Multi-scale approach to species–habitat relationships: juvenile fish in a large river section

1. Juvenile fish were sampled by electrofishing using the point abundance sampling method from August to November 1991 in the Lower Rhône River. Variations in fish abundance between point samples were partitioned according to three spatial scales of habitat heterogeneity.
2. The compartment scale, differentiating channel and 'dike fields' (an area marginal to the main channel partially enclosed by an artificial embankment), was associated with variation in water current, temperature and biological productivity. Every juvenile fish species studied was more abundant in the dike fields, using them as a nursery.
3. Within both compartments (i.e. main channel and dike fields), the same five bank types were represented, defining the intermediate scale of habitat heterogeneity. In the dike fields, beaches were selected by most of the species, whereas steep silt banks were more often selected in the main channel.
4. After having removed the variation explained at the two larger habitat scales, only depth and shelter were found to have a significant effect on fish abundance at the microhabitat scale.
5. When compared with a classical single-scale approach, the multi-scale approach was found to explain a greater amount of fish abundance variations and to facilitate the interpretation of observed patterns by scaling the species' responses to habitat heterogeneity. The difference between the amounts of variation explained was proposed as a measure of how much habitat selection is hierarchical.     

Influence of habitat heterogeneity on the distribution of larval Pacific lamprey (Lampetra tridentata) at two spatial scales

1. Spatial patterns in channel morphology and substratum composition at small (1–10 metres) and large scales (1–10 kilometres) were analysed to determine the influence of habitat heterogeneity on the distribution and abundance of larval lamprey. 2. We used a nested sampling design and multiple logistic regression to evaluate spatial heterogeneity in the abundance of larval Pacific lamprey, Lampetra tridentata, and habitat in 30 sites (each composed of twelve 1‐m² quadrat samples) distributed throughout a 55‐km section of the Middle Fork John Day River, OR, U.SA. Statistical models predicting the relative abundance of larvae both among sites (large scale) and among samples (small scale) were ranked using Akaike's Information Criterion (AIC) to identify the ‘best approximating’ models from a set of a priori candidate models determined from the literature on larval lamprey habitat associations. 3. Stream habitat variables predicted patterns in larval abundance but played different roles at different spatial scales. The abundance of larvae at large scales was positively associated with water depth and open riparian canopy, whereas patchiness in larval occurrence at small scales was associated with low water velocity, channel‐unit morphology (pool habitats), and the availability of habitat suitable for burrowing. 4. Habitat variables explained variation in larval abundance at large and small scales, but locational factors, such as longitudinal position (river km) and sample location within the channel unit, explained additional variation in the logistic regression model. The results emphasise the need for spatially explicit analysis, both in examining fish habitat relationships and in developing conservation plans for declining fish populations.     

Effects of longitudinal variations in stream habitat structure on fish abundance: an analysis based on subunit-scale habitat classification

SUMMARY 1. Stream reaches contain assortments of various habitat types that can be defined at different spatial scales, such as channel unit (e.g. pools, riffles) and subunit (patches within channel units). We described longitudinal (upstream–downstream) patterns of stream habitat structure by considering subunits as structural elements, and examined their effects on the abundance of masu salmon ( Oncorhynchus masou ) and rosyface dace ( Leuciscus ezoe ) in a third-order tributary of the Teshio River in northern Hokkaido, Japan.
2. Nine subunit types were determined on the basis of water depth, current velocity and substrate, using 0.5 × 0.5 m grids. Although both masu salmon and rosyface dace used pools as a major habitat, the former preferred a subunit type occurring at pool heads (PH subunit) while the latter preferred a slow-current edge type (SE-2 subunit).
3. Along the course of the stream, slow-edge subunits (SE-1, 2 and 3) increased in frequency downstream while fast-edge subunits (FE-1 and 2) decreased, suggesting a downstream development of slow-current edges. Regression analyses indicated that longitudinal variation in masu salmon abundance was explained by the area of PH, rather than pools. Masu salmon density increased with the area of PH. Rosyface dace abundance was explained by a combination of water depth and the area of SE-2, both effects being positive.
4. Longitudinal variations in the abundance of both species were related to the abundance of their preferred habitat at the subunit scale, rather than channel-unit scale. The results emphasise the importance of fine-scale patchiness when examining stream fish habitats.     

