Variability in stream macroinvertebrates at multiple spatial scales

1. We intensively sampled 16 western Oregon streams to characterize: (1) the variability in macroinvertebrate assemblages at seven spatial scales; and (2) the change in taxon richness with increasing sampling effort. An analysis of variance (ANOVA) model calculated spatial variance components for taxon richness, total density, percent individuals of Ephemeroptera, Plecoptera and Trichoptera (EPT), percent dominance and Shannon diversity.
2. At the landscape level, ecoregion and among‐streams components dominated variance for most metrics, accounting for 43–72% of total variance. However, ecoregion accounted for very little variance in total density and 36% of the variance was attributable to differences between streams. For other metrics, variance components were more evenly divided between stream and ecoregion effects.
3. Within streams, approximately 70% of variance was associated with unstructured local spatial variation and not associated with habitat type or transect position. The remaining variance was typically split about evenly between habitat and transect. Sample position within a transect (left, centre or right) accounted for virtually none of the variance for any metric.
4. New taxa per stream increased rapidly with sampling effort with the first four to eight Surber samples (500–1000 individuals counted), then increased more gradually. After counting more than 50 samples, new taxa continued to be added in stream reaches that were 80 times as long as their mean wetted width. Thus taxon richness was highly dependent on sampling effort, and comparisons between sites or streams must be normalized for sampling effort.
5. Characterization of spatial variance structure is fundamental to designing sampling programmes where spatial comparisons range from local to regional scales. Differences in metric responses across spatial scales demonstrate the importance of designing sampling strategies and analyses capable of discerning differences at the scale of interest.     

Using hierarchical sampling to understand scales of spatial variation in early coral recruitment

Ecological patterns are created by processes acting over multiple spatial and temporal scales. By combining spatially explicit sampling with variance components models, the relative importance of spatial scale to overall variability can be determined. We used a spatially structured experimental design in the Mombasa Marine National Park in Kenya to quantify variation in coral recruitment across four spatial scales (~1–1,000 m) and to generate hypotheses about processes affecting recruitment and potential sources of post-settlement mortality during early life history. For the dominant recruiting corals (Pocillopora spp.), variation in recruitment on surfaces protected from fish grazing was greatest at the largest spatial scale examined (1,000 m). We hypothesize that recruitment on protected surfaces varies mainly with larval delivery due to different lagoonal circulation and water flow between sites. Conversely, variation on surfaces exposed to fishes was greatest at the smallest spatial scale (1 m). We hypothesize that recruitment on exposed surfaces mainly reflects local differences in the scale and intensity of fish grazing, which may obscure larval delivery patterns. Spatial variation in recruitment can affect many ecological processes and factors, including growth, survival to maturity, the distribution of habitat, and variation in species interaction strengths. This study demonstrates how spatially explicit sampling, followed by variance components modeling to partition variance across scales, can help to identify potential drivers of patterns at each relevant scale.     

On the roles of landscape heterogeneity and environmental variation in determining population genomic structure in a dendritic system

Dispersal and natural selection are key evolutionary processes shaping the distribution of phenotypic and genetic diversity. For species inhabiting complex spatial environments however, it is unclear how the balance between gene flow and selection may be influenced by landscape heterogeneity and environmental variation. Here, we evaluated the effects of dendritic landscape structure and the selective forces of hydroclimatic variation on population genomic parameters for the Murray River rainbowfish, Melanotaenia fluviatilis across the Murray–Darling Basin, Australia. We genotyped 249 rainbowfish at 17,503 high‐quality SNP loci and integrated these with models of network connectivity and high‐resolution environmental data within a riverscape genomics framework. We tested competing models of gene flow before using multivariate genotype–environment association (GEA) analysis to test for signals of adaptive divergence associated with hydroclimatic variation. Patterns of neutral genetic variation were consistent with expectations based on the stream hierarchy model and M. fluviatilis’ moderate dispersal ability. Models incorporating dendritic network structure suggested that landscape heterogeneity is a more important factor determining connectivity and gene flow than waterway distance. Extending these results, we also introduce a novel approach to controlling for the unique effects of dendritic network structure in GEA analyses of populations of aquatic species. We identified 146 candidate loci potentially underlying a polygenic adaptive response to seasonal fluctuations in stream flow and variation in the relative timing of temperature and precipitation extremes. Our findings underscore an emerging predominant role for seasonal variation in hydroclimatic conditions driving local adaptation and are relevant for informing proactive conservation management.     

