The area-independent effects of habitat complexity on biodiversity vary between regions.

Potential explanatory variables often co-vary in studies of species richness. Where topography varies within a survey it is difficult to separate area and habitat-diversity effects. Topographically complex surfaces may contain more species due to increased habitat diversity or as a result of increased area per se. Fractal geometry can be used to adjust species richness estimates to control for increases in area on complex surfaces. Application of fractal techniques to a survey of rocky shores demonstrated an unambiguous area-independent effect of topography on species richness in the Isle of Man. In contrast, variation in species richness in south-west England reflected surface availability alone. Multivariate tests and variation in limpet abundances also demonstrated regional variation in the area-independent effects of topography. Community composition did not vary with increasing surface complexity in south-west England. These results suggest large-scale gradients in the effects of heterogeneity on community processes or demography.

Functional responses of the intertidal amphipod Echinogammarus marinus: effects of prey supply, model selection and habitat complexity.

The influence of predation in structuring ecological communities can be informed by examining the shape and magnitude of the functional response of predators towards prey. We derived functional responses of the ubiquitous intertidal amphipod Echinogammarus marinus towards one of its preferred prey species, the isopod Jaera nordmanni. First, we examined the form of the functional response where prey were replaced following consumption, as compared to the usual experimental design where prey density in each replicate is allowed to deplete. E. marinus exhibited Type II functional responses, i.e. inversely density-dependent predation of J. nordmanni that increased linearly with prey availability at low densities, but decreased with further prey supply. In both prey replacement and non-replacement experiments, handling times and maximum feeding rates were similar. The non-replacement design underestimated attack rates compared to when prey were replaced. We then compared the use of Holling’s disc equation (assuming constant prey density) with the more appropriate Rogers’ random predator equation (accounting for prey depletion) using the prey non-replacement data. Rogers’ equation returned significantly greater attack rates but lower maximum feeding rates, indicating that model choice has significant implications for parameter estimates. We then manipulated habitat complexity and found significantly reduced predation by the amphipod in complex as opposed to simple habitat structure. Further, the functional response changed from a Type II in simple habitats to a sigmoidal, density-dependent Type III response in complex habitats, which may impart stability on the predator−prey interaction. Enhanced habitat complexity returned significantly lower attack rates, higher handling times and lower maximum feeding rates. These findings illustrate the sensitivity of the functional response to variations in prey supply, model selection and habitat complexity and, further, that E. marinus could potentially determine the local exclusion and persistence of prey through habitat-mediated changes in its predatory functional responses.

Natural and artificially-induced habitat complexity and freshwater fish species composition

The effects of habitat complexity, induced by construction of artificial reefs, on the fish assemblages in the Barra Bonita reservoir, Brazil, and in the lotic zone immediately below the dam were studied. Four artificial reefs were constructed in each habitat at variable distances from the shore. Multiple correspondence analysis showed that the factors distance from the shore and type of habitat were determinants for the group formation, and artificial reefs had a lesser effect. Fish species composition was about the same at locations with and without reefs.

The effects of habitat complexity and hydraulic conditions on the establishment of benthic stream macroalgae

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Application of computer-aided tomography techniques to visualize kelp holdfast structure reveals the importance of habitat complexity for supporting marine biodiversity

Ecosystem engineers that increase habitat complexity are keystone species in marine systems, increasing shelter and niche availability, and therefore biodiversity. For example, kelp holdfasts form intricate structures and host the largest number of organisms in kelp ecosystems. However, methods that quantify 3D habitat complexity have only seldom been used in marine habitats, and never in kelp holdfast communities. This study investigated the role of kelp holdfasts (Laminaria hyperborea) in supporting benthic faunal biodiversity. Computer-aided tomography (CT-) scanning was used to quantify the three-dimensional geometrical complexity of holdfasts, including volume, surface area and surface fractal dimension (FD). Additionally, the number of haptera, number of haptera per unit of volume, and age of kelps were estimated. These measurements were compared to faunal biodiversity and community structure, using partial least-squares regression and multivariate ordination. Holdfast volume explained most of the variance observed in biodiversity indices, however all other complexity measures also strongly contributed to the variance observed. Multivariate ordinations further revealed that surface area and haptera per unit of volume accounted for the patterns observed in faunal community structure. Using 3D image analysis, this study makes a strong contribution to elucidate quantitative mechanisms underlying the observed relationship between biodiversity and habitat complexity. Furthermore, the potential of CT-scanning as an ecological tool is demonstrated, and a methodology for its use in future similar studies is established. Such spatially resolved imager analysis could help identify structurally complex areas as biodiversity hotspots, and may support the prioritization of areas for conservation.

