Geigeria alata in the Namib desert: seed heteromorphism in an extremely arid environment

Abstract. Seed heteromorphism in annual Geigeria alata in the Namib Desert was investigated in relation to environmental conditions. Geigeria alata shows heteromorphism in seed dispersal distance depending on the position of the seeds on the parent plant. Seeds produced in capitula on branches (aerial) travel long distances, while seeds produced at the base of the plant remain in the vicinity of the parent plant. The ratio of aerial to basal seed production was investigated under different environmental conditions following an aridity gradient. Although there were significant differences in seed and capitula production between seasons and between different positions on the plant, no overall trend was found with increasing aridity.     

Limiting factors for seedling emergence and establishment of the invasive non-native Rosa rugosa in a coastal dune system

The relative importance of seed, habitat and microhabitat limitation has rarely been investigated for invasive non-native species, although this is critical for their effective management and for predicting future range expansion. Rosa rugosa is an abundant non-native shrub in coastal habitats of NW Europe; it is common along the Baltic coast but more scarce in exposed dunes of the North Sea. To investigate whether invasion of exposed dunes is limited by seed, habitat and microhabitat limitation, seedling emergence and establishment were examined in a factorial sowing, transplant and disturbance experiment. Twenty plots were randomly placed in each of five dune habitats (white dune, Empetrum dune, grey dune, outer dune heath, inner dune heath), and studied over 2 years. Seedling emergence in control subplots was zero in all habitats, whereas 96% and 98% of the undisturbed and disturbed seeded subplots produced seedlings. Disturbance had a positive effect on emergence and subsequent survival in white dune, outer and inner dune heath. Seedling survival and growth, and growth of transplanted seedlings, were markedly lower in grey dune. These findings indicate that establishment of R. rugosa is seed-limited in coastal dune habitats, and that the species is able to establish in both active and fixed dunes once seeds have arrived. Although differential seedling emergence and growth indicate that habitats differ in their degree of invasion susceptibility by R. rugosa, the positive influence of small-scale disturbance suggests microhabitat limitation in some dune habitats as well. Dune management should aim to reduce seed production and dispersal of R. rugosa in near-natural sites, and anthropogenic changes of habitat dynamics should be prevented.     

In vitro culture of Geigeria aspera Harv.

Callus-, root-, shoot-, and rooted shoot cultures of Geigeria aspera were established on Murashige and Skoog's nutrient medium. Organogenesis of roots and shoots was induced with treatments containing BA or KN in combination with NAA, IAA, IBA or 2, 4-D. Rooted plantlets were successfully transplanted into soil.     

Microhabitat use in a mediterranean riverine fish assemblage

Summary We examined microhabitat use in Barbus graellsii, Barbus haasi, Chondrostoma toxostoma, Rutilus arcasii, and Salmo gairdneri over a 19 month period in the upper Rio Matarraña, Spain. B. graellsii and Ch. toxostoma exhibited non-random microhabitat use during all seasons and preferentially occupied deep microhabitats with heterogeneous substrates. During the majority of seasons in which they were present, B. haasi and R. arcasii occurred in microhabitats similar to those occupied by B. graellsii and Ch. toxostoma. S. gairdneri was over-represented in high velocity microhabitats with erosional substrates. We did not observe any evidence of interspecific interference competition or avoidance. Substrate composition did not appear to affect microhabitat use outside of its covariation with depth and velocity. Seaonal variation in microhabitat use by B. graellsii, B. haasi and Ch. toxostoma was strongly correlated with seasonal changes in microhabitat availability. S. gairdneri, however, occurred closer to the substrate when average velocities were high. Larger B. graellsii and B. haasi sometimes occupied deeper, higher velocity microhabitats than did smaller specimens. Larger B. graellsii also occasionally occurred farther from shelter than did smaller specimens; the reverse was true for B. haasi. Larger Ch. toxostoma sometimes were found farther from both the substrate and shelter than smaller individuals, whereas smaller specimens occasionally inhabited deeper areas with more depositional substrates than did larger Ch. toxostoma. During Late Summer 1985, smaller Ch. toxostoma also occupied microhabitats with higher velocities than did larger specimens. A comparison of microhabitat use for two species present in both upper and lower portions of the Matarraña indicated that most differences in microhabitat use could be attributed to inter-site differences in microhabitat availability. The data suggest, hovever, that both species shifted to more protected microhabitats in the higher velocity site. Assemblage members generally occupied statistically distinguishable microhabitats and could be classified as: 1) high-velocity upper water column (S. gairdneri), 2) low velocity lower water column (B. graellsii, Ch. toxostoma and R. arcasii), and 3) shelter-oriented benthic (B. haasi). The introduction of S. gairdneri during Winter 1984 did not produce microhabitat shifts in any of the native species. Whether or not the native species affected microhabitat use in S. gairdneri is unknown. Interspecific competition for space, however, did not appear to strongly influence microhabitat use among the native species.     