Habitat and climate heterogeneity maintain beta-diversity of birds among landscapes within ecoregions

Aim  Habitat and climate heterogeneity may affect patterns of species diversity from the relatively local scale of communities to the broad biogeographical scale of continents. However, the effects of heterogeneity on species diversity have not been studied as widely at intermediate scales although differences among landscapes in local climate and habitat should maintain beta-diversity.
Location  Bailey ecoregions in the USA.
Methods  Using a geographically extensive dataset on bird distribution and abundance in 35 ecoregions, we tested for the effects of habitat and climate heterogeneity on beta-diversity at two discrete spatial scales: among sample points within landscapes, and among landscapes within ecoregions.
Results  Landscape-level beta-diversity typically accounted for 50% or more of gamma-diversity and was significantly and positively related to habitat heterogeneity (elevational range within an ecoregion) and climate heterogeneity (variation in potential evapotranspiration). Contrary to predictions, point-level beta-diversity was negatively related to habitat and climate heterogeneity, perhaps because heterogeneity constrains alpha-diversity at the landscape level. The geographical spatial separation of landscapes within an ecoregion did not significantly affect beta-diversity at either scale.
Main conclusions  Our results suggest that habitat selection and adaptation to local climate may be the primary processes structuring bird diversity among landscapes within ecoregions, and that dispersal limitation has a lesser role in influencing beta-diversity among landscapes.     

The relationship between multi‐scale habitat heterogeneity and farmland bird abundance

It has recently been proposed that losses in farmland habitat heterogeneity may have been a primary driver of the profound declines exhibited by many farmland bird species in recent decades. However, it has yet to be demonstrated which facets of heterogeneity and what spatial scales are most important for birds. Here we analyse the relationship between abundance and features of landscape heterogeneity at three spatial scales (1, 9 and 25 km²) for 32 bird species commonly associated with farmland. Heterogeneity was quantified using three contrasting indices reflecting 1) the spatial mixing of land uses, 2) variation in field sizes and 3) the density of field boundaries. The spatial mixing of land‐uses explained, on average, the most variation in, and was most likely to be positively associated with, abundance at all spatial scales. The majority of species (66–75%, depending on the spatial scale) were more common in heterogeneous landscapes overall; however, migrants, those under a high level of conservation concern and farmland specialist species tended to be less abundant in more heterogeneous landscapes at all scales. Ground‐nesting species were also more likely to be found in more homogeneous habitats than non‐ground‐nesters, but only at the finest spatial scale. Relationships between abundance and heterogeneity were generally consistent across spatial scales; however, species of high conservation concern had more variable associations compared with other species. These results highlight a potential role for farmland habitat heterogeneity in determining the abundance of many farmland species but suggest that population responses to an increase in heterogeneity would not be unanimously positive and would probably have negative impacts on some species, notably those that are already threatened.     

Assessing the influence of environmental heterogeneity on bird spacing patterns: a case study with two raptors

Testing for aggregation or regularity in point patterns is difficult in the presence of spatial variation in abundance due to environmental heterogeneity. Using a recently developed method generalizing Ripley's K function for non homogeneous point patterns, we test the aggregation of the nests in two species of birds (little owl and Montagu's harrier) exhibiting heterogeneous distributions in response to landscape structure. We compare the results obtained under different null models accounting for environmental heterogeneity at large and/or small spatial scales.
Whereas both species were initially found to form clusters at some scale, taking spatial heterogeneity into account revealed that 1) territorial little owls showed no clustering of territories when habitat availability was considered; 2) semi-colonial harriers still formed significant clusters, but part of the aggregation in this species could be explained by landscape structure alone. Our results highlight that it is feasible and highly recommended to account for non-stationarity when testing for aggregation. Further, provided that sufficient knowledge of the study system is available, this approach helps to identify behavioural and environmental components of spatial variation in abundance. Additionally, we demonstrate that accounting for large or small-scale heterogeneity affects the perception of spacing behaviours differently, so that both need to be considered.     

