological cover, enhancing the overall diversity and biomass of this ecosystem.

An association between a cusk eel (Bassozetus sp.) and a black coral (Schizopathes sp.) in the deep western Indian Ocean

Detailed observations in the deep sea can reveal previously unknown behaviour, species interactions and fine-scale habitat heterogeneity. Here, the first in situ images of the black coral Schizopathes sp. (Anthozoa: Antipatharia) in the deep western Indian Ocean have been obtained from remotely operated vehicle video footage and time-lapse photography. In these images, there appears to be an association with the cusk eel Bassozetus (Family: Ophidiidae). In the primary observation, chance encounters revealed the fish interacted with the anitpatharian on multiple occasions over several days. Subsequent time-lapse camera footage showed the fish remained almost exclusively underneath the antipatharian for the duration of a 30-h deployment. Excursions from the cover of the antipatharian were for less than 2 min. The primary observation is supported by two similar encounters in the same region. Observed reduction in the tail-beat frequency of the fish under the antipatharian suggests reduced energy requirements for the ophidiid in this position. The observations demonstrate the role that even individual coral colonies play as a source of three-dimensional structure, providing habitat heterogeneity in the deep sea.

Getting into the groove: Opportunities to enhance the ecological value of hard coastal infrastructure using fine-scale surface textures

Concrete flood defences, erosion control structures, port and harbour facilities, and renewable energy infrastructure are increasingly being built in the world’s coastal regions. There is, however, strong evidence to suggest that these structures are poor surrogates for natural rocky shores, often supporting assemblages with lower species abundance and diversity. Ecological engineering opportunities to enhance structures for biodiversity conservation (and other management goals) are therefore being sought, but the majority of work so far has concentrated on structural design features at the centimetre–meter scale.We deployed concrete tiles with four easily-reproducible fine-scale (millimetre) textures (control, smoothed, grooved and exposed aggregate) in the intertidal zone to test opportunities for facilitating colonisation by a dominant ecosystem engineer (barnacles) relative to natural rock. Concrete texture had a significant effect on colonisation; smoothed tiles supported significantly fewer numbers of barnacles, and those with intermediate roughness (grooved concrete) significantly greater numbers, after one settlement season.The successful recruitment of early colonists is a critical stage in the development of more complex and diverse macrobenthic assemblages, especially those that provide physical habitat structure for other species. Our observations show that this can be facilitated relatively simply for barnacles on marine concrete by manipulating surface heterogeneity at a millimetre scale. Alongside other larger-scale manipulation (e.g. creating holes and pools), including fine-scale habitat heterogeneity in engineering designs can support international efforts to maximise the ecological value of marine urban infrastructure.

Predator avoidance during reproduction: diel movements by spawning sockeye salmon between stream and lake habitats

Daily movements of mobile organisms between habitats in response to changing trade-offs between predation risk and foraging gains are well established; however, less in known about whether similar tactics are used during reproduction, a time period when many organisms are particularly vulnerable to predators. We investigated the reproductive behaviour of adult sockeye salmon (Oncorhynchus nerka) and the activity of their principal predator, brown bears (Ursus arctos), on streams in south-western Alaska. Specifically, we continuously monitored movements of salmon between lake habitat, where salmon are invulnerable to bears, and three small streams, where salmon spawn and are highly vulnerable to bears. We conducted our study across 2 years that offered a distinct contrast in bear activity and predation rates. Diel movements by adult sockeye salmon between stream and lake habitat were observed in 51.3% ± 17.7% (mean ± SD) of individuals among years and sites. Fish that moved tended to hold in the lake for most of the day and then migrated into spawning streams during the night, coincident with when bear activity on streams tended to be lowest. Additionally, cyclic movements between lakes and spawning streams were concentrated earlier in the spawning season. Individuals that exhibited diel movements had longer average reproductive life spans than those who made only one directed movement into a stream. However, the relative effect was dependent on the timing of bear predation, which varied between years. When predation pressure primarily occurred early in the spawning run (i.e., during the height of the diel movements), movers lived 120-310% longer than non-movers. If predation pressure was concentrated later in the spawning run (i.e. when most movements had ceased), movers only lived 10-60% longer. Our results suggest a dynamic trade-off in reproductive strategies of sockeye salmon; adults must be in the stream to reproduce, but must also avoid predation long enough to spawn. Given the interannual variation in the timing and intensity of predation pressure, the advantages of a particular movement strategy will likely vary among years. Regardless, movements by salmon allowed individuals to exploit fine-scale habitat heterogeneity during reproduction, which appears to be a strategy to reduce predation risk on the spawning grounds.

