Patterns in the co-occurrence of fish species in streams: the role of site suitability,morphology and phylogeny versus species interactions

A number of studies at large scales have pointed out that abiotic factors and recolonization dynamics appear to be more important than biotic interactions in structuring stream-fish assemblages. In contrast, experimental and field studies at small scales show the importance of competition among stream fishes. However, given the highly variable nature of stream systems over time, competition may not be intense enough to generate large-scale complementary distributions via competitive exclusion. Complementary distribution is a recurrent pattern observed in fish communities across stream gradients, though it is not clear which instances of this pattern are due to competitive interactions and which to individual species requirements. In this study, I introduce a series of null models developed to provide a more robust evaluation of species associations by facilitating the distinction between different processes that may shape species distributions and community assembly. These null models were applied to test whether conspicuous patterns in species co-occurrences are more consistent with their differences in habitat use, morphological features and/or phylogenetic constraints, or with species interactions in fish communities in the streams of a watershed in eastern Brazil. I concluded that patterns in species co-occurrences within the studied system are driven by common species-habitat relationships and species interactions may not play a significant role in structuring these communities. I suggest that large-scale studies, where adequate designs and robust analytical tools are applied, can contribute substantially to understanding the importance of different types of processes in structuring stream-fish communities.     

The resource balance hypothesis of plant species diversity in grassland

Abstract: We hypothesize that plant species diversity is favoured when actual resource supply ratios are balanced according to the optimum resource supply ratios for the vegetation as a whole. This ‘resource balance hypothesis of plant species diversity’ (RBH) follows from two different mechanisms of plant species coexistence, namely: ‘differential resource limitation’, which allows species to coexist in a competitive equilibrium in a homogeneous environment and ‘micro-habitat differentiation’, which builds on spatial heterogeneity. Both mechanisms require that resource supply ratios are intermediate between the optimum supply ratios of the species present in the species pool. Additional conditions, concerning the resource acquisition and requirement ratios of the species, are easier to meet for the second mechanism than for the first. To test the RBH we measured species diversity parameters in 74 grassland plots, as well as the N, P and K concentrations in the above-ground biomass. We used a new ceiling detection algorithm to examine the relationship between maximum observed diversity and the N/P-, P/K- and K/N-ratios in the biomass. Most of these ceiling relationsips could be described by parabolic curves with significant quadratic terms. This indicates that high diversity does not occur at the extremes of the observed ranges of nutrient ratios. This supports the RBH.     

Experimental effects of habitat fragmentation on old-field canopy insects: community, guild and species responses

We examined the effects of habitat fragmentation on the species distributions, guild membership, and community structure of old-field insects using a fine-scale experimental approach. A continuous 1-ha goldenrod field was fragmented into four treatments that varied in both patch size and degree of isolation. Each treatment was replicated four times and arranged in a Latin square design. Canopy insects in fragmented patches were sampled with sweep nets during early and late summer 1995. The species richness of insects was significantly lower in fragmented than in unfragmented treatments during July, but was similar among treatments in September. Overall community abundance showed no treatment effect during either month. We also found significant row and column effects, suggesting there was spatial heterogeneity in species richness and abundance apart from treatment effects. Differences in species richness during July were primarily due to the loss of rare species in highly fragmented plots. Overall abundance was less responsive to community change because deletions of rare species in fragmented areas were not detected in abundance analyses. Four feeding guilds showed different responses to fragmentation: the species richness of sucking herbivores and the abundance of parasitoids were significantly reduced by fragmentation but predators and chewing herbivores were largely unaffected. Analyses of a subset of individual species within guilds suggest that the greater effects of fragmentation on sucking herbivores and parasitoids may be due to the degree of habitat specificity of guild members. The effects of small-scale habitat fragmentation were therefore detectable at the level of community, guild, and individual species. Changes in species richness, guild structure and species distributions were likely due to differential effects of habitat alteration on individual movements and patch selection rather than dispersal or demographic change. Nonetheless, the selective loss of rare species, differential guild effects and changes in species occupancy that we found in this small-scale experiment are also factors that are likely to operate in fragmented habitats over broader spatial scales. Received: 11 May 1998 / Accepted: 27 September 1998     