Local and ecoregion effects on fish assemblage structure in tributaries of the Rio Paraíba do Sul, Brazil

1.  We examined the effects of physical and chemical habitat variables and ecoregions on species occurrence and fish assemblage structure in streams of the Paraíba do Sul basin, in southeast Brazil.
2.  Fish and environmental data were collected from 42 sites on 26 first to fourth order streams (1 : 50 000 map scale) in three ecoregions. The sites occurred in one valley and two plateau ecoregions at altitudes of 40–1080 m and distances of 0.1–188 km from the main channel of the Rio Paraíba do Sul. Physical habitat (substratum, riparian cover, habitat types) and water quality (dissolved oxygen, pH, temperature and conductivity) variables were measured at each sampling site.
3.  A total of 2684 individuals in 16 families and 59 species were recorded.
4.  Ecoregion was a better predictor of the fish assemblage than the other environmental variables, according to the differences between the mean within-class and mean between-class similarities in assemblage data.
5.  Differing landscape characteristics were associated with differing local variables and thereby with differing fish assemblage structures. Riffles, shrub, grass, dissolved oxygen, conductivity and temperature were closely related to fish assemblage structure.
6.  Fish assemblages in sites far from the main river and at higher altitudes also differed from those near the Paraíba do Sul main channel, presumably as a result of differences in connectivity, covarying environmental factors and anthropogenic influence.
7.  These results reinforce the importance of understanding how stream communities are influenced by processes and patterns operating at local and regional scales, which will aid water resource managers to target those factors in their management and rehabilitation efforts.     

Fish-habitat associations across multiple spatial scales

Important to the study of reef fish ecology is understanding the degree to which fish community structure varies across space, what factors can account for such variation, and whether these factors are scale dependent. This study examined the structure of reef fish communities across four spatial scales (1, 10 100, and 200 m²) visually censused from seven sites within Tague Bay, St. Croix, U.S. Virgin Islands. Relative differences in the number of individuals and species among sites indicated a pattern that was consistent across spatial scales. Spearmans rank correlation revealed significant positive correlation in site rankings, in terms of species richness, between 1 and 10 m², and 100 and 200 m²; and for the abundance of individuals between 100 and 200 m². In order to understand the degree to which quantified habitat variables account for patterns in the abundance of individuals and species, and whether these fish-habitat relationships were consistent regardless of spatial scale, separate canonical correlation analyses were conducted at each scale. Independent of scale, the total number of individuals and species were correlated with specific habitat variables, either negatively (with areas of pavement, sand, no algae, and low structural complexity) or positively (with areas of Amphiroa rigida, Halimeda incrassata, high structural complexity, and diverse algae/seagrass communities). These habitat variables explained 31–81% (at scales of 1–200 m²) of the variation in the number of individuals and species. Similar analyses were also performed on the abundances of the nine most common species, and whether their specific habitat associations were independent of scale. Results indicated that habitat variables explained 19–73% (at scales of 1–200 m²) of the variation in abundances of each species. Unique fish-habitat relationships were observed for each species, and most such relationships were consistent across spatial scales. The structure of reef fish communities of Tague Bay was explained in large part by the composition of coral and algae communities present. Both the spatial variation in community structure and the fish-habitat relationships, at the community and population level, appeared to be largely independent of the spatial scale examined. This suggests that generalizations across Tague Bay are possible. Similar habitat associations reported in the literature are discussed with regard to the possibility for generalizations across regions.     

Hierarchy theory reveals multiscale predictors of Arkansas darter (Etheostoma cragini) abundance in a Great Plains riverscape

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Spatio-temporal distribution of young fish in tributaries of natural and flow-regulated sections of a neotropical river in French Guiana

1. We investigated which environmental parameters control the variation in density, in space and time, of young stages of fish in tributaries of a natural and a flow-regulated section of the Sinnamary River, French Guiana.
2. The density of the progeny in most taxa varied in space and/or time. However, most non-Perciformes responded differently to space and/or time in the two sections.
3. Oxygen, turbidity and habitat structure (i.e. bank length, occurrence of undercut bank, richness in litter, vegetation and substratum) were important, as was the position of the sampling site relatively to the main channel in the downstream tributaries, in explaining the variation of density in space in both sections. Both habitat complexity and distance from the main channel protect young fish against unpredictable flow releases downstream from the Petit Saut dam.
4. Hydrological events played an important role in the temporal variation in densities of many fish taxa. The density of most early life and many juvenile stages (mostly Characiformes) was positively related to hydrological events.
5. Some fish taxa had reproductive habits which were relatively independent of abiotic factors, such as flow variability, and the density of their progeny did not vary with time.
6. The nursery areas of more than 45% of species in the Sinnamary River have been degraded by flow regulation.     