Patterns of temporal variation in Lake Titicaca. A high altitude tropical lake. I. Background,physical and chemical processes,and primary production

A statistical analysis is presented of patterns of variation in some physical, chemical, and biological variables for a 6 year series of data from the tropical, high altitude Lake Titicaca (Peru-Bolivia). ANOVA techniques and autocorrelation analyses were used to partition the variance in Titicaca, and in some comparison tropical and temperate series, into components with repeatable annual cycles and components attributable to other kinds of patterns.In Titicaca, insolation and stratification are highly seasonal in pattern of variation, although the amount of variance relative to means is small compared to temperate lakes. However, the seasonal pattern of physical variation is only weakly imposed on chemical and biological processes, to judge from analyses of silicate, oxygen, and primary production series. Comparable temperate series of primary production and chlorophyll a are much more seasonal.     

PEMPHIGUS REVISITED: CHANGES IN GEOGRAPHIC VARIATION BUT CONSTANCY IN VARIABILITY AND COVARIATION

Samples of the gall-forming aphids Pemphigus populicaulis and P. populitransversus (both elongate and globular morphs) were re-collected at sites in eastern North America after 13 to 16 years. Twenty-three morphometric characters of the galls, stem mothers, and alate fundatrigeniae were analyzed by univariate and multivariate methods. Varying proportions of the variance of each character are attributable to the four levels of variation—locality, year, year by locality interaction, and among galls (within one year and locality). The year by locality interaction level generally has the greatest variation and is highly significant. Year and locality effects tend to be lower and not significant. The variance components do not exhibit trends with time. Geographic variation patterns of single variables or factor scores in the original and revisited populations show significant spatial structure overall but lack clear-cut spatial patterns, especially clines. Observable patterns of variation match results of the spatial analyses: most characters lack clear trends; patterns in the revisited data do not resemble patterns for the same variables in the original data. Variability profiles for characters change little over the time span and are comparable among and within localities. Covariation among characters over localities is largely maintained during the time interval despite the changes in patterns. Fluctuating interclonal competition among aphids on secondary hosts is believed to cause the marked heterogeneity in space and time among the aphids in the galls.     

Limnological observations of an intermittent tropical dry forest stream

This study examines aspects of the seasonal limnology of an intermittent stream in the dry forest region of northwest Costa Rica. It focuses on annual water level fluctuations and both seasonal and among pool variation in dissolved oxygen concentration and water temperature. Dry season pools differed in morphometry and the rate of decline in water levels subsequent to the seasonal floods. Rate of water level decline was related both to pool depth and to the exposure of the pool to the sun. Oxygen concentration was generally low in the dry season, but increased during the rainy season in association with rain events and seasonal flooding. A repeated-measures analysis of variance indicated that seasonal effects accounted for 40% of the variance in oxygen concentration for residual pools. Differences among pools were also significant. However, there was strong evidence for a significant interaction between seasonal and spatial influences on oxygen values in the system. Variation in water temperature was small, but differences among pools, sampling dates, and their interaction were all significant. Periphyton production increased significantly between the late wet season sample in November and the dry season sample in February. Incident light intensity explained 76% of the variation among pools in net periphyton production in the dry season.     

The influence of mean climate trends and climate variance on beaver survival and recruitment dynamics

Ecologists are increasingly aware of the importance of environmental variability in natural systems. Climate change is affecting both the mean and the variability in weather and, in particular, the effect of changes in variability is poorly understood. Organisms are subject to selection imposed by both the mean and the range of environmental variation experienced by their ancestors. Changes in the variability in a critical environmental factor may therefore have consequences for vital rates and population dynamics. Here, we examine ≥90‐year trends in different components of climate (precipitation mean and coefficient of variation (CV); temperature mean, seasonal amplitude and residual variance) and consider the effects of these components on survival and recruitment in a population of Eurasian beavers (n = 242) over 13 recent years. Within climatic data, no trends in precipitation were detected, but trends in all components of temperature were observed, with mean and residual variance increasing and seasonal amplitude decreasing over time. A higher survival rate was linked (in order of influence based on Akaike weights) to lower precipitation CV (kits, juveniles and dominant adults), lower residual variance of temperature (dominant adults) and lower mean precipitation (kits and juveniles). No significant effects were found on the survival of nondominant adults, although the sample size for this category was low. Greater recruitment was linked (in order of influence) to higher seasonal amplitude of temperature, lower mean precipitation, lower residual variance in temperature and higher precipitation CV. Both climate means and variance, thus proved significant to population dynamics; although, overall, components describing variance were more influential than those describing mean values. That environmental variation proves significant to a generalist, wide‐ranging species, at the slow end of the slow‐fast continuum of life histories, has broad implications for population regulation and the evolution of life histories.     

