Altitudinal gradients in stream fish diversity and the prevalence of diadromy in the Sixaola River basin,Costa Rica

Landscape-scale patterns of freshwater fish diversity and assemblage structure remain poorly documented in many areas of Central America, while aquatic ecosystems throughout the region have been impacted by habitat degradation and hydrologic alterations. Diadromous fishes may be especially vulnerable to these changes, but there is currently very little information available regarding their distribution and abundance in Central American river systems. We sampled small streams at 20 sites in the Sixaola River basin in southeastern Costa Rica to examine altitudinal variation in the diversity and species composition of stream fish assemblages, with a particular focus on diadromous species. A set of environmental variables was also measured in the study sites to evaluate how changes in fish assemblage structure were related to gradients in stream habitat. Overall, fish diversity and abundance declined steeply with increasing elevation, with very limited species turnover. The contribution of diadromous fishes to local species richness and abundance increased significantly with elevation, and diadromous species dominated assemblages at the highest elevation sites. Ordination of the sampling sites based on fish species composition generally arranged sites by elevation, but also showed some clustering based on geographic proximity. The dominant gradient in fish community structure was strongly correlated with an altitudinal habitat gradient identified through ordination of the environmental variables. The variation we observed in stream fish assemblages over relatively small spatial scales has significant conservation implications and highlights the ecological importance of longitudinal connectivity in Central American river systems.     

Environmental and spatial variables influence the composition of frog assemblages in sub-tropical eastern Australia

In community ecology, contrasting theories suggest that the distribution and abundance of species, and thus the composition of assemblages, are influenced by i) environmental gradients, or ii) contagious biotic processes such as predation, competition, dispersal and disease. In the former case, sites with similar environments would tend to support similar assemblages, while in the latter, geographically proximate sites would tend to support more similar assemblages than widely separated sites. I investigated the relative influence of environmental variables and spatial position on the composition of frog assemblages at forest streams in sub-tropical eastern Australia using redundancy analysis (RDA) and partial RDA. Data on the maximum abundance of the frog species at 65 survey sites were transformed such that RDA would yield the Hellinger distance between sites. The following analysis identified 11 environmental variables that explained 45% of the variation in the abundance of species at the survey sites (the species matrix), as a proportion of total variance. The geographic co-ordinates of the survey sites accounted for 12%, while the environmental and spatial variables combined accounted for 47% of the variation in the species matrix. Partial redundancy analysis indicated that of the explained variation, 74% was purely environmental, 5% was purely spatial and 21% was spatial environmental variation. This study is the first to quantify the relative influence of environmental and spatial variables on the composition of amphibian assemblages. It provides support for both the environmental control model and the biotic control model of species' distributions and assemblage composition, although environmental variables appear to have the greater effect at this scale of analysis.     

Spatial Variation in Fish Assemblages across small Mediterranean Drainages: Effects of Habitat and Landscape Context

Much debate about assemblage organization in stream fish may stem from analysing the effects of both local and large-scale processes on assemblage attributes over whole geographic regions. This study addresses this issue, by examining the contribution of local habitat attributes and landscape context to fish assemblage variation across small Mediterranean drainages in southern Portugal. Fish abundance and species composition was estimated in 28 sites, across 10 drainages, in both a dry year (1999) and a wet year (2001), and related to two sets of variables reflecting habitat and landscape characteristics. Fish showed responses to both sets of variables with variance partitioning indicating that landscape context had important effects on species richness whereas habitat attributes were the primary determinants of local fish abundance. In general, high species richness was associated with larger drainage area and higher rainfall variability, whereas variation in species abundances mostly reflected the influence of width, depth, conductivity, current velocity, substrate size and emergent vegetation. The relative contributions of both landscape context and habitat attributes to species richness and abundance were generally lower in 1999 than in 2001, with much less diversified species–habitat relationships being found in the former dry year. These results point to the dynamic nature of assemblage organization, emphasizing the importance of innovative, multi-scale approaches in advancing our knowledge of fish assemblage structure in Mediterranean streams.     