Moms are better nurses than dads: gender biased self‐facilitation in a dioecious Juniperus tree

Questions: Can gender of nurse plants affect regeneration patterns and spatial population structure? Is there a seed‐seedling conflict in the regeneration process? What factors are responsible for the clumped spatial population structure observed for adult trees? Location: Mediterranean cold semi‐arid high mountains in Spain. Methods: The spatial pattern of adult Juniperus thurifera trees was studied by means of Ripley's K‐analysis. χ² analyses were used to test for natural seedling frequency in each of three main microhabitats: (1) under female and (2) male tree canopies and (3) in open interspaces. The observed pattern was explained experimentally by studying seed and seedling survival for two years. Survival probabilities were calculated across life stages for each of three main microhabitats. Results: Adult J. thurifera trees were aggregated in space. Most seedlings were found underneath female J. thurifera trees. Experimental studies demonstrated that from seed dispersal to seedling survival all life stages showed the same positive or negative trend within a given microhabitat, indicating stage coupling and no seed‐seedling conflicts. Attraction of frugivo‐rous birds by reproductive female junipers and improvement of environmental conditions beneath tree canopies were the main factors responsible for the variation in seedling density among microhabitats; highest underneath female trees and lowest in open interspaces. Conclusions: In dioecious species, the gender of nurse plants can significantly determine the spatial population structure. In J. thurifera forests, facilitation beneath female trees occurs among all life stages without any sign of seed‐seedling conflict. The most critical factors shaping the spatial population structure were directed seed dispersal and environmental amelioration beneath female conspecific trees.     

Microhabitat and morphometric variation of two Chaetophoracean (Chaetophorales, Chlorophyta) species in tropical streams of southeastern Brazil

Two populations of Chaetophora elegans (Roth) C. Agardh and two of Stigeoclonium helveticum Vischer were investigated for microhabitat characteristics and morphological variation in streams of São Paulo State, southeastern Brazil. Different patterns of microhabitat distribution were found between species investigated. Populations of C, elegans were distributed under relatively narrow microhabitat conditions (high irradiance, low depth, moderate to high current velocity, rocky substrata and lower values of niche width) and showing little morphometric variation (colony diameter, main axis cell size, and apical branch number), Stigeoclonium helveticum occurred under more diverse microhabitat conditions, revealed by lack of significant difference between sampling units with and without the alga and wider niche width, but also exhibited relatively narrow morphometric variation (plant length, main axis cell and ateral branch cell sizes). The narrow microhabitat conditions and smaller niche width of C. elegans can explain its low abundance (percentage cover) in streams from the area studied as well as in other regions of São Paulo State, In contrast, the wider variation of microhabitat conditions and the higher niche widths of S. helveticum suggest that this green alga is able to grow in a high number of stream ecosystems in the region investigated, ranging from undisturbed to highly disturbed habitats. Thus. the results suggest that S. helveticum is a generalist species.     

A cladistic analysis ofAnisopappus (Asteraceae: Inuleae)

A cladistic analysis of the genusAnisopappus (Asteraceae: Inuleae) has been undertaken. A hypothesis of species interrelationships in the genus is presented for the first time. The analysis also includedArctotis (Arctoteae), used as outgroup, and five additional genera from theInuleae: Geigeria, Calostephane, Asteriscus, Buphthalmum, Pulicaria, andInula. It is concluded thatAnisopappus is a monophyletic group situated at the base of the tribe, diagnosed by, e.g., their obtuse stylar sweeping-hairs. The species with acute sweeping-hairs were found to be derived within the genus. Problems concerning species delimitation, biogeography and character evolution in the genus are briefly discussed.     

