Biodiversity, productivity and the temporal stability of productivity: patterns and processes

Theory predicts that the temporal stability of productivity, measured as the ratio of the mean to the standard deviation of community biomass, increases with species richness and evenness. We used experimental species mixtures of grassland plants to test this hypothesis and identified the mechanisms involved. Additionally, we tested whether biodiversity, productivity and temporal stability were similarly influenced by particular types of species interactions. We found that productivity was less variable among years in plots planted with more species. Temporal stability did not depend on whether the species were planted equally abundant (high evenness) or not (realistically low evenness). Greater richness increased temporal stability by increasing overyielding, asynchrony of species fluctuations and statistical averaging. Species interactions that favoured unproductive species increased both biodiversity and temporal stability. Species interactions that resulted in niche partitioning or facilitation increased both productivity and temporal stability. Thus, species interactions can promote biodiversity and ecosystem services.     

Resource stoichiometry and consumers control the biodiversity-productivity relationship in pelagic metacommunities

Recent theory suggests that both biodiversity and productivity are constrained by resource supply rates and ratios and that resource stoichiometry is the key to understanding the relationship between biodiversity and productivity. We experimentally tested this theory using pelagic metacommunities. We amended existing predictions by explicitly considering evenness as an aspect of biodiversity and including control of algal biomass by consumption in addition to competition. The metacommunities received a different phosphorus (P) supply and the three patches within each metacommunity differed in their nitrogen (N) supply, which created different N∶P ratios (2, 16, and 128). All patches were inoculated with a phytoplankton assemblage consisting of five species, and half of the metacommunities received two ciliate species as consumers. At the level of the entire metacommunity, algal biomass increased with increasing P supply, whereas species richness and evenness decreased with increasing P supply. Without consumers, resource use efficiency (RUE; realized biomass per unit of P) increased with increasing richness and evenness. Consumer presence reduced overall biomass and richness and precluded a correlation between RUE and biodiversity. At the patch level, local evenness correlated with higher RUE at both imbalanced N∶P ratios (2 and 128) but not at a balanced N∶P ratio. In conclusion, overall P supply constrained realized biomass and altered diversity, whereas resource stoichiometry shaped the relationship between biodiversity and RUE.     

Consistent Effects of Biodiversity on Ecosystem Functioning Under Varying Density and Evenness

Biodiversity experiments typically vary only species richness and composition, yet the generality of their results relies on consistent effects of these factors even under varying starting conditions of density and evenness. We tested this assumption in a factorial species richness x density x evenness experiment using a pool of 60 common grassland species divided into four functional groups (grasses, legumes, tall herbs and short herbs). Richness varied from 1, 2, 4, 8 to 16 species, total planting density was 1,000 or 2,000 seeds/m², and species were sown in even or uneven proportions, where one functional group was made dominant. Aboveground plant biomass increased linearly with the logarithm of species richness in all density and evenness treatments during all three years of the experiment. This was due to a convergence of realized density and evenness within species richness levels, although functional groups which were initially made dominant retained their dominance. Between species richness levels, realized density increased, and realized evenness decreased with species richness. Thus, more individuals could coexist if they belonged to different species. Within species richness levels, higher biomass values were correlated with lower density, suggesting an underlying thinning process. However, communities with low realized evenness also had low biomass values; thus high biomass did not result from species dominance. So-called complementarity and selection effects were similar across density and evenness treatments, indicating that the mechanisms underpinning the biodiversity effects were not altered. Species richness was the dominant driver of aboveground biomass, irrespective of variations in total densities and species abundance distributions at the start of the experiment; rejecting the hypothesis that initial differences in species abundance distributions might lead to different “stable states” in community structure or biomass. Thus, results from previous biodiversity experiments that only manipulated species richness and composition should be quite robust and broadly generalizable.     