Specialization and habitat: spatial and environmental effects on abundance and genetic diversity of forest generalist and specialist Carabus species

Habitat specialist species are supposed to be more susceptible to variations in local environmental characteristics than generalists. To test this hypothesis, we conducted a comparative analysis on abundance and genetic diversity of forest carabids differing in their habitat requirements. Four species were sampled in forests characterized by abiotic, landscape and biotic environmental variables. A statistical framework based on canonical correspondence analysis was used for one habitat generalist and one habitat specialist species to determine the relative contribution of environmental variables in structuring inter- and intrapopulational genetic diversity depicted by microsatellites. Our results showed that sympatric species differed in their sensitivity to environmental variables. The same variables were found to be important in analyses of abundance and genetic data. However, specialization was not related to a greater sensitivity to local environmental characteristics. The strong impact of spatial variables on genetic data suggested that genetic variation among populations would largely reflect the response of individual species to dispersal opportunities more than the effect of habitat quality.     

Population structure on breeding grounds of Lake Malawi’s ‘twilight zone’ cichlid fishes

Aim Free‐ranging benthopelagic fishes often have large population sizes and high rates of dispersal. These traits can act to homogenize population structure across the distributional range of a species and to reduce the likelihood of allopatric speciation. The apparent absence of any barriers to gene flow among populations, together with prior molecular evidence for panmixia across the ranges of three species, has resulted in Diplotaxodon, a genus of benthopelagic cichlid fishes of Lake Malawi, being proposed as a candidate case of sympatric speciation. Our aim was to further investigate this possibility by testing for intraspecific genetic subdivision among breeding populations, and intraspecific differences in breeding habitat. Location Lake Malawi, central‐east Africa. Methods We analysed eight microsatellite DNA loci to test for spatial genetic differences among populations on breeding grounds of eight Diplotaxodon species. We also tested for temporal population genetic differences within breeding grounds of three species. Records of ripe Diplotaxodon encountered during sampling were analysed to test if spatial variation in assemblage structure was linked to nearshore water depth and geographic proximity of sampling sites. Results Consistent with previous molecular evidence, within four of the eight species tested we found no evidence of spatial genetic structuring among breeding populations. However, within the other four species we found slight yet significant spatial genetic differences, indicating restricted gene flow among breeding grounds. There was no evidence of temporal genetic differences within sites. Analyses of the distributions of ripe Diplotaxodon revealed differences in assemblage structure linked to nearshore water depth. Main conclusions Together, these results demonstrate both the evolution of fidelity to deep‐water breeding locations in some Diplotaxodon species, and differences in breeding habitat among species. These findings are consistent with a role for divergence of breeding habitat in speciation of these cichlids, possibly promoted by dispersal limitation among geographically segregated spawning aggregations.     

Consistent spatial patterns across biogeographic gradients in temperate reef fishes

Biogeographic gradients may facilitate divergent evolution between populations of the same species, leading to geographic variation and possibly reproductive isolation. Previous work has shown that New Zealand triplefin species (family Tripterygiidae) have diversified in habitat use, however, knowledge about the consistency of this pattern throughout their geographic range is lacking. Here we examine the spatial habitat associations of 15 New Zealand triplefin species at nine locations on a latitudinal gradient from 35°50'S to 46°70'S to establish whether distant populations differ in habitat use. Triplefin diversity and density varied between locations, as did habitat variables such as percentage cover of the substratum, onshore-offshore location, microposition, depth and exposure. Canonical discriminant analysis identified specific species-habitat combinations, and when habitat was statistically partialled from location, most species exhibited consistent habitat associations throughout their range. However, the density of a few species at some locations was lower or higher than expected given the habitat availability. This indicates that the habitat variables recorded were not the sole predictors of assemblage structure, and it is likely that factors influencing larval dispersal (e.g. the low salinity layer in Fiordland and geographic isolation of the Three Kings Islands) play an additional role in structuring assemblage composition. Together these results suggest that New Zealand triplefin species show strong and consistent habitat use across potential biogeographical barriers, but this pattern appears to be modified by variation in larval supply and survival. This indicates that species with broad geographic distributions do not necessarily show phenotypic variation between populations.     