Application of computer-aided tomography techniques to visualize kelp holdfast structure reveals the importance of habitat complexity for supporting marine biodiversity

Ecosystem engineers that increase habitat complexity are keystone species in marine systems, increasing shelter and niche availability, and therefore biodiversity. For example, kelp holdfasts form intricate structures and host the largest number of organisms in kelp ecosystems. However, methods that quantify 3D habitat complexity have only seldom been used in marine habitats, and never in kelp holdfast communities. This study investigated the role of kelp holdfasts (Laminaria hyperborea) in supporting benthic faunal biodiversity. Computer-aided tomography (CT-) scanning was used to quantify the three-dimensional geometrical complexity of holdfasts, including volume, surface area and surface fractal dimension (FD). Additionally, the number of haptera, number of haptera per unit of volume, and age of kelps were estimated. These measurements were compared to faunal biodiversity and community structure, using partial least-squares regression and multivariate ordination. Holdfast volume explained most of the variance observed in biodiversity indices, however all other complexity measures also strongly contributed to the variance observed. Multivariate ordinations further revealed that surface area and haptera per unit of volume accounted for the patterns observed in faunal community structure. Using 3D image analysis, this study makes a strong contribution to elucidate quantitative mechanisms underlying the observed relationship between biodiversity and habitat complexity. Furthermore, the potential of CT-scanning as an ecological tool is demonstrated, and a methodology for its use in future similar studies is established. Such spatially resolved imager analysis could help identify structurally complex areas as biodiversity hotspots, and may support the prioritization of areas for conservation.

The comparative effects of habitat complexity on faunal assemblages of northern temperate artificial and natural reefs

Several north temperate marine species were recorded on subtidal hard-substratum reef sites selected to produce a gradient of structural complexity. The study employed an established scuba-based census method, the belt transect. The three types of reef examined, with a measured gradient of increasing structural complexity, were natural rocky reef, artificial reef constructed of solid concrete blocks, and artificial reef made of concrete blocks with voids. Surveys were undertaken monthly over a calendar year using randomly placed fixed rope transects. For a number of conspicuous species of fish and invertebrates, significant differences were found between the levels of habitat complexity and abundance. Overall abundance for many of the species examined was 2-3 times higher on the complex artificial habitats than on simple artificial or natural reef habitats. The enhanced habitat availability produced by the increased structural complexity delivered through specifically designed artificial reefs may have the potential to augment faunal abundance while promoting species diversity. 

Fish Habitat Utilization Patterns and Evaluation of the Efficacy of Marine Protected Areas in Hawaii: Integration of NOAA Digital Benthic Habitat Mapping and Coral Reef Ecological Studies

Over the past four decades, the state of Hawaii has developed a system of eleven Marine Life Conservation Districts (MLCDs) to conserve and replenish marine resources around the state. Initially established to provide opportunities for public interaction with the marine environment, these MLCDs vary in size, habitat quality, and management regimes, providing an excellent opportunity to test hypotheses concerning marine protected area (MPA) design and function using multiple discreet sampling units. NOAA/NOS/NCCOS/Center for Coastal Monitoring and Assessment’s Biogeography Team developed digital benthic habitat maps for all MLCD and adjacent habitats. These maps were used to evaluate the efficacy of existing MLCDs for biodiversity conservation and fisheries replenishment, using a spatially explicit stratified random sampling design. Coupling the distribution of habitats and species habitat affinities using GIS technology elucidates species habitat utilization patterns at scales that are commensurate with ecosystem processes and is useful in defining essential fish habitat and biologically relevant boundaries for MPAs. Analysis of benthic cover validated the a priori classification of habitat types and provided justification for using these habitat strata to conduct stratified random sampling and analyses of fish habitat utilization patterns. Results showed that the abundance and distribution of species and assemblages exhibited strong correlations with habitat types. Fish assemblages in the colonized and uncolonized hardbottom habitats were found to be most similar among all of the habitat types. Much of the macroalgae habitat sampled was macroalgae growing on hard substrate, and as a result showed similarities with the other hardbottom assemblages. The fish assemblages in the sand habitats were highly variable but distinct from the other habitat types. Management regime also played an important role in the abundance and distribution of fish assemblages. MLCDs had higher values for most fish assemblage characteristics (e.g. biomass, size, diversity) compared with adjacent fished areas and Fisheries Management Areas (FMAs) across all habitat types. In addition, apex predators and other targeted resources species were more abundant and larger in the MLCDs, illustrating the effectiveness of these closures in conserving fish populations. Habitat complexity, quality, size and level of protection from fishing were important determinates of MLCD effectiveness with respect to their associated fish assemblages. (PDF contains 217 pages)