The trade-off between dispersability and longevity - an important aspect of plant species diversity

Abstract. Local presence of plant species is determined by population colonizations and extinctions. All traits that influence the capacity of individuals to colonize patches and survive within patches, are therefore important for community diversity. Spatial models can explain the coexistence of species provided that the inferior competitor has a greater spatial mobility and thereby can avoid competition. We searched the literature for empirical evidence for such trade-offs and included all available information on correlations between traits associated with the capacity to colonize and traits promoting the ability to survive. A lower reproductive effort of a species is associated with a longer life span and a higher competitive ability. Morphological adaptations for dispersal are less common in species which better tolerate stress, that are better competitors or possess seed dormancy. Such patterns suggest that species that are good survivors may have a limited ability to colonize new patches and vice versa. A negative correlation between dispersability and longevity has important effects on the regional dynamics of single species as well as on the coexistence of species. From a conservation perspective differences in the colonization capacity among species imply that restoration of plant biodiversity must not only focus on conditions within patches, but also consider the spatial arrangement of patches in order to enable plants to bridge gaps in time and space.     

Changes in plant species diversity along a chronosequence of vegetation restoration in the humid evergreen broad-leaved forest in the Rainy Zone of West China

Plant species diversity has been recognized as one of the vital attributes for assessing vegetation restoration. Changes in the diversity may be related to different stages of succession. In this study, 54 sites of humid, evergreen, broad-leaved forest were selected in the Rainy Zone of West China. A chronosequence of the sites was used to study the successive patterns of the diversity in the forest that had undergone natural regeneration for 5 to 350 years and to test the hypothesis that the diversity is maximized in mid-succession. Data were collected simultaneously at different stages of succession, and four α-diversity indices (species richness, Margalef index, Shannon-Wiener index, Pielou Evenness index) and two β-diversity indices (Whittaker index, Sørensen’s index) were calculated for each stratum in each plot. A total of 394 vascular plant species were recorded. From the β-diversity indices, the forest succession may be divided into the early-successional stage (before 50 years), mid-successional stage (from 50 to 300 years), and late-successional stage (after 300 years). In this community, the species diversity and richness were found to be the greatest at the mid-successional stage, followed by the late- and early-successional stages. The results of regression analysis indicated that the richness and Margalef index peaked around the 175th and 165th year, respectively. Shannon-Wiener index values also appeared to follow an approximately humped pattern of succession and were maximal around the 100th year. However, the species evenness did not show any significant relationship with successional age. Our results demonstrate (1) forest restoration is a long-term process and the formation of climax forest requires at least 300 years and (2) the forest has a strong capacity for restoration. Our results also suggest Lindera limprichitii and Machilus pingii as ideal tree species for afforestation because of their wide niche.     

Diversity of forest plant species at the community and landscape scales in Switzerland

Abstract

Conservation strategies increasingly refer to indicators derived from large biological data. However, such data are often unique with respect to scale and species groups considered. To compare richness patterns emerging from different inventories, we analysed forest species richness at both the landscape and the community scales in Switzerland. Numbers of forest species were displayed using nationwide distributional species data and referring to three different definitions of forest species. The best regression models on a level of four predictor variables ranged between adj. R ² = 0.50 and 0.66 and revealed environmental heterogeneity/energy, substrate (rocky outcrops) and precipitation as best explanatory variables of forest species richness at the landscape scale. A systematic sample of community data (n = 729; 30 m², 200 m², 500 m²) was examined with respect to nationwide community diversity and plot species richness. More than 50% of all plots were assigned to beech forests (Eu-Fagion, Cephalanthero-Fagion, Luzulo-Fagion and Abieti-Fagion), 14% to Norway spruce forests (Vaccinio-Piceion) and 13% to silver fir forests (Piceo-Abietion). Explanatory variables were derived from averaged indicator values per plot, and from biophysical and disturbance factors. The best models for plot species richness using four predictor variables ranged between adj. R ² = 0.31 and 0.34. Light (averaged L-indicator, tree canopy) and substrate (averaged R-indicator and pH) had the highest explanatory power at all community scales. By contrast, the influence of disturbance variables was very small, as only a small portion of plots were affected by this factor. The effects of disturbances caused by extreme events or by management would reduce the tree canopies and lead to an increase in plant species richness at the community scale. Nevertheless, such community scale processes will not change the species richness at the landscape scale. Instead, the variety of different results derived from different biological data confirms the diversity of aspects to consider. Therefore, conservation strategies should refer to value systems.     