Biome transitions as centres of diversity: habitat heterogeneity and diversity patterns of West African bat assemblages across spatial scales

It is widely accepted that species diversity is contingent upon the spatial scale used to analyze patterns and processes. Recent studies using coarse sampling grains over large extents have contributed much to our understanding of factors driving global diversity patterns. This advance is largely unmatched on the level of local to landscape scales despite being critical for our understanding of functional relationships across spatial scales. In our study on West African bat assemblages we employed a spatially explicit and nested design covering local to regional scales. Specifically, we analyzed diversity patterns in two contrasting, largely undisturbed landscapes, comprising a rainforest area and a forest‐savanna mosaic in Ivory Coast, West Africa. We employed additive partitioning, rarefaction, and species richness estimation to show that bat diversity increased significantly with habitat heterogeneity on the landscape scale through the effects of beta diversity. Within the extent of our study areas, habitat type rather than geographic distance explained assemblage composition across spatial scales. Null models showed structure of functional groups to be partly filtered on local scales through the effects of vegetation density while on the landscape scale both assemblages represented random draws from regional species pools. We present a mixture model that combines the effects of habitat heterogeneity and complexity on species richness along a biome transect, predicting a unimodal rather than a monotonic relationship with environmental variables related to water. The bat assemblages of our study by far exceed previous figures of species richness in Africa, and refute the notion of low species richness of Afrotropical bat assemblages, which appears to be based largely on sampling biases. Biome transitions should receive increased attention in conservation strategies aiming at the maintenance of ecological and evolutionary processes.     

Soil‐based habitat partitioning in understorey palms in lower montane tropical forests

Aim Dispersal assembly and niche assembly are two competing theories proposed to explain the maintenance of species diversity in tropical forests. Dispersal theory emphasizes the role of chance colonization events and distance‐limited seed dispersal in explaining species abundance and distribution, whereas niche theory emphasizes differences among species in requirements for potentially limiting resources. Species distribution patterns in tropical forests often correlate with geology and topography, but tests of the relative importance of dispersal and niche partitioning have been hampered by an inadequate characterization of resource availability. The aim of this study was to explore how soil chemical and physical properties, climate, and geographic distance affect understorey palm communities in lower montane forests. Location Fortuna Forest Reserve, Chiriqui Province, and Palo Seco Forest Reserve, Bocas del Toro Province, in western Panama. Methods Understorey palms and soil nutrient concentrations were surveyed within 10 sites on different soil types across a 13‐km transect. Variation in palm community composition was examined in relation to spatial and environmental variables. Results The 25 understorey palm species recorded in the study were non‐randomly distributed among forests differing in soil nutrient availability. In support of dispersal theory, floristic similarity decreased predictably with increasing geographic distance. However, environmental and soil variables were also correlated with geographic distance. Floristic similarity was also highly associated with a subset of environmental variables. Variation in palm community similarity was most strongly correlated with inorganic nitrogen availability and cation concentration. A subset of soil variables had a stronger relationship with floristic similarity when geographic distance was controlled for than did geographic distance when differences in soils were controlled for. Main conclusions Both dispersal and niche processes affect palm species distribution patterns. Although spatially limited dispersal may influence species distribution patterns, soil‐based habitat associations, particularly with respect to soil nitrogen, cation availability and aluminium concentrations, remain important factors influencing palm community composition at the mesoscale level in this tropical montane forest.     

Ecological biogeography of Mexican bats: the relative contributions of habitat heterogeneity,beta diversity,and environmental gradients to species richness and composition patterns

Mexico has higher mammalian diversity than expected for its size and geographic position. High environmental hetero geneity throughout Mexico is hypothesized to promote high turnover rates (β‐diversity), thus contributing more to observed species richness and composition than within‐habitat (α) diversity. This is true if species are strongly associated with their environments, such that changes in environmental attributes will result in changes in species composition. Also, greater heterogeneity in an area will result in greater species richness. This hypothesis has been deemed false for bats, as their ability to fly would reduce opportunities for habitat specialization. If so, we would expect no significant relationships between 1) species composition and environmental variables, 2) species richness and environmental heterogeneity, 3) β‐diversity and environmental heterogeneity. We tested these predictions using 31 bat assemblages distributed across Mexico. Using variance partitioning we evaluated the relative contribution of vegetation, climate, elevation, horizontal heterogeneity (a variate including vegetation, climate, and elevational heterogeneity), spatial variation (lat‐long), and vertical hetero geneity (of vegetation strata) to variation in bat species composition and richness. Variation in vegetation explained 92% of the variation in species composition and was correlated with all other variables examined, indicating that bats respond directly to habitat composition and structure. Beta‐diversity and vegetational heterogeneity were significantly correlated. Bat species richness was significantly correlated with vertical, but not horizontal, heterogeneity. Nonetheless, neither horizontal nor vertical heterogeneity were random; both were related to latitude and to elevation. Variation in bat community composition and richness in Mexico were primarily explained by local landscape heterogeneity and environmental factors. Significant relationships between β‐diversity and environmental variation reveal differences in habitat specialization by bats, and explain their high diversity in Mexico. Understanding mechanisms acting along environmental or geographic gradients is as important for understanding spatial variation in community composition as studying mechanisms that operate at local scales.     