Importance of shoreface sand ridges as habitat for fishes off the northeast coast of the United States

To determine if shoreface sand ridges provide unique habitats for fish on the inner continental shelf, two cross-shelf trawl surveys (23 km in length) were conducted in southern New Jersey (July and September 1991−95 with a beam trawl and July and September 1997−06 with an otter trawl) to assess whether species abundance, richness, and assemblages differed on and away from the ridge. The dominant species collected with both gears were from the families Paralichthyidae, Triglidae, Gobiidae, Serranidae, Engraulidae, Stromateidae, and Sciaenidae. Overall abundance (n=41,451 individuals) and species richness (n=61 species) were distributed bimodally across the nearshore to offshore transect, and the highest values were found on either side of the sand ridge regardless of gear type. Canonical correspondence analysis revealed three species assemblages: inshore (<5 meters depth), near-ridge (9−14 meters depth), and offshore (>14 meters depth), and variation in species composition between gear types. Environmental factors that corresponded with the assemblage changes included depth, temperature, distance from the top of the ridge, and habitat complexity. The most abundant near-ridge assemblages were distinct and included economically important species. Sand ridges of the inner continental shelf appear to be important habitat for a number of fish species and therefore may not be a suitable area for sand and gravel mining.

Multiscale habitat associations of deepwater demersal fishes off central California

Fish-habitat associations were examined at three spatial scales in Monterey Bay, California, to determine how benthic habitats and landscape configuration have structured deepwater demersal fish assemblages. Fish counts and habitat variables were quantified by using observer and video data collected from a submersible. Fish responded to benthic habitats at scales ranging from cm’s to km’s. At broad-scales (km’s), habitat strata classified from acoustic maps were a strong predictor of fish assemblage composition. At intermediate-scales (m’s−100 m’s), fish species were associated with specific substratum patch types. At fine-scales (<1 m), microhabitat associations revealed differing degrees of microhabitat specificity, and for some species revealed niche separation within patches. The use of habitat characteristics in ecosystembased management, particularly as a surrogate for species distributions, will depend on resolving fish-habitat associations and habitat complexity over multiple scales.

Habitat simplification increases the impact of a freshwater invasive fish

Biodiversity continues to decline at a range of spatial scales and there is an urgent requirement to understand how multiple drivers interact in causing such declines. Further, we require methodologies that can facilitate predictions of the effects of such drivers in the future. Habitat degradation and biological invasions are two of the most important threats to biodiversity and here we investigate their combined effects, both in terms of understanding and predicting impacts on native species. The predatory largemouth bass Micropterus salmoides is one of the World’s Worst Invaders, causing declines in native prey species, and its introduction often coincides with habitat simplification. We investigated the predatory functional response, as a measure of ecological impact, of juvenile largemouth bass in artificial vegetation over a range of habitat complexities (high, intermediate, low and zero). Prey, the female guppy Poecilia reticulata, were representative of native fish. As habitats became less complex, significantly more prey were consumed, since, even although attack rates declined, reduced handling times resulted in higher maximum feeding rates by bass. At all levels of habitat complexity, bass exhibited potentially population destabilising Type II functional responses, with no emergence of more stabilising Type III functional responses as often occurs in predator-prey relationships in complex habitats. Thus, habitat degradation and simplification potentially exacerbate the impact of this invasive species, but even highly complex habitats may ultimately not protect native species. The utilisation of functional responses under varying environmental contexts provides a method for the understanding and prediction of invasive species impacts.

Spatial prediction of rufous bristlebird habitat in a coastal heathland: a GIS-based approach