Weak impact of fine‐scale landscape heterogeneity on evolutionary potential in Arabidopsis lyrata

Microhabitat heterogeneity can lead to fine-scale local adaptation when gene flow is restricted, which may be important for the maintenance of genetic variation within populations. This study tested whether microhabitat heterogeneity was associated with trait differences in a population of Arabidopsis lyrata and studied its impact on the genetic variance-covariance (G) matrix. Maternal seed families were collected from dune tops and bottoms, two microhabitats known to vary significantly in water availability. In a common garden experiment, replicate individuals per family were raised under wet and dry conditions, and physiological, morphological and life-history traits were assessed. Plants from the two microenvironments differed in their response to treatment in two performance components, in stomata density and most strongly in flowering time. Under wet conditions, plants originating from dune bottoms flowered 4weeks earlier than those from dune tops. Only one of three G-matrix comparisons revealed that habitat heterogeneity and evolutionary potential were positively linked. The number of independent trait dimensions was larger in the entire population than within subpopulations separated by microhabitat under wet conditions. However, the size of the G-matrix was no larger in the entire population than within subpopulations separated by microhabitat, and trait correlation structure between microhabitats and treatments was not significantly different. These results indicate that fine-scale habitat heterogeneity likely led to local adaptation, which weakly affected levels of across-trait genetic variation.

Cork oak woodlands in the north Tyrrhenian area (Italy): distribution and plant species diversity of a relict forest ecosystem

Italian north Tyrrhenian regions harbour pure or nearly pure cork oak woodlands towards the NE limit of the species range and the highest proportion of the habitat on mainland Italy, but their distribution, biodiversity and conservation status are still poorly known. Focusing on the key region of Tuscany, we analysed distribution and plant species richness of these forests based on literature and original field data. The habitat covers about 5.730 ha and is strongly fragmented for natural and non-natural reasons, underscoring its relict character. Out of the 420 plant taxa recorded, 19 were listed in IUCN categories at the national or regional level. Taxonomic singularity was especially high at the genus rank. Three neighbouring areas with different soil conditions and forest structure were compared based on field surveys, sample plots and linear transects from dense stands to shrub and herbaceous communities of small openings. The Versegge site showed the highest species richness at the area-scale and the mean diversity at the plot scale, representing a key biotope for the conservation of the habitat. However, the geographic–floristic covariation between the three sites suggested that a network of small areas would be more effective than a single, large area to protect the most significant components of the local cork oak flora. The Lattaia and Terzo areas may play an important role as corridor sites between the four SCIs of the Rete Natura 2000 network including cork oak communities on the Tuscan mainland. Species richness was higher in the stands with lower cork oak basal area, and there was a significant increase in species number along transects reaching openings caused by traditional human activities. A minor anthropic disturbance of traditional type may help to preserve the fine-scale habitat heterogeneity and the biodiversity of the whole ecosystem.