Across species‐pool aggregation alters grassland productivity and diversity

Plant performance is determined by the balance of intra‐ and interspecific neighbors within an individual's zone of influence. If individuals interact over smaller scales than the scales at which communities are measured, then altering neighborhood interactions may fundamentally affect community responses. These interactions can be altered by changing the number (species richness), abundances (species evenness), and positions (species pattern) of the resident plant species, and we aimed to test whether aggregating species at planting would alter effects of species richness and evenness on biomass production at a common scale of observation in grasslands. We varied plant species richness (2, 4, or 8 species and monocultures), evenness (0.64, 0.8, or 1.0), and pattern (planted randomly or aggregated in groups of four individuals) within 1 × 1 m plots established with transplants from a pool of 16 tallgrass prairie species and assessed plot‐scale biomass production and diversity over the first three growing seasons. As expected, more species‐rich plots produced more biomass by the end of the third growing season, an effect associated with a shift from selection to complementarity effects over time. Aggregating conspecifics at a 0.25‐m scale marginally reduced biomass production across all treatments and increased diversity in the most even plots, but did not alter biodiversity effects or richness–productivity relationships. Results support the hypothesis that fine‐scale species aggregation affects diversity by promoting species coexistence in this system. However, results indicate that inherent changes in species neighborhood relationships along grassland diversity gradients may only minimally affect community (meter) – scale responses among similarly designed biodiversity–ecosystem function studies. Given that species varied in their responses to local aggregation, it may be possible to use such species‐specific results to spatially design larger‐scale grassland communities to achieve desired diversity and productivity responses.     

Influence of Plant Community Composition on Biomass Production in Planted Grasslands

United States energy policy mandates increased use of renewable fuels. Restoring grasslands could contribute to a portion of this requirement through biomass harvest for bioenergy use. We investigated which plant community characteristics are associated with differences in biomass yield from a range of realistic native prairie plantings (n = 11; i.e., conservation planting, restoration, and wildlife cover). Our primary goal was to understand whether patterns in plant community composition and the Floristic Quality Index (FQI) were related to productivity as evidenced by dormant season biomass yield. FQI is an objective measure of how closely a plant community represents that of a pre-European settlement community. Our research was conducted in planted fields of native tallgrass prairie species, and provided a gradient in floristic quality index, species richness, species diversity, and species evenness in south-central Wisconsin during 2008 and 2009. We used a network of 15 randomly located 1 m² plots within each field to characterize the plant community and estimate biomass yield by clipping the plots at the end of each growing season. While plant community composition and diversity varied significantly by planting type, biomass yield did not vary significantly among planting types (ANOVA; P >0.05). Biomass yield was positively correlated with plant community evenness, richness, C4 grass cover, and floristic quality index, but negatively correlated with plant species diversity in our multi-season multiple linear mixed effects models. Concordantly, plots with biomass yield in the lowest quartile (biomass yield < 3500 kh/ha) had 8% lower plant community evenness and 9% lower FQI scores than those in the upper quartile (biomass yield > 5800 kh/ha). Our results suggest that promoting the establishment of fields with high species evenness and floristic quality may increase biomass yield, while simultaneously supporting biodiversity.     

Compositional and functional consequences of environmental change in Belgian farmland ponds

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Climate mediates the effects of disturbance on ant assemblage structure

Many studies have focused on the impacts of climate change on biological assemblages, yet little is known about how climate interacts with other major anthropogenic influences on biodiversity, such as habitat disturbance. Using a unique global database of 1128 local ant assemblages, we examined whether climate mediates the effects of habitat disturbance on assemblage structure at a global scale. Species richness and evenness were associated positively with temperature, and negatively with disturbance. However, the interaction among temperature, precipitation and disturbance shaped species richness and evenness. The effect was manifested through a failure of species richness to increase substantially with temperature in transformed habitats at low precipitation. At low precipitation levels, evenness increased with temperature in undisturbed sites, peaked at medium temperatures in disturbed sites and remained low in transformed sites. In warmer climates with lower rainfall, the effects of increasing disturbance on species richness and evenness were akin to decreases in temperature of up to 9°C. Anthropogenic disturbance and ongoing climate change may interact in complicated ways to shape the structure of assemblages, with hot, arid environments likely to be at greatest risk.     

Hierarchical determinants of the functional richness, evenness and divergence of a vertebrate predator assemblage