1. To develop a conservation management plan for a species, knowledge of its distribution and spatial arrangement of preferred habitat is essential. This is a difficult task, especially when the species of concern is in low   abundance. In south-western Victoria, Australia, populations of the rare rufous bristlebird Dasyornis broadbenti are threatened by fragmentation of suitable habitat. In order to improve the conservation status of this species, critical habitat requirements must be identified and a system of corridors must be established to link known populations. A predictive spatial model of rufous bristlebird habitat was developed in order to identify critical areas requiring preservation, such as corridors for dispersal.
2. Habitat models generated using generalized linear modelling techniques can assist in delineating the specific habitat requirements of a species. Coupled with geographic information system (GIS) technology, these models can be extrapolated to produce maps displaying the spatial configuration of suitable habitat.
3. Models were generated using logistic regression, with bristlebird presence or absence as the dependent variable and landscape variables, extracted from both GIS data layers and multispectral digital imagery, as the predictors. A multimodel inference approach based on Akaike’s information criterion was used and the resulting model was applied in a GIS to extrapolate predicted likelihood of occurrence across the entire area of concern. The predictive performance of the selected model was evaluated using the receiver operating characteristic (ROC) technique. A hierarchical partitioning protocol was used to identify the predictor variables most likely to influence variation in the dependent variable. Probability of species presence was used as an index of habitat suitability.
4. Negative associations between rufous bristlebird presence and  increasing elevation, 'distance to cree', 'distance to coast' and sun index were evident, suggesting a preference for areas relatively low in altitude, in close proximity to the coastal fringe and drainage lines, and receiving less direct sunlight. A positive association with increasing habitat complexity also suggested that this species prefers areas containing high vertical density of vegetation.
5. The predictive performance of the selected model was shown to be high (area under the curve 0·97), indicating a good fit of the model to the data. Hierarchical partitioning analysis showed that all the variables considered had significant  independent contributions towards explaining the variation in the dependent variable. The proportion of the total study area that was predicted as suitable habitat for the rufous bristlebird (using probability of occurrence at a ≥0·5 level ) was 16%.
6. Synthesis and applications. The spatial model clearly delineated areas predicted as highly suitable rufous bristlebird habitat, with evidence of potential corridors linking coastal and inland populations via gullies. Conservation of this species will depend on management actions that protect the critical habitats identified in the model. A multi-scale  approach to the modelling process is recommended whereby a spatially explicit model is first generated using landscape variables extracted from a GIS, and a second model at site level is developed using fine-scale habitat variables measured on the ground. Where there are constraints on the time and cost involved in measuring finer scale variables, the first step alone can be used for conservation planning.

Modelling habitat suitability of the swamp antechinus (Antechinus minimus maritimus) in the coastal heathlands of southern Victoria, Australia

In recent years, predictive habitat distribution models, derived by combining multivariate statistical analyses with Geographic Information System (GIS) technology, have been recognised for their utility in conservation planning. The size and spatial arrangement of suitable habitat can influence the long-term persistence of some faunal species. In southwestern Victoria, Australia, populations of the rare swamp antechinus (Antechinus minimus maritimus) are threatened by further fragmentation of suitable habitat. In the current study, a spatially explicit habitat suitability model was developed for A. minimus that incorporated a measure of vegetation structure. Models were generated using logistic regression with species presence or absence as the dependent variable and landscape variables, extracted from both GIS data layers and multi-spectral digital imagery, as the predictors. The most parsimonious model, based on the Akaike Information Criterion, was spatially extrapolated in the GIS. Probability of species presence was used as an index of habitat suitability. A negative association between A. minimus presence and both elevation and habitat complexity was evidenced, suggesting a preference for relatively low altitudes and a vegetation structure of low vertical complexity. The predictive performance of the selected model was shown to be high (91%), indicating a good fit of the model to the data. The proportion of the study area predicted as suitable habitat for A. minimus (Probability of occurrence greater-or-equal, slanted0.5) was 11.7%. Habitat suitability maps not only provide baseline information about the spatial arrangement of potentially suitable habitat for a species, but they also help to refine the search for other populations, making them an important conservation tool.

Methodological and conceptual issues in the search for a relationship between animal body-size distributions and benthic habitat architecture

Benthic ecologists have studied the distribution of animal body sizes because it is a form of ‘taxon-free’ classification that may be a useful metric for describing variation within and between ecological communities. In particular, the idea that the allometry of physiological and life-history traits may control species composition and relative abundances implies a functional link between body-size distributions and communities. The physical structure of aquatic habitats has often been cited as the mechanism by which habitat may determine body-size distributions in communities. However, further progress is hindered by a lack of theoretical clarity regarding the mechanisms that connect body size to the characteristics of ecological communities, leading to methods that may obscure interesting trends in body-size data. This review examines the methodological and conceptual issues hindering progress in the search for a relationship between animal body size and habitat architecture and suggests ways to resolve these issues. Problems are identified with current methods for the measurement of animal body size, the data and measures used to quantify body-size distributions and the methods used to identify patterns therein. Fundamentally, renewed emphasis on the mechanisms by which animal body sizes are influenced by habitat architecture is required to refine methodology and synthesise results from pattern-seeking and mechanistic studies.