Biogeography,ecoregions, and geomorphology affect fish species composition in streams of eastern Oklahoma,USA

Stream fish assemblages are structured by biogeographical, physical and biological factors acting on different spatial scales. We determined how physical factors, geomorphology and stream habitat, influenced fish species composition (presence–absence) in eastern Oklahoma, USA relative to the ecoregion and biogeographic effects previously reported. We sampled fish assemblages and surveyed geomorphology and habitat at 107 stream sites in the Boston Mountains, Ouachita Mountains, and Ozark Highlands ecoregions in eastern Oklahoma. Partial canonical correspondence analyses (pCCAs) and variance partitioning showed that patterns of endemism related to drainage basins and ecoregions explained important variation in fish species composition in all streams, but stream size and local channel morphology explained more variation overall. Stream size effects were most important in explaining variability in fish species composition in both northeastern and southeastern Oklahoma streams. Local channel morphology and substrate characteristics were secondarily important. Variables typically considered important as fish habitat (aquatic vegetation, etc.) had little effect on fish species composition.     

Small-scale recruitment variation in a temperate fish: the roles of macrophytes and food supply

Synopsis The availability of reef-related resources, particularly food and shelter can play a significant role in determining the distribution and abundance of reef fishes. Much of the structure on temperate reefs is provided by macroalgae, and variability in the density of temperate reef fishes at large spatial scales (100's of meters) can often be explained by variation in macroalgal cover or density. In this study I investigated the role of macrophytes and associated food resources on the recruitment of a temperate fish, Tautogolabrus adspersus, at a small spatial scale (0.25 m²). No relationship between the density of new recruits and the percent cover of kelp, foliose or filamentous algae was observed. Multiple regressions revealed that less than 8% of variability in recruitment could be explained by variability in macroalgal cover. However, recruits were found in higher abundance in patches containing many functional forms of seaweeds than in patches dominated by a single form. A wide variety of prey were available for use by cunner recruits; however, crustaceans and mussels were the only common components of their diet, and crustaceans were clearly the most preferred prey. The prey composition in patches where fish were present was compared to randomly selected patches. Significantly greater numbers of isopods, amphipods and newly settled mussels were present in patches where fish were present than in randomly selected patches. The data presented in this study contradict previous work that has shown algal structure to be important in determining patterns of abundance and food supply to be of little significance. A conceptual model is proposed suggesting that settling fish select habitats in a hierarchical manner largely based on their dispersal tendencies. Hierarchical selection of habitats results in different attributes of the habitat being selected during different life-history intervals.     

Spatial variability of the soil seed bank in a heterogeneous ephemeral wetland system in semi-arid Australia

Soil seed banks are an important component of plant community diversity in ephemeral wetlands, allowing many species to persist through unpredictable periods of flood and drought. Spatial variation of extant vegetation in such habitats commonly reflects patterns of flood history and often varies predictably between broadly differing hydro-geomorphic habitat types. Here we investigate whether spatial variation of soil seed banks is similarly controlled by fluvial processes at this scale. Results are presented from a seedling emergence trial using samples collected from a range of habitat types, and at different scales within these, in the ephemeral Narran Lakes system in semi-arid Australia. Composition and structure of soil seed banks varied significantly between habitat types reflecting broad differences in flood frequency. As predicted, germinable seed abundance was found to be highest in intermediately flooded habitats. Variability in soil seed bank composition at a local scale was also found to be influenced by hydrology with greater spatial heterogeneity evident in the river channel as well as amongst the least frequently inundated riparian and floodplain habitats.     