Microhabitat segregation in two desert rodent species: the relation of prey availability to diet

Summary I studied diet in relation to microhabitat use in two desert rodents:Microdipodops megacephalus, the dark kangaroo mouse, andPeromyscus maniculatus, the deer mouse. Contrary to expectation, both species ate primarily arthropods, which were most abundant near shrubs.Peromyscus used the area near shrubs, in contrast toMicrodipodops, which used open microhabitat. As a consequence, the diet ofPeromyscus was narrower and more concentrated on abundant prey types than that ofMicrodipodops. Thus microhabitat segregation, which is frequently reported for desert rodents, is related to a diet-breadth difference between these rodents. The use of open microhabitat and low density resources byMicrodipodops, when compared with the large bipedalDipodomys and small quadrupedalPerognatus, suggests that bipedal locomotion in desert rodents is related to use of open microhabitat, and that body size is related to density of food resources.     

Importance of microhabitat and acorn burial on Quercus ilex early recruitment: non-additive effects on multiple demographic processes

This study investigates the effect of microhabitat and seed burial on the main demographic processes operating during the early recruitment of Quercus ilex, such as postdispersal seed predation, seed germination, and seedling emergence, survival and growth. The effect of burial was positive over all the processes analysed in this study, since predation rate was lower (63.6% vs. 88%), whereas germination (53.1% vs. 21.8%) and emergence (32.0% vs. 5.5%) were higher for buried acorns. The quality of some microhabitats remained similar throughout the stages and processes studied. Thus, afforestation provided especially suitable microhabitats for oak establishment, since seed predation was lower, while germination, emergence and seedling survival were higher, than in any other microhabitat. By contrast, the quality of some microhabitats, such as open sites and Holm oaks, differed between recruitment stages. Acorns in open sites escaped predation and germinated easily, but most seedlings died due to summer drought. Similarly, although acorns under Holm oaks can germinate and survive drought, they cannot survive to postdispersal predators. This uncoupling results in a post-dispersal change in the spatial distribution of Q. ilex recruits. Furthermore, there were significant interactions between burial and microhabitat for some demographic processes. The recruitment was in afforestations high irrespective of burial, suggesting that burial is not as beneficial in high-quality habitats as it is in lower-quality ones. An accurate understanding of plant recruitment requires the determination not only of the direct effects of limiting factors but also the potential interactions occurring between them.     

Stream microhabitat selectivity,resource partitioning,and niche shifts in grazing caddisfly larvae

Microhabitat selectivity, resource partitioning, and niche shifts in five species of grazing caddisfly larvae (Glossosoma califica, G. penitum, Dicosmoecus gilvipes, Neophylax rickeri, and N. splendens) were quantified by underwater measurement of microhabitat availability and utilization in three northern California streams. The microhabitat parameters water depth and velocity and rock size, roughness, and slope were measured. Comparisons of habitat available to habitat used revealed significant selection for at least two microhabitat parameters by each population, with depth and velocity being the most important. Comparisons of habitat used by different species showed significant partitioning of at least two microhabitat parameters at each site, with depth being partitioned at all sites. Non-parametric discriminant analysis revealed significant microhabitat partitioning on a multivariate level at two sites. Comparisons of habitat used at different sites quantified a major niche shift by D. gilvipes in its preference for riffles versus pools. Size-selective predation by dippers (Cinclus mexicanus) and steelhead (Salmo gairdneri gairdneri) is proposed as a hypothesis to explain the observed resource partitioning and niche shift.     

Microhabitat Selection of three Forest Understory Birds in the Brazilian Atlantic Rainforest

When assessing fragmentation effects on species, not only habitat preferences on the landscape scale, but also microhabitat selection is an important factor to consider, as microhabitat is also affected by habitat disturbance, but nevertheless essential for species for foraging, nesting and sheltering. In the Atlantic Rainforest of Brazil we examined microhabitat selection of six Pyriglena leucoptera (white-shouldered fire-eye), 10 Sclerurus scansor (rufous-breasted leaftosser), and 30 Chiroxiphia caudata (blue manakin). We radio-tracked the individuals between May 2004 and February 2005 to gain home ranges based on individual fixed kernels. Vegetation structures in core plots and fringe plots were compared. In C. caudata, we additionally assessed the influence of behavioural traits on microhabitat selection. Further, we compared microhabitat structures in the fragmented forest with those in the contiguous, and contrasted the results with the birds' preferences. Pyriglena leucoptera preferred liana tangles that were more common in the fragmented forest, whereas S. scansor preferred woody debris, open forest floor (up to 0.5 m), and a thin closed leaf litter cover which all occurred significantly more often in the contiguous forest. Significant differences were detected in C. caudata for vegetation densities in the different strata; the distance of core plots to the nearest lek site was significantly influenced by sex and age. However, core sites of C. caudata in fragmented and contiguous forests showed no significant differences in structure. Exploring microhabitat selection and behavior may greatly support the understanding of habitat selection of species and their susceptibility to fragmentation on the landscape scale.     