Photosynthetic activity of US biomes: responses to the spatial variability and seasonality of precipitation and temperature

We examined the response of the normalized difference vegetation index, integrated over the growing season (gNDVI) to mean precipitation, maximum temperature (T_max), and minimum temperature (T_min) over an 11‐year period (1990–2000) for six biomes in the conterminous United States. We focused on within‐ and across‐biome variance in long‐term average gNDVI, emphasizing the degree to which this variance is explained by spatial gradients in long‐term average seasonal climate. Since direct measurements of ecosystem function are unavailable at the spatial and temporal scales studied, we used the satellite‐based gNDVI as a proxy for net photosynthetic activity. Forested and nonforested biomes differed sharply in their response to spatial gradients in temperature and precipitation. Gradients in mean spring and fall precipitation totals explained much of the variance in mean annual gNDVI within arid biomes. For forested biomes, mean annual gNDVI was positively associated with mean annual and seasonal T_min and T_max. These trends highlight the importance of the seasonal components of precipitation and temperature regimes in controlling productivity, and reflect the influence of these climatic components on water balance and growing‐season length. According to the International Panel on Climate Change (IPCC) (2001) increases in temperature minima and fall precipitation have contributed the dominant components of US increases in temperature and precipitation, respectively. Within the range of conditions observed over the study region, our results suggest that these trends have particularly significant consequences for above‐ground plant productivity, especially for Grassland, Open Shrubland, and Evergreen Needleleaf Forest. If historical climatic trends and the biotic responses suggested in this analysis continue to hold, we can anticipate further increases in productivity for both forested and nonforested ecoregions in the conterminous US, with associated implications for carbon budgets and woody proliferation.     

Spatial, seasonal and diel distribution patterns of two species of mojarras (Pisces: Gerreidae) in a Mexican tropical coastal lagoon

Spatial, seasonal and diel distribution patterns were analysed over a 2‐year period for Diapterus auratus and Eucinostomus melanopterus in Pueblo Viejo lagoon, a tropical coastal lagoon in Veracruz, Mexico. A total of 216 samples were collected with 329 individuals of D. auratus and 170 of E. melanopterus. Diapterus auratus individuals were captured throughout a wider range of environmental conditions, and consequently its spatial niche breadth was significantly greater (P < 0.001) than that of E. melanopterus. In general, more individuals were captured at localities with submerged vegetation (Ruppia maritima), with 70% for D. auratus and 87.2% for E. melanopterus, although only the latter species showed significant spatial differences. A spatial segregation influenced by distance from the lagoon inlet and salinity was observed between the species. It is considered that this segregation may allow resource partitioning and minimize interspecific competition. A relatively low niche overlap (4.9%) was observed for the two species. Seasonal capture peaks of D. auratus were related to rainfall and the productivity pattern in the lagoon system. Seasonal fluctuations in monthly mean number of D. auratus were correlated with preceding variations in local rainfall (r = 0.86, P < 0.02). Moreover, seasonal capture peaks of D. auratus coincided with ecosystem primary production peaks. Although E. melanopterus exhibited no significant seasonal differences, its maximum numbers also coincided with an ecosystem primary production peak and recruitment patterns. In 24‐h cycle analysis, both species were captured mainly around mid‐day, with minor peaks at dawn and/or at dusk. Generally, few individuals were collected during hours of darkness. It was proposed that diel abundance variation of gerreids might be related to both light penetration in the water column and feeding activity. Food availability within Pueblo Viejo lagoon appears to be an important factor influencing fish abundance.     