Local and regional determinants of stream fish assemblage structure: inferences based on taxonomic vs. functional groups

Aim  To examine the roles of local and regional environmental variables and biotic interactions in determining the structure of local stream fish assemblages, and to compare results derived from analyses based on taxonomic and functional groups.
Location  Texas, USA.
Methods  Species abundance data were compiled for 157 stream fish assemblages in several river basins across Texas. Species were condensed into functional groups based on trophic and life-history characteristics. Local and regional environmental variables were either measured at each location or determined from scale maps and public-access data bases. The original taxonomic and functional group data sets were analysed using similarity indices, null models of co-occurrence, and direct and indirect ordination techniques. Results derived from taxonomic and functional group data sets are compared.
Results  Inferences regarding the relative roles of local and larger-scale factors in determining stream fish assemblage structure differ dramatically between analyses of taxonomic and functional groups. Taxonomic analyses suggest a prominent role of regional-scale environmental factors, and local assemblages sorted according to a biogeographic pattern. Functional group analyses suggest almost equal roles of factors representative of local and larger scales, and assemblages were distinguished by a habitat template irrespective of geographic region.
Main conclusions  The structure of local stream fish assemblages is determined ultimately by factors representing multiple scales, with the relative importance of each depending on the biological unit employed (species or functional groups). We suggest that analyses using functional groups can more directly infer ecological responses to environmental variation, and therefore may provide a more fruitful avenue for developing and testing ecological theory of community organization across biogeographic scales.     

Consequences for a host–parasitoid interaction of host-plant aggregation, isolation, and phenology

Abstract.  1. Spatial habitat structure can influence the likelihood of patch colonisation by dispersing individuals, and this likelihood may differ according to trophic position, potentially leading to a refuge from parasitism for hosts.
2. Whether habitat patch size, isolation, and host-plant heterogeneity differentially affected host and parasitoid abundance, and parasitism rates was tested using a tri-trophic thistle–herbivore–parasitoid system.
3.  Cirsium palustre thistles ( n = 240) were transplanted in 24 blocks replicated in two sites, creating a range of habitat patch sizes at increasing distance from a pre-existing source population. Plant architecture and phenological stage were measured for each plant and the numbers of the herbivore Tephritis conura and parasitoid Pteromalus elevatus recorded.
4. Mean herbivore numbers per plant increased with host-plant density per patch, but parasitoid numbers and parasitism rates were unaffected. Patch distance from the source population did not influence insect abundance or parasitism rates. Parasitoid abundance was positively correlated with host insect number, and parasitism rates were negatively density dependent. Host-plant phenological stage was positively correlated with herbivore and parasitoid abundance, and parasitism rates at both patch and host-plant scales.
5. The differential response between herbivore and parasitoid to host-plant density did not lead to a spatial refuge but may have contributed to the observed parasitism rates being negatively density dependent. Heterogeneity in patch quality, mediated by variation in host-plant phenology, was more important than spatial habitat structure for both the herbivore and parasitoid populations, and for parasitism rates.     

Three ecological factors influencing riverine fish diversity in the Shubuto River system,Japan: habitat capacity,habitat heterogeneity and immigration

A major goal of stream ecology is to identify environmental gradients that shape riverine communities. We examined the relative importance of three ecological factors that have been hypothesized to influence a longitudinal pattern of fish diversity: habitat capacity, heterogeneity and immigration of diadromous fishes. Field surveys were carried out in the entire network of the Shubuto River system, Hokkaido, Japan. A hierarchical partitioning approach revealed that distance from the sea, a proxy for immigration potential of diadromous fishes, had the greatest explanatory capacity, by which 24.9 % of variation in fish species richness was explained. Habitat capacity (approximated by catchment area) was also identified as a significant predictor of fish diversity, whereas habitat heterogeneity brought little improvement to the model performance. These results reflect the fish fauna of the Shubuto River system, in which diadromous fishes are dominant in both abundance and species richness.     