Effects of river level fluctuation on plant species richness, diversity, and distribution in a floodplain forest in Central Amazonia

River levels in Central Amazonia fluctuate up to 14 m annually, with the flooding period ranging from 50 to 270 days between the rising and falling phases. Vast areas of forest along the rivers contain plant species that are well adapted to annual flooding. We studied the effect of flooding level on tree species richness, diversity, density, and composition in lake, river, and stream habitats in Jaú National Park, Brazil. 3051 trees >10 cm diameter (at 1.3 m diameter at breast height, dbh) were measured and identified in 25 10 m × 40 m randomly selected plots in each habitat. Ordination methods and analysis of variance results showed that forested areas near lakes had significantly lower species richness of trees than riverine and streamside habitats. Plot species richness and diversity were strongly negatively correlated with the water level and duration of flooding. The drier (stream) habitat had more total species (54 species of trees) and more unique species of trees (6 tree species) than the riverine (52 tree species; 3 unique species) and lake (33 tree species; 3 unique species) habitats. Species composition overlap among habitats was surprisingly high (42.6–60.6% overlap), almost one-third of the species were found in all three habitat types, and few species were unique to each habitat. We conclude that: (1) duration of flooding has a strong impact on species richness, diversity and plant distribution patterns; (2) most species are adapted to a wide range of habitats and flood durations; and (3) while flood duration may decrease local diversity, it also creates and maintains high landscape-scale diversity by increasing landscape heterogeneity. Received: 20 April 1997 / Accepted: 14 January 1999     

Plant species diversity and environmental heterogeneity: spatial scale and competing hypotheses

Questions: What is the observed relationship between plant species diversity and spatial environmental heterogeneity? Does the relationship scale predictably with sample plot size? What are the relative contributions to diversity patterns of variables linked to productivity or available energy compared to those corresponding to spatial heterogeneity? Methods: Observational and experimental studies that quantified relationships between plant species richness and within‐sample spatial environmental heterogeneity were reviewed. Effect size in experimental studies was quantified as the standardized mean difference between control (homogeneous) and heterogeneous treatments. For observational studies, effect sizes in individual studies were examined graphically across a gradient of plot size (focal scale). Relative contributions of variables representing spatial heterogeneity were compared to those representing available energy using a response ratio. Results: Forty‐one observational and 11 experimental studies quantified plant species diversity and spatial environmental heterogeneity. Observational studies reported positive species diversity‐spatial heterogeneity correlations at all points across a plot size gradient from ～1.0 × 10^?1 to ～1.0 × 10¹¹ m², although many studies reported spatial heterogeneity variables with no significant relationships to species diversity. The cross‐study effect size in experimental studies was not significantly different from zero. Available energy variables explained consistently more of the variance in species richness than spatial heterogeneity variables, especially at the smallest and largest plot sizes. Main conclusions: Species diversity was not related to spatial heterogeneity in a way predictable by plot size. Positive heterogeneity‐diversity relationships were common, confirming the importance of niche differentiation in species diversity patterns, but future studies examining a range of spatial scales in the same system are required to determine the role of dispersal and available energy in these patterns.     

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.     