Environmental factors are hypothesized to affect the functional diversity of assemblages hierarchically. First, ecological filters constrain the range of functional traits potentially displayed by an assemblage by determining its taxonomic composition. Second, some factors determine actual functional traits through the phenotypic plasticity of species. Little is known about the relative importance of each set of factors and, by using a priori functional information, most empirical studies report only the variability expected from species turnover and neglect that associated to phenotypic plasticity. Herein, we use structural equation models to assess the factors driving the functional richness, evenness, and divergence of a top‐predator assemblage faced with high variability in resource availability and assemblage structure (i.e. species richness and abundance). We measured actual functional traits (i.e. diet composition and predation pressure) in the field and contrasted the effects of environmental filters and phenotypic plasticity (i.e. behavioral responses) by controlling for species turnover and sample size with a null model. We found that a priori estimations (i.e. null‐model expectations) tended to significantly underestimate (richness and divergence) or overestimate (evenness) functional‐diversity components, explaining just a fraction (13–63%) of the variability in observed values. Furthermore, while species richness strongly affected functional richness (positively) and evenness (negatively), and resource availability slightly affected functional divergence, via compositional effects, changes in functional‐diversity components attributable to behavioral responses of predators showed little or no association with environmental variables. As a consequence, results indicated that in productive years, functionally‐distinctive species exerting relatively low predation pressure entered the assemblage, increasing functional richness and decreasing functional evenness. However, the strong behavioral responses of functionally dominant species buffered such compositional changes, affecting to different extents the three functional‐diversity components. Thus, we argue in favor of considering phenotypic plasticity in future studies of functional diversity.     

Effects of species evenness can be derived from species richness – ecosystem functioning relationships

Although species richness effects on ecosystem functioning have been studied thoroughly in countless experiments, the effects of the other side of diversity – species evenness – remain less identified, especially at high species richness. Due to the large number of different model ecosystems that need to be created, the explanatory power of the experimental approach for evenness is indeed limited. We show here that experimental studies on the influence of species richness on ecosystem functions contain hidden information on the influence of species evenness. Both the effects of maximum and minimum evenness, and of a key set of intermediate evenness levels, can be derived from species richness – ecosystem function curves, and that for every richness level, by using communities with low species richness as the equivalent of highly uneven communities with higher richness. We show that evenness effects on ecosystem functioning have the same direction as richness effects, however with increasing effect sizes at higher richness levels. We validated our technique for a wide range of ecosystem functions and applied it to the species richness – community biomass data from an existing biodiversity experiment. Our approach could provide a fast and easy alternative to resource‐intensive experiments in which evenness is experimentally varied, as we can build on the elaborate existing literature on species richness to assess its effects.     

Effects of experimental warming on biodiversity depend on ecosystem type and local species composition

Climatic warming is a primary driver of change in ecosystems worldwide. Here, we synthesize responses of species richness and evenness from 187 experimental warming studies in a quantitative meta‐analysis. We asked 1) whether effects of warming on diversity were detectable and consistent across terrestrial, freshwater and marine ecosystems, 2) if effects on diversity correlated with intensity, duration, and experimental unit size of temperature change manipulations, and 3) whether these experimental effects on diversity interacted with ecosystem types. Using multilevel mixed linear models and model averaging, we also tested the relative importance of variables that described uncontrolled environmental variation and attributes of experimental units. Overall, experimental warming reduced richness across ecosystems (mean log‐response ratio = –0.091, 95% bootstrapped CI: –0.13, –0.05) representing an 8.9% decline relative to ambient temperature treatments. Richness did not change in response to warming in freshwater systems, but was more strongly negative in terrestrial (–11.8%) and marine (–10.5%) experiments. In contrast, warming impacts on evenness were neutral overall and in aquatic systems, but weakly negative on land (7.6%). Intensity and duration of experimental warming did not explain variation in diversity responses, but negative effects on richness were stronger in smaller experimental units, particularly in marine systems. Model‐averaged parameter estimation confirmed these main effects while accounting for variation in latitude, ambient temperature at the sites of manipulations, venue (field versus lab), community trophic type, and whether experiments were open or closed to colonization. These analyses synthesize extensive experimental evidence showing declines in local richness with increased temperature, particularly in terrestrial and marine communities. However, the more variable effects of warming on evenness were better explained by the random effect of site identity, suggesting that effects on species’ relative abundances were contingent on local species composition. Synthesis A global research priority is to understand the consequences of climate change for biodiversity. A growing number of experimental studies have manipulated climatic drivers, in particular changes in temperature, in local communities. In the first quantitative meta‐analysis of community‐level studies across freshwater, marine and terrestrial experiments, species richness declined consistently with experimental warming. This effect was insensitive to warming intensity, duration, and multiple environmental and procedural covariates. However, evenness responses were weakly negative only in terrestrial systems and more variable across ecosystem types. Linear mixed model analyses revealed that the identity of local sites explained nearly 50% of variance in evenness effect sizes, compared to only 10% for richness. This result provides evidence that local species composition strongly constrains changes in relative species abundances in response to warming.     