Microhabitat use in coexisting desert rodents—The role of population density

Summary Among some species of Sonoran Desert rodents microhabitat differences are density dependent. I studied the differences in microhabitat use among four species of heteromyid rodents (Dipodomys merriami, Perognathus amplus, P. baileyi, and P. penicillatus) at low and at high population densities. Microhabitats are defined by the abundance and size distribution of desert shrubs. During a period of low population density the rodent species showed substantial microhabitat differentiation. Following a large increase in pocket mouse (Perognathus spp.) numbers differences in microhabitat use between species disappeared. The lack of microhabitat differentiation at high density is due to microhabitat shifts rather than an expansion in the number of microhabitats used. The shifts lead to increased similarity among species in microhabitat use. Microhabitat overlap is not constant but it is highly variable and sensitive to changes in rodent abundance.     

Variability in life history parameters of four serotinous plants in the Namib Desert

Plants growing in extreme habitats, such as deserts, are expected to show high plasticity (variability) in life history parameters to compensate for the unpredictability of their environment. In this study variability in life history parameters and possible sources of variability were investigated in four serotinous (= seed retaining) plant species from the Namib Desert.Blepharis grossa, Geigeria alata, Geigeria ornativa andPetalidium setosum were studied in a mosaic of spatio-temporal environmental conditions represented by three sites and three rain seasons. The coefficient of variation and whether or not significant differences occurred between environmental conditions were used to determine the most variable species and parameters. The most variable species was the strictly annualB. grossa, which might be related to its life form strategy and poor competitive ability as a pioneer species. Life history parameters which responded most frequently to varying environmental conditions were source pool, flowering time and seed production. Seedling density was influenced by the source pool and intensity of rainfall, seedling establishment by timing and intensity of rain events, as well as by seed mass. Although it was expected that high within population variability in life history parameters would also result in a response to differing environmental conditions, there was no direct correlation. Variable responses in life history parameters in most species were caused by differences in site condition as well as by differences in seasons, especially with regard to timing and intensity of rain events.     

Effects of seed dispersal on Juniperus communis recruitment on a Mediterranean mountain

Abstract. The recruitment of the relict shrub Juniperus communis on a mountain in SE Spain was studied during the period 1994–1998. The main objective was to determine both the quantitative and qualitative effects of bird dispersal on seedling establishment. Seed removal by birds, seed rain, post‐dispersal seed predation, germination, and seedling emergence and survival were analysed in different microhabitats. Birds removed 53 ‐ 89% of the seeds produced by plants. Seed rain was spatially irregular as most seeds accumulated near stones used by birds as perches and below mother plants while a few seeds were dropped in wet meadows and open ground areas. Post‐dispersal seed predation by rodents affected < 10% of dispersed seeds but varied significantly among microhabitats. Only 3.6 ‐ 5.5% of dispersed seeds appeared viable, as many seeds had aborted or showed wasp damage. Seeds germinated in the second and third springs after sowing, reaching a germination percentage of 36%. Seedling emergence was concentrated in wet meadows. Seedling mortality was high (75–80%), but significantly lower in wet meadows, the only microhabitat where seedlings could escape from summer drought, the main mortality cause. Seed abortion, germination and seedling mortality proved to be the main regeneration constraints of J. communis on Mediterranean mountains. Birds exerted a strong demographic effect, although their qualitative effect was limited by abiotic factors which caused the pattern of seed rain to differ from the final pattern of recruitment between microhabitats.     

Niche partitioning and the effect of interspecific competition on microhabitat use by two sympatric galaxiid stream fishes