Spatial and temporal variation in the morphology (and thus, predicted impact) of an invasive species in Australia

The impact of an invasive species is unlikely to be uniform in space or time, due to variation in key traits of the invader (e.g. morphology, physiology, behaviour) as well as in resilience of the local ecosystem. The weak phylogeographic structure typical of an invasive population suggests that much of the variation in an invading taxon is likely to be generated by the environment and recent colonisation history. Here we describe effects of the environment and colonisation history on key morphological traits of an invader (the cane toad Bufo marinus ). These "key traits" (body size and relative toxicity) mediate the impact of toads on Australian native predators, which often die as a consequence of ingesting a fatal dose of toad toxin. Measurements of museum specimens collected over >60 yr from a wide area show that seasonal variation in toad body size (due to seasonal recruitment) effectively swamps much of the spatial variance in this trait. However, relative toxicity of toads showed strong spatial variation and little seasonal variation. Thus, the risk to a native predator ingesting a toad will vary on both spatial and temporal scales. For native predators capable of eating a wide range of toad sizes (e.g. quolls, varanid lizards), seasonal variation in overall toad size will be the most significant predictor of risk. In contrast, gape-limited predators restricted to a specific range of toad sizes (such as snakes) will be most strongly affected by the relative toxicity of toads. Gape-limited predators will thus experience strong spatial variation in risk from toad consumption.     

The structure of sculpin populations along a stream size gradient

Synopsis Data on spatial variation of sculpin density, growth and fecundity support the hypothesis that populations of stream fish are structured by changes in risk of predation and prey availability along a gradient in stream size. Cottus bairdi in warm streams and C. cognatus in cold streams exhibit similar patterns. Sculpins in large streams have faster individual growth rates and higher fecundities than those in small streams, but occur at lower density. The patterns appear to be persistent and suggest that predation reduces sculpin density in larger streams. Competitive release, variations in prey productivity, and local factors probably contribute to the variation in sculpin growth.     

Reproductive strategies of Amazonian stream fishes and their fine‐scale use of habitat are ordered along a hydrological gradient

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Multivariate analysis of a fine-scale breeding bird atlas using a geographical information system and partial canonical correspondence analysis: environmental and spatial effects

Aim To assess the relative roles of environment and space in driving bird species distribution and to identify relevant drivers of bird assemblage composition, in the case of a fine‐scale bird atlas data set. Location The study was carried out in southern Belgium using grid cells of 1 × 1 km, based on the distribution maps of the Oiseaux nicheurs de Famenne: Atlas de Lesse et Lomme which contains abundance for 103 bird species. Methods Species found in < 10% or > 90% of the atlas cells were omitted from the bird data set for the analysis. Each cell was characterized by 59 landscape metrics, quantifying its composition and spatial patterns, using a Geographical Information System. Partial canonical correspondence analysis was used to partition the variance of bird species matrix into independent components: (a) ‘pure’ environmental variation, (b) spatially‐structured environmental variation, (c) ‘pure’ spatial variation and (d) unexplained, non‐spatial variation. Results The variance partitioning method shows that the selected landscape metrics explain 27.5% of the variation, whilst ‘pure’ spatial and spatially‐structured environmental variables explain only a weak percentage of the variation in the bird species matrix (2.5% and 4%, respectively). Avian community composition is primarily related to the degree of urbanization and the amount and composition of forested and open areas. These variables explain more than half of the variation for three species and over one‐third of the variation for 12 species. Main conclusions The results seem to indicate that the majority of explained variation in species assemblages is attributable to local environmental factors. At such a fine spatial resolution, however, the method does not seem to be appropriated for detecting and extracting the spatial variation of assemblages. Consequently, the large amount of unexplained variation is probably because of missing spatial structures and ‘noise’ in species abundance data. Furthermore, it is possible that other relevant environmental factors, that were not taken into account in this study and which may operate at different spatial scales, can drive bird assemblage structure. As a large proportion of ecological variation can be shared by environment and space, the applied partitioning method was found to be useful when analysing multispecific atlas data, but it needs improvement to factor out all‐scale spatial components of this variation (the source of ‘false correlation’) and to bring out the ‘pure’ environmental variation for ecological interpretation.     

Scale-related patterns in the spatial and environmental components of stream macroinvertebrate assemblage variation