Climate and geographic distance are more influential than rivers on the beta diversity of passerine birds in Amazonia

Variation in the spatial structure of communities in terms of species composition (beta diversity) is affected by different ecological processes, such as environmental filtering and dispersal limitation. Large rivers are known as barriers for species dispersal (riverine hypothesis) in tropical regions. However, when organisms are not dispersal limited by geographic barriers, other factors, such as climatic conditions and geographic distance per se, may affect species distribution. In order to investigate the relative contribution of major rivers, climate and geographic distance on Passeriformes beta diversity, we divided Amazonia into 549 grid cells (1° of latitude and longitude) and obtained data of species occurrence, climate and geographic position for each cell. Beta diversity was measured using taxonomic, phylogenetic and functional metrics of composition. The influence of climatic variables, geographic distance and rivers on these metrics was tested using regression analyses. Passerine beta diversity is characterized mainly by the change in species taxonomic identity and in phylogenetic lineages across climatic gradients and over geographic distance. However, species with similar traits are found throughout the entire Amazonia. The size of rivers was proportional to their effect on species composition. However, climate and geographic distance are relatively more important than rivers for Amazonian taxonomic and phylogenetic species composition.     

地理距离及环境差异对云南元江干热河谷植物群落beta多样性的影响

beta多样性反映了群落间物种组成的差异, 是生物多样性研究的热点之一。本研究通过对云南元江干热河谷41个植物群落样方进行调查, 用Jaccard相异系数表征物种beta多样性, 用样方之间的最近谱系距离(mean nearest taxon distance, MNTD)及平均谱系距离(mean pairwise distance, MPD)表征谱系beta多样性, 采用基于距离矩阵的多元回归和方差分解方法, 探讨了该区域干热河谷典型植物群落的物种beta多样性和谱系beta多样性与样方间环境差异(主要是气候)及地理距离之间的关系。结果表明: (1)群落间的地理距离和年平均温度差异对干热河谷植物群落的物种beta多样性和谱系beta多样性有显著影响; (2)地理距离对物种beta多样性和MNTD的影响最大; 地理距离和年平均温度差异对MPD的影响均较大; (3)样方间年平均温度与年平均降水量的差异和地理距离能够解释群落间beta多样性及谱系beta多样性11-13%的变异。以上结果表明, 生态位分化和扩散限制对该地区植物群落的beta多样性均有显著影响, 其中扩散限制的影响可能更大。此外, 人类活动等其他因素也很可能对元江干热河谷的群落组成具有非常重要的影响。     

Isolation by distance,not rivers,control the distribution of termite species in the Amazonian rain forest

The spatial distribution of species is affected by dispersal barriers, local environmental conditions and climate. However, the effect of species dispersal and their adaptation to the environment across geographic scales is poorly understood. To investigate the distribution of species from local to broad geographic scales, we sampled termites in 198 transects distributed in 13 sampling grids in the Brazilian Amazonian forest. The sampling grids encompassed an area of 271 500 km² and included the five major biogeographic regions delimited by Amazonian rivers. Environmental data for each transect were obtained from local measurements and remote sensing. Similar to previous studies, termite species composition at the local scale was mostly associated with measures of soil texture and chemistry. In contrast, termite species composition at broad geographic scales was associated with soil nutrients, and the geographic position of the transects. Between 17 and 30% of the variance in termite species composition could be attributed exclusively to the geographic position of the transects, but could not be attributed to measured environmental variables or the presence of major rivers. Isolation by distance may have strong effects on termite species composition due to historic processes and the spatially structured environments along distinct geological formations of Amazonia. However, in contrast to many taxa in Amazonia, there is no evidence that major rivers are important barriers to termite dispersal.     