Diversity of bird feeding guilds in relation to habitat heterogeneity and land-use cover in a human-modified landscape in southern Mexico

Abstract:Habitat heterogeneity is an important ecological determinant of species richness. We evaluated the diversity within bird feeding guilds as related to habitat heterogeneity and land-use cover in a human-modified tropical landscape. To quantify this process, fine-scale bird census and habitat heterogeneity data were collected for a bird community in a 22.5-km2fragmented landscape in southern Mexico. Land-use cover data derived from field surveys were used to calculate habitat heterogeneity index values and the extent of each land-use cover type in 239 grid cells of 300 × 300 m. Bird diversity values were obtained based on 1195 point-counts in these cells. Product-moment correlations and linear regression analyses were used to evaluate the relationship between bird-guild diversity values and habitat heterogeneity. A total of 109 resident bird species grouped in six feeding guilds were recorded: insectivores (42%), frugivores (21%), granivores (17%), nectarivores (9%), omnivores (8%) and carnivores (3%). Diversity values for the entire bird community were significantly positively related to habitat heterogeneity, but feeding guilds showed contrasting responses to habitat heterogeneity and the amount of land-use cover: insectivores and frugivores were more diverse and abundant in secondary forests than in any other land-cover. Our findings illustrate the importance of small landscape fragments as potential key refuges for the most diverse and specialized feeding guilds, such as granivores and insectivores.

The influence of fine-scale habitat heterogeneity on invertebrate assemblage structure in upland semi-natural grassland

The Scottish uplands consist of a variety of semi-natural habitats of high nature value that are typically managed by extensive livestock grazing. However, as a result of the decoupling of financial subsidies from production in the recent Common Agricultural Policy reform, sheep numbers in the uplands have significantly declined and there is concern that the conservation value of these habitats is at risk. A large-scale experiment that manipulated sheep grazing management provided an excellent opportunity to examine the influence of fine-scale habitat characteristics on invertebrate assemblages. A Geographical Information System was used to extract habitat data at four spatial scales ranging from 0.25 m to 5.0 m in ellipses around pitfall transects. Multivariate analyses were applied to determine the most appropriate spatial scale and the primary habitat characteristics influencing three categories of invertebrates (i.e., mobile arthropods, immobile invertebrates and carabid beetles). Sites that were grazed year-round, and thus more intensively, were associated with smaller mobile arthropods (e.g., money spiders), tipulid larvae and earthworms, while sites that were grazed during summer months only were associated with larger mobile arthropods (e.g., wolf spiders and Carabus beetles) and sawfly and lepidopteran larvae. Grazing regime and the area of fine and broad-leaved grasses were found to influence all three groups of invertebrates at a range of spatial scales, thus indicating the importance of these variables in predicting assemblage structure. The optimum spatial scale at which to collect associated habitat information was from within 0.25 m for immobile invertebrates, and from within 3 m for mobile arthropods and carabids. This supports the hypothesis that immobile invertebrates react to their surrounding habitat at a finer spatial scale than mobile invertebrates and draws into question the effectiveness of collecting habitat information from directly adjacent to the pitfall traps when studying mobile arthropods.

Relationship Between Distance From Major Larval Habitats and Abundance of Adult Mosquitoes in Semiarid Plains Landscapes in Colorado

We examined the relationship between distance from major larval habitats and abundance of adult mosquitoes in the semiarid plains landscape characteristic of eastern Colorado. Mosquito collection was conducted from late June to early August 2007 and included trap locations at distances ranging from <10 m up to 20-150 m and 160-373 m from three major larval habitats: one area along a river corridor and two small reservoirs. The study yielded 65,140 mosquitoes of 14 species, and five species were sufficiently abundant to be included in statistical analyses: Aedes vexans (Meigen), Culex tarsalis Coquillett, Ochlerotatus dorsalis (Meigen) (=Ae. dorsalis), Ochlerotatus melanimon (Dyar) (=Ae. melanimon), and Culex pipiens L. Distance to nearest major larval habitat was not strongly related to Culex abundance within the approximately = 400-m range from larval habitats examined in this study. Abundance of Ae. vexans declined significantly with distance from the larval habitat, whereas abundance was significantly higher in the 20-150- and 160-373-m classes compared with areas within 10 m of the larval habitat for both Ochlerotatus species. Except for Ae. vexans, however, we did not find monotonic increasing or decreasing abundance trends associated with distance from larval habitats for the 400-m range examined. This, combined with a finding that fine-scale habitat heterogeneity influenced abundance for most of the mosquitoes examined, underscores the importance of considering not only distance from larval habitat but also fine-scale habitat heterogeneity to understand how important nuisance-biters and West Nile virus (family Flaviviridae, genus Flavivirus, WNV) vectors use the landscape. We also discuss how these results relate to previous studies from western North America and explore their relevance to operational implementation of adulticides to suppress mosquito vectors during WNV disease outbreaks in the Great Plains.