Aim We examined the relative contributions of spatial gradients and local environmental conditions to macroinvertebrate assemblages of boreal headwater streams at three hierarchical extents: bioregion, ecoregion and drainage system. We also aimed to identify the environmental variables most strongly related to assemblage structure at each study scale, and to assess how the importance of these variables is related to regional context and spatial structuring at different scales. Location Northern Finland ( 62 – 68° N, 25–32° E). Methods Variation in macroinvertebrate data was partitioned using partial canonical correspondence analysis into components explained by spatial variables (nine terms from the cubic trend surface regression), local environmental variables (15 variables) and spatially structured environmental variation. Results The strength of the relationship between assemblage structure and local environmental variables increased with decreasing spatial extent, whereas assemblage variation related to spatial variables and spatially structured environmental variation showed the opposite pattern. At the largest extents, spatial variation was related to latitudinal gradients, whereas spatial autocorrelation among neighbouring streams was the likely mechanism creating spatial structure within drainage systems. Only stream size and water acidity were consistently important in explaining assemblage structure at all study scales, while the importance of other environmental variables was more context‐dependent. Main conclusions The importance of local environmental factors in explaining macroinvertebrate assemblage structure increases with decreasing spatial extent. This scale‐related pattern is not caused solely by changes in study extent, however, but also by variable sample sizes at different regional extents. The importance of environmental gradients is context‐dependent and few factors are likely to be universally important correlates of macroinvertebrate assemblage structure. Finally, our results suggest that bioassessment should give due attention to spatial structuring of stream assemblages, because important assemblage gradients may not only be related to local factors but also to biogeographical constraints and neighbourhood dispersal processes.     

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.     

GENOTYPE-ENVIRONMENT INTERACTION FOR CLIMATE AND COMPETITION IN A NATURAL POPULATION OF FLOUR BEETLES,TRIBOLIUM CASTANEUM

The norm of reaction, the set of average phenotypes produced by a genotype in different environments, can be affected by spatial variation in natural selection especially when there exists genotype-environment interaction. In subdivided populations, the greater the genotype-environment interaction variance and the lower the migration rate, the more independent are the possible evolutionary trajectories for local adaptation. I examined genotype-environment interaction in the rate of population increase for lineages randomly derived from a wild population of Tribolium castaneum across a series of ecologically important environments. The lineages were derived from an outbred, wild-caught population by 14 generations of random genetic drift, during which the effective size of each lineage was approximately 22 breeding adults. The environments studied were the classic temperate-wet and cold-dry climates of Park (1954) in factorial combination with two genetic strains of a congeneric competitor, T. confusum. Much among-lineage genetic variation for rate of population increase was found for each of these ecologically important environments of climate and competition. Genotype-environment interaction accounted for 40.5% of the total among-lineage variance in rate of population increase signifying that the performance of a lineage in one environment is not necessarily a good predictor of its performance in another. Changing the genetic identity of the competitor changed the rate of increase of some lineages as much or more than changing the climatic conditions of temperature and humidity. This is the first empirical study to characterize the genotype-environment interaction variance associated with genetic variation in a competing congeneric species. This competitor-specific genetic variation in competitive ability may play an important role in coevolution in subdivided populations.     

Seasonal and spatial variability of soil respiration in four Sitka spruce stands

We investigated the causes for the seasonal and spatial variation of soil respiration in a first rotation Sitka spruce chronosequence composed of four age classes (10, 15, 31, and 47 year old) in Central Ireland. The study aimed at identifying easily determinable environmental parameters that explained the variation in soil respiration rates. The variation in temperature and soil water content influenced the seasonal trend observed in the spatial variability of soil respiration. The highest coefficients of variation in soil respiration were observed during autumn drought, while lower coefficients were generally observed during periods with highest soil respiration rates. On average, the sampling strategy of 30 sampling points per stand was adequate to obtain an average rate of soil respiration within 20% of its actual value at the 95% confidence level. Significantly higher soil respiration rates were observed at locations with high accumulation of organic matter and in collars established in close vicinity to tree stems. The organic layer thickness was the only variable that yielded significant regressions for explaining spatial variation in soil respiration in all the stands. Correlation analyses between the studied variables and soil respiration suggested the relative importance of heterotrophic and autotrophic components differed in their annual contribution to total soil respiration at each forest stand. Multiple regression analyses were used to assess the relative importance of primary temporal and spatial controls over soil respiration. Soil temperature and organic layer thickness explained most of the variance of soil respiration for the different sampling periods, while soil water content had a weaker effect as well as a different influence on soil respiration depending on the time of the year. The strong linear correlation between forest floor carbon and soil carbon stock further confirmed organic layer thickness as an integrative factor encompassing the effect of soil carbon pools on soil respiration. Moreover, its inclusion in the multiple regression analyses overrode the influence of both distance and fine root biomass. Overall, a multiple linear regression model driven by easily determinable environmental variables such as soil temperature, organic thickness, soil water content, soil bulk density, and soil organic carbon concentration allowed us to explain 54% of total variance of soil respiration over the different stand ages for the entire year (P < 0.05). Our results show that the adoption of an adequate sampling strategy, and the determination of some key environmental variables may help to explain a large proportion of total variation of soil respiration over the entire rotation length of afforested ecosystems.