Drivers of the beta diversity of spider assemblages in southern Brazilian temporary wetlands

1. The composition of local assemblages is assembled by an interplay of species sorting, mass effects and dispersal limitation processes. The contributions of assembly processes to metacommunity structure can change with ecosystem type and specificities of the study area. Spider composition is influenced by environmental features such as habitat structure and climate, and also by spatial distances between patches. However, little is known about the roles of assembly processes in spider metacommunity structure in wetlands. 2. The beta diversity patterns of spider assemblages were assessed in 24 temporary wetlands distributed along a latitudinal gradient in southern Brazil. The study also assessed the individual correspondence of beta diversity (and its turnover and richness components) with dissimilarities in habitat structure and climate, as well as with geographic distances, using Mantel and partial Mantel correlation tests and multivariate correlograms. 3. Turnover was the most important component of spider beta diversity. Mantel tests detected significant correlations of spider beta diversity with habitat structure. Partial Mantel tests detected significant relationships only between spider beta diversity (and the richness component) and geographic distances. Additionally, spider composition was more similar than chance on smaller scales. 4. These results evidenced a complex interplay of assembly processes explaining spider metacommunity structure in temporary wetlands. Although species-sorting processes associated with habitat structure were important in structuring local spider composition, mass effects and dispersal limitation across climatic zones played an important role on a broader scale.     

Spatial variation in insect community and species responses to habitat loss and plant community composition

Several experimental studies have examined species responses to manipulations of habitat area and spatial arrangement, but plant composition and spatial variation in species distributions also affect animal responses to habitat alteration. We used an experimental approach to study the combined effects of habitat area, edge, and plant community composition on the spatial structure of insect species richness and composition. The abundance of three guilds (herbivores, predators and parasitoids) and individual species were also analyzed. Habitat patches were created that differed in area and edge by selectively mowing portions of 15 m×15 m plots in a 1.7-ha old field. Spatial and environmental variables were used to predict insect responses in separate multiple regression and ordination models. The variation in species responses due to spatial and environmental variables was then partitioned by combining these variables into an overall regression or ordination. Spatial and environmental variables contributed similar percentages to the total variance in insect species richness, abundance or composition. No significant effects of habitat area were observed in any response variable. Herbivore abundance showed positive responses to legume or grass cover, as well as spatial variation that was unrelated to environmental variables. Predators and parasitoids had greater effects of plant species richness and habitat edge, and less unexplained spatial variation. Individual species differed in their responses to plant variables, depending on host specialization or intraspecific aggregation. Our study highlights the importance of plant community composition and spatial variation apart from environmental variables. Spatial variation stems both from species responses to environmental features as well as species differences in habitat specialization and intraspecific aggregation.     

阿拉善荒漠啮齿动物集合群落实证研究

当生态学家探求在破碎化的栖息地中,群落物种的共存机制、多样性、局域尺度的性质和过程被放到更广阔的时空框架内时,就出现了"集合群落"这一概念。Leibold提出了集合群落概念,他们将一个集合群落定义为局域群落集,这些群落由各个潜在的相互作用的物种的扩散连接在一起。集合群落理论描述了那些发生在集合群落尺度上的过程,并且提出思考关于物种相互作用的新方法。集合群落概念为群落生态学提供了一个新的革命性的范式,集合群落研究的最基本问题是同一系统中多物种共存的机理、多样性的形成原因与维持机制。该范式强调区域范围内群落中的综合变异,强调环境特证和栖息地之间通过扩散调节的生物相互作用和空间变化。Leibold等提出了解释集合群落结果理论上的4个生态范式,即(1)中性理论;(2)斑块动态理论;(3)物种分配理论;(4)集团效应理论。之后有大量有关检验这4种生态理论的研究,但是有关陆地脊椎动物系统的集合群落的研究较少。2010—2012年,通过在内蒙古阿拉善荒漠景观中的8个固定样地中,对啮齿动物、栖息地环境因子进行调查。利用冗余分析和偏冗余分析,评估环境特征和空间特征对物种组成的影响。结果表明,环境特征独自解释72.8%的啮齿动物物种组成变化,空间特征独自解释33.8%的物种组成变化,环境特征和空间特征共同解释86.5%的啮齿动物物种组成变化,结果显著(P=0.032);去除环境特征之后,空间特征解释13.7%的变化(P=0.246),结果不显著;去除空间特征之后,栖息地变化解释52.7%的变化(P=0.016);环境特征和空间特征的交互作用解释20.1%的物种组成的变化,该区域啮齿动物群落构成集合群落,物种共存中环境特征起着主导作用,由物种分配理论解释该集合群落结构。     