宁夏六盘山国家级自然保护区眼蝶群落多样性

蝴蝶是地球生物多样性的重要指示物种之一, 其种群动态及群落结构变化能够揭示环境改变对生态系统的影响。六盘山国家级自然保护区生境优越, 植物区系多样, 孕育了丰富的蝶类资源。为了有效利用环境和保护蝶类资源, 2016-2018年, 作者采用样线调查法, 对宁夏六盘山国家级自然保护区人工植被、针阔混交林、阔叶林、灌木林4种生境中眼蝶亚科昆虫进行了调查, 分析了眼蝶在空间及时间上的动态变化。共记录眼蝶亚科13属19种4,363只, 其中阿芬眼蝶(Aphantopus hyperanthus, 961只)、亚洲白眼蝶(Melanargia asiatica, 637只)、爱珍眼蝶(Coenonympha oedippus, 594只)分别占总数的22%、15%和13%, 为保护区优势种; 蟾眼蝶(Triphysa phryne, 17只)仅占总数的0.39%, 为保护区的稀有种。在空间分布上, 4类生境中针阔混交林眼蝶物种丰富度和优势度低, 多样性指数最高, 不同生境眼蝶群落物种丰富度指数、多样性指数、均匀度指数存在明显差异(P < 0.05), 优势度指数差异不显著。在时间动态上, 2016-2018年眼蝶个体数量呈下降趋势, 物种个体数量高峰期主要发生在8月, 优势种变化明显。因此, 保护宁夏六盘山国家级自然保护区蝴蝶种类多样性刻不容缓。     

Effects of monoculture and polyculture practices in oil palm smallholdings on tropical farmland birds

Industrial oil palm expansion has led to dramatic landscape changes that have negatively affected forest biodiversity in the tropics. In contrast to large-scale plantations, oil palm smallholdings may support greater levels of biodiversity through the implementation of multi-cropping system or polyculture. We examined bird species richness, together with community structure, conservation status, and feeding guild of existing smallholdings in Peninsular Malaysia. Based on point transect sampling, we sampled birds in 100 smallholdings that practiced either monoculture or polyculture farming. Our results revealed that bird species richness was significantly greater in monoculture smallholdings than in polyculture smallholdings, but the opposite was true for bird abundance. Non-forest birds constituted the major species of bird communities in oil palm smallholdings. However, we found that the abundances of insectivores and frugivores were greater in polyculture smallholdings than in monoculture smallholdings. In the monoculture models, predictor variables explained 11.31–19.98% of the variation in bird species richness. When polyculture was being practiced, bird species richness increased significantly with the height of ground vegetation cover, distance to major roads, and distance to rice fields. In the polyculture models, predictor variables accounted for 11.71–24.85% of the variation in bird species richness. We also found that bird species richness increased significantly with height of ground vegetation, but it decreased with ground vegetation cover and distance to rivers. The evidence from this study suggests that monoculture and polyculture farming were able to maintain farmland biodiversity in smallholdings, at least for birds, but differed in richness, population, and feeding guild.     

A Comparative Study of the Far Eastern Seas and the Northern Pacific Ocean Based on Integral Parameters of Net Zooplankton in the Epipelagic Layer

Integral parameters of zooplankton community, including species diversity and its components were compared between the Chukchi Sea, Bering Sea, Sea of Okhotsk, Sea of Japan, and adjacent Pacific waters based on the data obtained by standard Juday net with a mouth area of 0.1 m² during the large-scale surveys conducted by the Pacific Fisheries Research Center (TINRO Center) in 1984–2013. These parameters were calculated for the total surveyed area of approximately 7.0 million km² and separately for each of the considered water bodies. In Pacific waters, species richness is higher than that in all the seas, while the concentration of individuals (expressed in terms of abundance, ind./m³) and evenness of their distribution over species were lower. The only sea with a larger mean size of organisms compared to the ocean is the Bering Sea. A lower species diversity than in the ocean has been recorded only from the Chukchi Sea; a lower density (in terms of biomass, g/m³) was determined only from the Sea of Japan. Among the four seas, the Chukchi Sea ranks first in terms of biomass and abundance of zooplankton, second in species evenness, third in the mean size of individuals, and last in species richness and diversity. The Bering Sea ranks first in terms of mean size of plankton organisms, second in species richness, diversity, and biomass, third in abundance, and last in species evenness. The Sea of Okhotsk ranks second in terms of mean size of individuals, last in their abundance, and third in the other parameters. The Sea of Japan ranks first in terms of species richness, evenness, and diversity, second in abundance, and last in mean size of zooplankton organisms, and, therefore, their biomass. The biomass of zooplankton, in accordance with the concentration of nutrients, increases in the southto-north direction (while its absolute abundance depends largely on the size of the body of water). The mean size of organisms increases in the same direction; the evenness of their distribution over species increases in the reverse direction (with the exception of both parameters for the Chukchi Sea). The rank of a water body for its biodiversity coincides with the species richness rank. The latter increases from north to south (except for the Okhotsk Sea), but greatly depends on the surveyed area and, even more, on the surveyed volume of water. A study of the literature data found some unexpected statistically significant relationships of the integral parameters of zooplankton with those of pelagic and bottom macrofauna, as well as with the parameters of zooplankton production, on the size of the considered bodies of water. The causes and the biological meanings of most of these relationships still do not have any rational interpretation. Their testing at other spatial scales will be continued in future works.     