Influences of wind-wave exposure on the distribution and density of recruit reef fishes at Kure and Pearl and Hermes Atolls, Northwestern Hawaiian Islands

This paper describes the results of a field survey designed to test the prediction that the density of benthic juveniles of shallow-reef fishes is greater on wind-wave “exposed” sectors of a pair of isolated oceanic atolls (Kure, Pearl and Hermes) at the far northwestern end of the Hawaiian Islands, an archipelago in which east-northeasterly trade winds dominate onshore water flow and transport by surface currents. The densities of recruits (juveniles ≤5 cm total length) were higher overall on windward versus leeward sectors of carbonate rock-rubble back reefs at both atolls, and the pattern was stronger for smaller (likely younger, more recently settled) recruits of four of the five most abundant species and the remainder pooled as an “Other” taxon. The windward-leeward disparity was four-fold greater at Pearl Hermes (the atoll with a three-fold longer perimeter) than at Kure. Resident predator biomass also was correlated with recruit densities, but habitat (benthic substratum) effects were generally weak. The distribution and abundance of recruits and juveniles of the primarily endemic reef fishes on shallow back reefs at these atolls appear partly influenced by relative rates of water flow over windward vs. leeward sectors of barrier reef and by the size, shape, and orientation of habitat parcels that filter out postlarval fishes with relatively weak swimming capabilities like labroids. Whole-reef geomorphology as well as fine-scale habitat heterogeneity and rugosity should be considered among the suite of many factors used to interpret observed spatial patterns of post-settlement juvenile fish distribution at atolls and perhaps some other tropical reefs.

Government targets for protected area management: will threatened butterflies benefit?

The UK Government has set targets for biodiversity conservation in England based on several indicators, including the status of protected areas [e.g. Sites of Special Scientific Interest (SSSIs)]. Specifically, the Government aims to achieve favourable condition [defined by Common Standards Monitoring (CSM)] on 95% by area of SSSIs by 2010. SSSIs are important for threatened butterflies and management to achieve favourable condition will play a key role in determining future population levels of these high-profile insects. Because only notified features of SSSIs are considered within CSM, we investigated the level of notification for three threatened butterflies. We found that these species were notified at only 15–33% of SSSIs where they occurred; though most site managers did manage for them under broader site conservation objectives. We investigated the relationship between SSSI condition status and population trend for eight butterfly species of conservation concern to assess the benefit to butterflies of sites attaining favourable condition status. The majority (80%) of population trends on SSSIs in favourable condition were positive, suggesting an overall beneficial impact of SSSI management. However, four of the eight species maintained lower populations at favourable condition SSSIs than at sites in one of the unfavourable condition categories. We suggest that current condition assessment based chiefly on notified vegetation communities lacks the sensitivity to identify the complex habitat conditions for these (mosaic) species. As butterflies are good indictors for a wide range of invertebrates, many species requiring fine-scale habitat heterogeneity may be at risk from the Government’s target.