1. Numerous interacting abiotic and biotic factors influence niche use and assemblage structure of freshwater fishes, but the strength of each factor changes with spatial scale. Few studies have examined the role of interspecific competition in structuring stream fish assemblages across spatial scales. We used field and laboratory approaches to examine microhabitat partitioning and the effect of interspecific competition on microhabitat use in two sympatric stream fishes (Galaxias‘southern’ and Galaxias gollumoides) at large (among streams and among sites within streams) and small (within artificial stream channels) spatial scales. 2. Diurnal microhabitat partitioning and interspecific competition at large spatial scales were analysed among three sympatry streams (streams with allotopic and syntopic sites; three separate catchments) and four allopatry streams (streams with only allotopic sites; two separate catchments). Electro‐fishing was used to sample habitat use of fishes at 30 random points within each site by quantifying four variables for each individual: water velocity, depth, distance to nearest cover and substratum size. Habitat availability was then quantified for each site by measuring those variables at each of 50 random points. Diet and stable isotope partitioning was analysed from syntopic sites only. Diel cycles of microhabitat use and interspecific competition at small spatial scales were examined by monitoring water velocity use over 48 h in artificial stream channels for three treatments: (i) allopatric G. ‘southern’ (10 G. ‘southern’); (ii) allopatric G. gollumoides (10 G. gollumoides) and (iii) sympatry (five individuals of each species). 3. One hundred and ninety‐four G. ‘southern’ and 239 G. gollumoides were sampled across all seven streams, and habitat availability between the two species was similar among all sites. Galaxias‘southern’ utilised faster water velocities than G. gollumoides in both the field and in channel experiments. Both species utilised faster water velocities in channels at night than during the day. Diet differences were observed and were supported by isotopic differences (two of three sites). No interspecific differences were observed for the other three microhabitat variables in the field, and multivariate habitat selection did not differ between species. Interspecific competition had no effect on microhabitat use of either species against any variable either in the field (large scale) or in channels (small scale). 4. The results suggest that niche partitioning occurs along a subset of microhabitat variables (water velocity use and diet). Interspecific competition does not appear to be a major biotic factor controlling microhabitat use by these sympatric taxa at any spatial scale. The results further suggest that stream fish assemblages are not primarily structured by biotic factors, reinforcing other studies de‐emphasising interspecific competition.     

Implications of microhabitat selection and patterns of activity on the thermal ecology of high elevation neotropical anurans

This paper focuses on whether microhabitat selection reduces exposure to cold temperatures in highelevation tropical amphibians. I studied the microhabitat associations and thermal ecology of four anurans (Colostethus subpunctatus, Atelopus sp. nov., Eleutherodactylus bogotensis, and Hyla labialis) that live in the Andes of central Colombia. I compared two populations (3500 m and 2900 m) to evaluate the impact of a change in elevation on microhabitat thermal regimes and frog behavior. Ambient temperatures of frequently used microhabitats (soil surface, retreat sites, bogs, and ponds) were significantly lower at 3500 m. However, for each species the microhabitat associations and general patterns of activity were similar at both elevations. Body temperatures of active frogs were mainly determined by the interaction between microhabitat selection and time of the day, therefore frogs at 3500 m had lower activity temperatures than conspecifics at 2900 m. Terrestrial and diurnal anurans (Atelopus sp. nov.) experienced highly variable activity temperatures. Frogs associated with water bodies (C. subpunctatus, and H. labialis) experienced more moderate and less variable temperatures. Terrestrial, nocturnal frogs (E. bogotensis) exhibited the lowest activity temperatures, and their thermal ecology was strongly affected by increase in altitude. The physical nature of high tropical elevations limits the opportunities for behavioral thermoregulation in amphibians, and increases the importance of physiological adjustments to tolerate a wide thermal range. This may partially explain why only a small number of amphibian species, representing genera that are much more diverse at lower elevations, have been able to exploit these environments.     

The influence of conditioning on olfactory microhabitat and host location in Asobara tabida (Nees) and A. rufescens (Foerster) (Braconidae: Alysiinae) larval parasitoids of Drosophilidae

Olfactory responses of parasitoids can be variable. This was shown by olfactometer experiments with females of two sibling Asobara species, larval endoparasitoids of Drosophilidae. Oviposition experiences of adult female parasitoids significantly altered their behavioural responses in microhabitat and host location. Females needed prior exposure to a host before they were capable of using volatile compounds related to the presence of their host. Asobara tabida (Nees) attacks Drosophila in fermenting fruits and A. rufescens (Foerster) attacks Drosophilidae in decaying plant materials. Naïve females show a strong preference for the odour of their own microhabitat. After experience with their own host and microhabitat, females were repelled by odours of the other microhabitat. Enforced experience with this repellent microhabitat in the laboratory modified the olfactory response from repellency to attraction. It was shown that even the microhabitat odour preference pattern could be changed through experience. This kind of behavioural flexibility may be the rule rather than the exception in many other Hymenopterous parasitoids.     