A multiple-site dissimilarity measure for species presence/absence data and its relationship with nestedness and turnover

Multiple-site dissimilarity may be caused by two opposite processes of meta-community organization, such as species nestedness and turnover. Therefore, discriminating among these contributions is necessary for linking multiple-site dissimilarity to ecosystem functioning. This paper introduces a measure of multiple-site dissimilarity or beta diversity for presence/absence data that is based on information on species absences from the species × sites matrix. It is also shown that the newly proposed dissimilarity index can be additively partitioned into species nestedness and turnover.     

Multi-scale analysis on species diversity within a 40-ha old-growth temperate forest

In order to better explore the maintenance mechanisms of biodiversity,data collected from a 40-ha undisturbed Pinus forest were applied to the Individual SpecieseArea Relationship model (ISAR) to determine distribution patterns for species richness.The ecological processes influencing species abundance distribution patterns were assessed by applying the same data set to five models:a LogNormal Model (LNM),a Broken Stick Model (BSM),a Zipf Model (ZM),a Niche Preemption Model (NPM),and a Neutral Model (NM).Each of the five models was used at six different sampling scales (10 m×10 m,20 m×20 m,40 m×40 m,60 m×60 m,80 m×80 m,and 100 m×100 m).Model outputs showed that:(1) Accumulators and neutral species strongly influenced species diversity,but the relative importance of the two types of species varied across spatial scales.(2) Distribution patterns of species abundance were best explained by the NPM at small scales (10 me20 m),whereas the NM was the best fit model at large spatial scales.(3) Species richness and abundance distribution patterns appeared to be driven by similar ecological processes.At small scales,the niche theory could be applied to describe species richness and abundance,while at larger scales the neutral theory was more applicable.     

High shoot plasticity favours plant coexistence in herbaceous vegetation

Several theoretical considerations imply that high shoot morphological plasticity could increase competition symmetry and favour plant coexistence. We tested whether mean plasticity across co-occurring species is a key trait for explaining ramet density and species richness in herbaceous vegetation. We used three data sets to test the hypotheses: (a) experimentally achieved estimates of plasticity to light availability for 35 herbaceous species; (b) richness, ramet density and canopy architecture data from 17 herbaceous communities; (c) species richness data from a 5-year permanent-plot study in a calcareous grassland. In herbaceous communities containing species with relatively higher shoot plasticity, ramet density was significantly higher. Consequently, relatively more species were growing per unit area—a greater proportion of the community species pool was represented on 1 m². In the permanent plot study species-richness was higher in those 40×40 cm quadrats where species with high shoot plasticity prevailed—there was a positive regression of richness on the mean plasticity of species. This relationship was highly significant in five consecutive years. Our results suggest that shoot plasticity to light availability is evidently one of the key traits in processes that alter the density of co-existing plants and, therefore, species diversity in herbaceous communities. Electronic Supplementary Material Supplementary material is available for this article at     

Does diversity beget diversity? A case study of crops and weeds

Abstract. The ecological literature is ambiguous as to whether the initial diversity of a plant community facilitates or deters the diversity of colonizing species. We experimentally planted annual crop species in monoculture and polyculture, and examined the resulting weed communities. The species composition of weeds was similar among treatments, but the species richness of weeds was significantly higher in the polycultures than in the monocultures. This supports the ‘diversity begets diversity’ hypothesis. Environmental microheterogeneity, diversity promoters, and ecological equivalency do not seem able to explain the observed patterns.     

Enhancement of habitat heterogeneity and species richness on rocky shores inundated by sand

Summary Many rocky shores are subject to periodic inundation by sand, which is often thought to reduce species richness by eliminating organisms intolerant of sand scour or sand smothering. However, regular disturbance (e.g. inundation) should promote richness by preventing the development of low diversity climax communities. A study of faunal richness on 10 regularly inundated shores showed that inundation does promote richness, but by increasing habitat heterogeneity. Some species are excluded from parts of the shore by sand, but because of the patchiness of sand deposits they are rarely excluded from the entire shore. Other species are found only on rocks associated with sand, while typically sandy shore animals occur in the sand deposits themselves. Total richness (281 species) was greater than for local noninundated shores and sandy beaches combined.     