高寒草甸植物群落中相似性、物种多样性与地上生物量的年际变异性

相似性似说通过物种构成的相似性来解释物种丧失是如何影响生物量的变异性的,但还没有得到检验。本研究通过设置在青藏高原东部地区的高寒草甸植物群落中的74个永久样方．采集3年(1999～2001)植物生长高峰期的群落数据,试图检验物种构成的相似性是如何解释物种多样性对地上生物量年际变异性的影响。结果表明：随着物种丰富度增加,生物量变异性降低;而随着均匀度的增加,生物量的变异性尽管在均匀度中等程度时似乎保持在同一水平,但总体上呈下降趋势;物种构成上的相似性解释了地上生物量变异性的大部分,而且随着物种构成上的相似性的增加,生物量的变异性降低;物种丰富度和均匀度均与物种构成上的相似性没有显著相关关系。这些结果表明：尽管生物多样性的丧失可能不必导致物种丰富群落中物种构成上的相似性,但相似性与地上生物量的变异性的因果联系可能是稳健的．由于本研究是在自然群落中进行的,对物种构成的相似性没有进行直接控制,因此,要深入理解相似性是如何影响生物多样性对生态系统功能变异性的效应的机制,可能还需要直接对物种构成的相似性进行控制的实验研究。     

Identifying hotspots for biodiversity management using rank abundance distributions

Aim Identification of biodiversity hotspots has typically relied on species richness. We extend this approach to include prediction to regional scales of other attributes of biodiversity based on the prediction of Rank Abundance Distributions (RADs). This allows us to identify areas that have high numbers of rare species and areas that have a rare assemblage structure. Location Continental slope and shelf of south‐western Australia, between 20.5 and 30° S and depths of 100–1500 m. Methods We use a recently developed method to analyse RADs from biological surveys and predict attributes of RADs to regional scales from spatially abundant physical data for demersal fish and invertebrates. Predictions were made for total abundance (N), species richness (S) and relative evenness at 147,996 unsampled locations using data from two spatially limited surveys. The predictions for S and relative evenness were then independently split into categories, creating a bivariate distribution. The RAD categories are mapped spatially between 20.5 and 30° S to depths of 1500 m to allow identification of areas with rare species and assemblage structure across this region. Results Rank abundance distributions for demersal fish vary with large scale oceanographic patterns. Peaks in abundance and unevenness are found on the shelf break. The bivariate distributions for richness and evenness for both fish and invertebrates show that all assemblage structures are not equally likely. The RAD categories identify regions that have high numbers of rare species and areas with unique assemblage structure. Main conclusions Predicted RADs over large regions can be used to identify biodiversity hotspots in more detail than richness alone. Areas of rare species and rare assemblage structure identified from fish and invertebrates largely overlap, despite the underlying data coming from two different data sets with two different collection methods. This approach allows us to target conservation management at species that would otherwise be missed.     