Coral reef development drives molluscan diversity increase at local and regional scales in the late Neogene and Quaternary of the southwestern Caribbean

The late Neogene was a time of major environmental change in Tropical America. Global cooling and associated oceanographic reorganization and the onset and intensification of glaciation in the Northern Hemisphere during the past ten million years coincided with the uplift of the Central American isthmus and resulting changes in regional oceanographic conditions. Previous analyses of patterns of taxonomic turnover and the shifting abundances of major ecological guilds indicated that the regional shallow-water marine biota responded to these environmental changes through extinction and via a restructuring of local benthic food webs, but it is not clear whether this ecological response had an effect on the diversity of molluscan assemblages in the region. Changes in regional and local diversity are often used as proxies for similar ecological response to environmental change in large-scale paleontological studies, but a clear relationship between diversity and ecological function has rarely been demonstrated in marine systems dominated by mollusks. To explore this relationship, we have compiled a data set of the stratigraphic and environmental distribution of genera of mollusks in large new collections of fossil specimens from the late Neogene and Recent of the southwestern Caribbean. Analysis of a selection of ecological diversity measures indicates that within shelf depths, assemblages from deeper water (51–200 m) were more diverse than shallow-water (&lt;50 m) assemblages in the Pliocene. Lower diversity for shallow-water assemblages is caused by increased dominance of a few superabundant taxa in each assemblage. This implies that studies of diversity of shelf benthos need to control for relatively fine scaled environmental conditions if they are to avoid interpreting artifacts of uneven sampling as true change of diversity. For shallow-water assemblages only, there was significant increase in local and regional diversity of bivalve assemblages after the late Pliocene. No parallel increase in gastropods could be detected, but this likely is because sample size was inadequate for documenting the diversity of gastropod assemblages following a steep post-Pliocene decline of average gastropod abundance. Both the increasing bivalve diversity and the decrease in average abundance of gastropod taxa correspond to an interval of increasing carbonate deposition and reef building in the region, and are likely a result of increased fine-scale habitat heterogeneity controlled by the local distribution of carbonate buildups. Each of these results demonstrates that documenting the ecological response of tropical marine ecosystems to regional environmental change requires a large volume of fine-scaled samples with detailed paleoenvironmental control. Such data sets are rarely available from the fossil record.

Spatial population structure in the banner-tailed kangaroo rat, Dipodomys spectabilis

I attempted to characterize spatial units of local dynamics and dispersal in banner-tailed kangaroo rats (Dipodomys spectabilis), to determine if spatial structure influenced population dynamics in the way predicted by current metapopulation models. D. spectabilis exhibited a hierarchical spatial structure. "Local populations" that appeared as discrete entities on a scale of kilometers were subdivided into clusters of mounds on a scale of meters. This structure, however, cannot be characyerized in terms of the discrete habitat patches envisioned by the metapopulation models. Occupied areas were statistically distinguishable from the surrounding matrix, but this difference was only quantitative. There were no discrete boundaries between occupied areas and the matrix. Habitat within occupied areas was heterogeneous, and occupied areas in different locations were statistically distinguishable from each other. High heterogeneity within occupied areas, and high contrast among them, make it difficult to define what is a suitable habitat patch for D. spectabilis. On a smaller spatial scale, there was significant aggregation of resident mounds within occupied areas. These aggregations, however, do not correspond to discrete habitat patches. Rather, they appear to result from an interaction between fine-scale habitat heterogeneity and limited dispersal due to natal philopatry and low adult vagility. These complications make it difficult to identify habitat patches independent of the species' distribution. For species like D. spectabilis that are patchily distributed but do not occupy discrete habitat patches, a patch occupancy approach does not seem appropriate for describing spatial structure. Hierarchical spatial structure underscores the need for a framework that incorporates multiple scales of spatial structure, rather than one that pre-imposes a single spatial scale as being important for population dynamics. A framework that (i) considers patchiness as a combination of both habitat heterogeneity, and life-history and behavioral characteristics, and (ii) incorporates hierarchical spatial structure, appears to be the most suitable for conceptualizing spatial dynamics of behaviorally complex vertebrates such as D. spectabilis.