Are Assemblages of Aquatic‐Breeding Anurans (Amphibia) Niches Structured or Neutral?

Local niche‐based processes and dispersal are important determinants of assemblage composition and species diversity. However, there is no consensus about the relative importance of niche and spatial processes to explain the distribution of anuran species in tropical systems. In our study, we analyzed the niche and neutral effects on anuran assemblages and found that biotic interactions were a predictor of assemblage structure. The Eltonian concept of niche was the best predictor for the structure of aquatic‐breeding anuran assemblages, as species tended to co‐occur more often than would be expected by chance. We suggest that the lack of environmental effect could be explained by differences in the pattern of movement between arboreal and non‐arboreal anurans. Once there is a reduction in the number of arboreal anurans in open areas, the importance of habitat heterogeneity to explain assemblage composition should decrease. The lack of correlation between the spatial component in our model and species composition is evidence that spatial processes, such as migration, did not play a major role in structuring local assemblages. Anurans are generally assumed as having poor dispersal ability, yet this assumption is not true for all anuran species. We suggest that future studies should include key behavioral traits, such as site fidelity and homing behavior, as these traits can represent the dispersal abilities of anurans and dispersal ability seems to be important when we try to predict patterns of anuran distribution.     

Habitat distribution models,spatial autocorrelation,functional traits and dispersal capacity of alpine plant species

Abstract. We evaluate the potential influence of disturbance on the predictability of alpine plant species distribution from equilibrium‐based habitat distribution models. Firstly, abundance data of 71 plant species were correlated with a comprehensive set of environmental variables using ordinal regression models. Subsequently, the residual spatial autocorrelation (at distances of 40 to 320 m) in these models was explored. The additional amount of variance explained by spatial structuring was compared with a set of functional traits assumed to confer advantages in disturbed or undisturbed habitats. We found significant residual spatial autocorrelation in the habitat models of most of the species that were analysed. The amount of this autocorrelation was positively correlated with the dispersal capacity of the species, levelling off with increasing spatial scale. Both trends indicate that dispersal and colonization processes, whose frequency is enhanced by disturbance, influence the distribution of many alpine plant species. Since habitat distribution models commonly ignore such spatial processes they miss an important driver of local‐ to landscape‐scale plant distribution.     

黄土高原腹地人工林下草本层群落构建机制

群落构建机制是生态学研究的核心论题。生态位理论和中性理论是阐明群落构建的两种主要理论, 但这两种理论分别强调的环境过滤和扩散限制对群落构建的影响尚存争议。该研究以黄土高原人工林下草本层群落为研究对象, 将群落物种组成数据与地理空间、气候、地形以及生物因子相结合, 运用随机森林模型(RF)和基于距离矩阵的多元回归方法(MRM), 探究了各个因子的重要性, 并通过邻体主坐标矩阵(PCNM)和基于距离的冗余分析(db-RDA)分别筛选显著的空间结构和环境因子, 最后结合筛选出的PCNM特征值和环境变量进行变差分解。研究结果显示: 1)林下草本层群落组成相似性随地理距离和生境差异的增加而减小。2)地理距离与生境差异共同解释了群落组成相似性变化的47.8%。其中, 空间因子和环境因子的独立解释率分别为14.1%和9.8%, 两者的联合解释率为23.9%。3)地理距离和年降水量是引起研究区内物种组成变化的关键因子, 且地理距离的重要性大于年降水量。在黄土高原腹地, 扩散限制与环境过滤共同主导了人工林下草本层群落的构建过程。     

Habitat filtering and adult dispersal determine the taxonomic composition of stream insects in an urbanizing landscape

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