Land use types influenced avian assemblage structure in a forest–agriculture landscape in Ghana

The conservation of biodiversity within tropical forest regions does not lie only in the maintenance of natural forest areas, but on conservation strategies directed toward agricultural land types within which they are embedded. This study investigated variations in bird assemblages of different functional groups of forest‐dependent birds in three agricultural land types, relative to distance from the interior of 34 tropical forest patches of varying sizes. Point counts were used to sample birds at each study site visited. Data from counts were used to estimate species richness, species evenness, and Simpson's diversity of birds. Mean species richness, evenness, and diversity were modeled as responses and as a function of agricultural land type, distance from the forest interior and three site‐scale vegetation covariates (density of large trees, fruiting trees, and patch size) using generalized linear mixed‐effect models. Mean observed species richness of birds varied significantly within habitat types. Mean observed species richness was highest in forest interior sites while sites located in farm centers recorded the lowest mean species richness. Species richness of forest specialists was strongly influenced by the type of agricultural land use. Fallow lands, density of large trees, and patch size strongly positively influenced forest specialists. Insectivorous and frugivorous birds were more species‐rich in fallow lands while monoculture plantations favored nectarivorous birds. Our results suggest that poor agricultural practices can lead to population declines of forest‐dependent birds particularly specialist species. Conservation actions should include proper land use management that ensures heterogeneity through retention of native tree species on farms in tropical forest‐agriculture landscapes.     

Species richness changes across two trophic levels simultaneously affect prey and consumer biomass

Increasing species richness of primary producers or consumers is proposed to increase primary and secondary production; however, the consequences of biodiversity change across trophic levels has been poorly investigated. We used a controlled marine microbial system to investigate the effects of simultaneous changes in biodiversity of consumer and prey species. Consumer (ciliates) and prey (algae) richness and identity were manipulated independently in a complete factorial design. The results showed clear biodiversity effects of both consumers and prey, within and across trophic levels. We found reduced prey and increased consumer biomass with increased consumer richness, with the most diverse prey assemblage supporting the highest biomass of consumers at the highest richness of consumers. Increasing prey richness did not increase resistance to consumption when consumers were present. Instead, our results indicated enhanced energy transfer with simultaneous increasing richness of consumers and prey.     

The relationship between species richness and evenness: a meta-analysis of studies across aquatic ecosystems

Biological diversity comprises both species richness, i.e., the number of species in a community, and evenness, measuring how similar species are in their abundances. The relationship between species richness and evenness (RRE) across communities remains, however, a controversial issue in ecology because no consistent pattern has been reported. We conducted a systematic meta-review of RRE in aquatic ecosystems along regional to continental gradients and across trophic groups, differing in body size by 13 orders of magnitude. Hypotheses that RRE responded to latitudinal and scale variability across trophic groups were tested by regression analyses. Significant correlations of species richness and evenness only existed in 71 out of 229 datasets. Among the RRE, 89 were negative and 140 were positive. RRE did not vary with latitude but showed a positive response to scale. In a meta-analysis with ecosystem type as a single explaining variable, RRE did not vary among ecosystem types, i.e. between marine and freshwater. Finally, autotrophs had more positive RRE than heterotrophs. The weak RRE in many aquatic datasets suggests that richness and evenness often reflect independent components of biodiversity, highlighting that richness alone may be an incomplete surrogate for biodiversity. Our results further elucidate that RRE is driven by organismal and environmental properties, both of which must be considered to gain a deeper understanding of large-scale patterns of biodiversity.     

What Factors Affect Diversity and Species Composition of Endangered Tumbesian Dry Forests in Southern Ecuador?

This paper reports a study on species richness and composition of Tumbesian dry forest communities. We tested two alternative hypotheses about species assemblage processes in tropical dry forests: (1) species assemblage is determined by the filtering effect of environmental conditions and (2) species assemblage is determined by facilitative processes along the gradient of water availability, and thus, species richness and evenness increase as water becomes limited. In addition, we also explored the effect of climate and soil conditions on species composition in tropical dry forests. Species composition was sampled in 109 plots in terms of cover and tree diameter at breast height. Climatic, edaphic, topographic and anthropogenic degradation variables were obtained for each plot. We used generalized linear models and canonical correspondence analyses to evaluate the effect of environmental variables on species composition, richness and evenness. Water availability negatively affected richness and significantly determined the species assemblage. Species richness increased from ridges to valleys and evenness increased at higher altitudes. Soil characteristics showed no effect on richness and evenness but soil moisture, nitrogen concentration and soil temperature explained significant fractions of species composition. Although timber extraction and livestock in our study area were of low intensity, it negatively affected richness but had only a minor effect on species composition. Our results suggest that species composition in these endangered tropical dry forests may be at least partially explained by the stress‐gradient hypothesis, with higher species richness at drier conditions probably induced by facilitation processes.