A general test of the interactive-isolationist continuum in gastrointestinal parasite communities of fish

Parasite communities are generally believed to lie somewhere along the interactive-to-isolationist continuum, i.e. from rich assemblages of species with high colonisation rates in which interspecific interactions play an important structuring role, to species-poor assemblages where interactions are unlikely. This framework has become one of the paradigms of parasite community ecology. There is, however, no objective way of ranking a set of parasite communities in terms of the extent of interactivity among their constituent species. Here, we propose a simple index of interactivity based on the general likelihood of species co-occurrence, and thus on the potential for interactions, and we apply it to component communities of gastrointestinal helminth parasites from 37 species of marine fish hosts. The index essentially collapses several features of parasite communities thought to influence the degree of interactivity into a single number independent of the number of hosts examined or the total number of species in a component community. The range of values obtained here suggests that the potential interactivity in helminth communities of fish covers almost the full spectrum of possibilities, i.e. from isolationist to highly interactive communities. Although derived from presence/absence data only, the index correlates relatively strongly with the total parasite abundance per host, as well as the total prevalence of infection and the mean infracommunity richness. In other words, it captures properties of the community that influence interactivity. The use of the index in comparative studies may help in determining whether interactive helminth communities are, as widely believed, more common in endothermic vertebrate hosts than in fish hosts.     

Constructive Species’ Coexistence Mechanisms of Pine Oak Mixed Forest in Qinling Mountains Based on the Niche Differentiation

Based on the 15 plots of Pinus tabuliformis and Quercus aliena var. accuteserrata mixed forest in Qinling Mountains, we analyzed this community’s characteristics from the aspects of species composition, niche width and diameter class, also, we used the Canonical Correspondence Analysis to explore the effects of environment on plants’ distributions and conductive species’ morphological characteristics, the main results were as follows: (1) Fifteen plots record 139 vascular plants, belonging to 53 families 102 genera, the families which possessing higher richness were Rosaceae, Asteraceae, Liliaceae and Poaceae; (2) The niche widths of Ptabuliformis (Bi=2674) and Q var. accuteserrata (Bi=2679) were all higher than other accompanying plants, and there was little niche overlap value (< 01) between them; (3) Organic matter, total nitrogen, elevation and slope mainly effected plant’s distributions; (4) Four morphological characteristics of constructive species in CCA diagram presented a opposite distribution patterns, while elevation, slope position and available nitrogen were the main induced factors; (5) Community stability of this pine oak mixed forest was relatively higher based on Godron index, and two conductive species’ diameter class were all growing type. In conclusion, high habitat heterogeneity of terrain and soil nutrient brought niche differentiation and promoted species coexistence of pine oak mixed forest in Qinling Mountains.     

Macroecological patterns of spider species richness across Europe

We analysed the pattern of covariation of European spider species richness with various environmental variables at different scales. Four layers of perception ranging from single investigation sites to the whole European continent were selected. Species richness was determined using published data from all four scales. Correlation analyses and stepwise multiple linear regression were used to relate richness to topographic, climatic and biotic variables. Up to nine environmental variables were included in the analyses (area, latitude, elevation range, mean annual temperature, local variation in mean annual temperature, mean annual precipitation, mean July temperature, local variation in mean July temperature, plant species richness). At the local and at the continental scale, no significant correlations with surface area were found, whereas at the landscape and regional scale, surface area had a significant positive effect on species richness. Factors that were positively correlated with species richness at both broader scales were plant species richness, elevation range, and specific temperature variables (regional scale: local variation in mean annual, and mean July temperature; continental scale: mean July temperature). Latitude was significantly negatively correlated with the species richness at the continental scale. Multiple models for spider species richness data accounted for up to 77% of the total variance in spider species richness data. Furthermore, multiple models explained variation in plant species richness up to 79% through the variables mean July temperature and elevation range. We conclude that these first continental wide analyses grasp the overall pattern in spider species richness of Europe quite well, although some of the observed patterns are not directly causal. Climatic variables are expected to be among the most important direct factors, although other variables (e.g. elevation range, plant species richness) are important (surrogate) correlates of spider species richness.