Seasonal changes in habitat suitability for adult shovelnose sturgeon in the lower Mississippi River

Shovelnose sturgeon Scaphirhynchus platorynchus are a large‐river fish distributed throughout the Mississippi River basin, including the lower 1,533 km of the Mississippi River where riverine habitat has been and continues to be modified for navigation and is a potential site for development of instream hydrokinetic electric power generation. Information about habitat use and preference is essential to future conservation efforts. Shovelnose sturgeon have previously been found to select particular habitat types, and these selected habitats vary seasonally; although these past analyses do not consider the selected habitats in a landscape context. We used ecological niche factor analysis (ENFA) that uses distributions of telemetry locations and environmental variables to model habitat suitability in a landscape context. We recorded 333 locations of shovelnose sturgeon during July–December 2013 that included periods of relatively high and low river stages. The ENFA analysis indicated high‐suitability locations were in or near deep water during both high and low river stages. During high river stages, high‐suitability locations were near island tip habitat, deep water, and steep bottom slope and far from main channel habitat. During low stages, high‐suitability locations were in or near deep water and main channel habitat and far from secondary channel and wing dike habitats. This landscape‐scale analysis supports seasonal shifts in habitat use and provides insights that can be used to inform habitat conservation and management to benefit shovelnose sturgeon in the lower Mississippi River and possibly other large rivers.     

The relative influence of local to regional drivers of variation in reef fishes

In this study, fishes and habitat attributes were quantified, four times over 1 year, on three reefs within four regions encompassing a c. 6° latitudinal gradient across south-western Australia. The variability observed was partitioned at these spatio-temporal scales in relation to reef fish variables and the influence of environmental drivers quantified at local scales, i.e. at the scale of reefs (the number of small and large topographic elements, the cover of kelp, fucalean and red algae, depth and wave exposure) and at the scale of regions (mean and maximum nutrient concentrations and mean seawater temperature) with regard to the total abundance, species density, species diversity and the multivariate structure of reef fishes. Variation in reef fish species density and diversity was significant at the regional scale, whereas variation in the total abundance and assemblage structure of fishes was also significant at local scales. Spatial variation was greater than temporal variation in all cases. A systematic and gradual species turnover in assemblage structure was observed between adjacent regions across the latitudinal gradient. The cover of red algae within larger patches of brown macroalgae (a biological attribute of the reef) and the number of large topographic elements (a structural attribute of the reef) were correlated with variation observed at local scales, while seawater temperature correlated with variation at the scale of regions. In conclusion, conservation efforts on reef fishes need to incorporate processes operating at regional scales with processes that shape local reef fish communities at local scales.     

Gradients in stream fish assemblages across a Mediterranean landscape: contributions of environmental factors and spatial structure

1. Despite wide recognition that fish assemblages are influenced by factors operating over a range of spatial scales, little effort has been devoted to quantifying large‐scale variation and the multiscale dependencies of assemblage patterns and processes. This is particularly true for Mediterranean streams, where seasonally predictable drying‐up may lead to a strong association between assemblage attributes and large‐scale factors affecting the distribution of population sources and extinction likelihood. 2. The contribution of large‐scale factors to stream fish assemblage variation was quantified across a Mediterranean landscape, in south‐west Portugal. Fish abundance and species composition were estimated at 166 sites across third‐ to sixth‐order streams, in March–July 1998. Variance partitioning by redundancy analyses was used to analyse assemblage variation against three sets of predictor variables: environmental (catchment position, and geomorphic and hydrological factors), large‐scale spatial trends and neighbourhood effects. 3. Environmental variables and spatial trends accounted for 34.6% of the assemblage variation across the entire region, and for 36.6 and 57.8% within the two largest catchments (Mira and Seixe). Neighbourhood effects were analysed at the catchment scale, increasing the explained variation to 56.1% (Mira) and 70.7% (Seixe). 4. A prevailing environmental gradient was reflected in an increase in the abundance of all species and size‐classes in relation to catchment position, with more fish present in larger streams and in downstream reaches. Variables describing geomorphic and hydrological settings were less important in explaining assemblage variation. 5. Spatial trends always accounted for the smallest fraction of assemblage variation, and they were probably associated with historical barriers to fish dispersal. The strong neighbourhood effects may be related to spatially autocorrelated habitat conditions, but they are also a likely consequence of fish emigration/extinction and colonisation processes. 6. These results emphasise that a substantial proportion of fish assemblage variation in Mediterranean streams may be explained by large‐scale factors, irrespective of microhabitats and local biotic interactions. It is suggested that this pattern results to a large extent from the seasonal drying‐up, with the summer shortage of surface water limiting fish occurrence in headwaters, and consequently the key core areas for fish concentrating in larger streams and tributaries adjacent to large streams because of neighbourhood effects.