Comparative microhabitat use of cyprinid larvae and juveniles in a lotic floodplain channel

Synopsis The microhabitat of six cyprinids in a braided side-channel of the Upper Rhône River, France, was studied June to September 1985, using point abundance sampling by electrofishing. Correspondence analysis of the samples-by-species data and Jacob&s electivity index (calculated from the availability and exploitation of eight environmental variables) revealed three groups of species: (1) Leuciscus cephalus and Alburnus alburnus, which used relatively similar microhabitat during both larval and 0 + juvenile development; (2) Rutilus rutilus, Leuciscus leuciscus and Chondrostoma nasus, which used different microhabitats as larvae but whose microhabitat overlapped notably during 0 + juvenile development; (3) and Gobio gobio, whose juvenile microhabitat overlapped only marginally with that of other 0 + juveniles (no data were available on larvae). Overlap in microhabitat use by 0 + juveniles increased notably during a period of reduced discharge, when the amount of available vegetal and ligneous structures decreased; while most species exploited the increased area of shallow waters, juvenile A. alburnus appeared to be the only species that exploited the deeper waters off the channel&s steeper banks. Although not quantified, numerous field observations of predation threat, particularly during the period of reduced discharge (which reduced the amount of cover), suggest that predation risk may influence the microhabitat use of young fishes. Compared with similar species in European lakes, the overlap in microhabitat use of these fluvial cyprinids appears to be more intensive.     

Habitat selection by two streamside plethodontid salamanders

Summary A series of experiments was conducted to test the hypothesis that habitat selection by the salamanders Desmognathus monticola and Desmognathus quadramaculatus is influenced differently by interspecific interference than by intraspecific interference. Individuals of neither species were influenced differently by the presence of heterospecifics than by the presence of conspecifics in their activity away from cover sites, in selection of different substrate moisture levels, in selection of different substrate textures, nor in selection of microhabitats relative to the streambed. Both species generally selected substrates with the highest moisture and the coarsest texture. Selection of different cover sizes by D. monticola depended on microhabitat, with larger rocks being selected most frequently when salamanders were farther from the streambed. D. quadramaculatus was seldom found away from the streambed and did not discriminate among the cover sizes available. The strong affinity of D. quadramaculatus for streambed microhabitat, compared with the much more variable microhabitat selection of D. monticola, resulted in spatial separation of individuals of the two species most of the time. Results of this study do not support the hypothesis that interspecific inference is more important than intraspecific interference in the microhabitat distributions of D. monticola and D. quadramaculatus.     

Scale-dependence of fish habitat associations in a North American river system

1. Determining the appropriate measurement scale to assess habitat variables is critical for ecologists assessing biological or ecological conditions. Depth, velocity, substrate, woody debris and other fish cover variables occur on both reach and microhabitat scales, and fish habitat associations with these variables may be scale-dependent. The aim of this work was to better understand the importance of scale for fish–habitat associations with these variables in a framework consistent with environmental filtering and to test the hypothesis that habitat variable importance is scale-dependent.
2. I used prepositioned areal electrofishing in wadeable streams of the Delaware River basin to evaluate the associations of fish with the same variables summarised on different reach and microhabitat scales. The importance of scale for fish–habitat associations was assessed using two approaches that approximate an environmental filtering framework: variance partitioning with (1) ordination and (2) generalised linear mixed models.
3. Variables on both the reach and microhabitat scales explained a significant fraction of the total variation in fish community composition (p < 0.05). Variation decomposition of reach- and microhabitat-scale effects revealed 20.2% and 2.0% of all variation were due uniquely to reach and microhabitat scales, respectively. Measures of coarseness, embeddedness, amount of riffle and areal coverage of five fish cover variables were significant explanatory variables of community composition at the reach scale only (p < 0.05). Velocity and mesohabitat (amount or presence of riffle) were the only two habitat features that were significant explanatory variables of fish community composition at both the reach and microhabitat scales (p < 0.05). Individual models of species occurrence revealed similar patterns as seen with analyses of community composition.
4. For many fishes, habitat features quantified at the reach scale were more explanatory than at the microhabitat scale. Longnose dace (Rhinichthys cataractae) were more dependent upon microhabitat variables than reach-scale variables, relative to other fishes. Mean velocity at the reach scale was the most important explanatory variable for explaining fish community composition and indicated support for the concept of environmental filtering at the reach and microhabitat scales.
5. Few studies of fish occurrence have incorporated a study design and analytical framework that approximates the hierarchical nature of habitat. This study identifies important scales and predictors, demonstrates the importance of a multiscale approach, and provides support for the environmental filtering concept at the reach and microhabitat scales. These findings will allow ecologists to better account for scale-dependent habitat associations and justify the use of fish habitat associations on reach and microhabitat scales for assessing biotic integrity, restoration and conservation of fishes.