Natural Environmental Gradients Research Articles

Submarine groundwater discharge drives both direct and indirect effects on organismal and community metabolism on coral reefs.

Coral reefs experience numerous environmental gradients affecting organismal physiology and species biodiversity, which ultimately impact community metabolism. This study shows that submarine groundwater discharge (SGD), a common natural environmental gradient in coastal ecosystems associated with decreasing temperatures, salinity and pH with increasing nutrients, has both direct and indirect effects on coral reef community metabolism by altering individual growth rates and community composition. Our data revealed that SGD exposure hindered the growth of two algae, Halimeda opuntia and Valonia fastigiata, by 67 and 200%, respectively, and one coral, Porites rus, by 20%. Community metabolic rates showed altered community production, respiration and calcification between naturally high and low exposure areas mostly due to differences in community identity (i.e. species composition), rather than a direct effect of SGD on physiology. Production and calcification were 1.5 and 6.5 times lower in assemblages representing high SGD communities regardless of environment. However, the compounding effect of community identity and SGD exposure on respiration resulted in the low SGD community exhibiting the highest respiration rates under higher SGD exposure. By demonstrating SGD's role in altering community composition and metabolism, this research highlights the critical need to consider compounding environmental gradients (i.e. nutrients, salinity and temperature) in the broader context of ecosystem functions.

Conspecific interactions between corals mediate the effect of submarine groundwater discharge on coral physiology.

Land-based inputs, such as runoff, rivers, and submarine groundwater, can alter biologic processes on coral reefs. While the abiotic factors associated with land-based inputs have strong effects on corals, corals are also affected by biotic interactions, including other neighboring corals. The biologic responses of corals to changing environmental conditions and their neighbors are likely interactive; however, few studies address both biotic and abiotic interactions in concert. In a manipulative field experiment, we tested how the natural environmental gradient created by submarine groundwater discharge (SGD) affected holobiont and symbiont metabolic rates and endosymbiont physiology of Porites rus. We further tested how the effect of SGD on the coral was mediated by intra and interspecific interactions. SGD is a natural land-sea connection that delivers nutrients, inorganic carbon, and other solutes to coastal ecosystems worldwide. Our results show that a natural gradient of nutrient enrichment and pH variability as a result of acute SGD exposure generally benefited P. rus, increasing gross photosynthesis, respiration, endosymbiont densities, and chlorophyll a content. Conspecifics in direct contact with the a neighboring coral, however, altered the relationship between coral physiology and SGD, lowering the photosynthetic and respiration rates from expected values when the coral had no neighbor. We show that the response of corals to environmental change is dependent on the types of nearby neighbor corals and how neighbors alter the chemical or physical environment around the coral. Our study underscores the importance of considering biotic interactions when predicting the physiologic responses of corals to the environment.

Deconstructing the geography of human impacts on species’ natural distribution

It remains unknown how species’ populations across their geographic range are constrained by multiple coincident natural and anthropogenic environmental gradients. Conservation actions are likely undermined without this knowledge because the relative importance of the multiple anthropogenic threats is not set within the context of the natural determinants of species’ distributions. We introduce the concept of a species ‘shadow distribution’ to address this knowledge gap, using explainable artificial intelligence to deconstruct the environmental building blocks of current species distributions. We assess shadow distributions for multiple threatened freshwater fishes in Switzerland which indicated how and where species respond negatively to threats — with negative threat impacts covering 88% of locations inside species’ environmental niches leading to a 25% reduction in environmental suitability. Our findings highlight that conservation of species’ geographic distributions is likely insufficient when biodiversity mapping is based on species distribution models, or threat mapping, without also quantifying species’ expected or shadow distributions. Overall, we show how priority actions for nature’s recovery can be identified and contextualised within the multiple natural constraints on biodiversity to better meet national and international biodiversity targets.

Below‐ground traits, rare species and environmental stress regulate the biodiversity–ecosystem function relationship

Abstract Understanding the relationship between biodiversity and ecosystem functioning (BEF) is crucial to predicting the consequences of ongoing global biodiversity loss. However, what drives BEF relationships in natural ecosystems under globally changing conditions remains poorly understood. To address this knowledge gap, we applied a trait‐based approach to data from coastal dune plant communities distributed along a natural environmental stress gradient. Specifically, we compared the relative importance of below‐ground and above‐ground traits in predicting productivity, decomposition, water regulation, carbon stock and nutrient pools, and tested how these BEF relationships were modulated by environmental stress and the presence of rare species that are typically excluded from experimental systems. Below‐ground traits were just as important as above‐ground traits in driving ecosystem functioning. Moreover, despite having low abundances, rare species positively influenced ecosystem multifunctionality (EMF). However, most biodiversity effects became weaker as environmental stress increased. Our study shows that to understand variation in ecosystem functioning we must consider below‐ground traits as much as above‐ground ones. Moreover, it highlights the importance of conserving rare species for maintaining EMF. However, our findings also suggest that rapid global change could dampen the positive effects of diversity on ecosystem functioning. Read the free Plain Language Summary for this article on the Journal blog.

Quantifying the functional genes of C, N, P, and S cycling in a deep lake: Depth patterns and drivers

Microbial functional composition is important for biogeochemical cycles, and is usually constrained by taxonomic species pool and natural environmental gradients. However, the distribution of functional genes in deep lakes and the driving factors remains elusive. Here, we quantified the abundance of 71 functional genes relevant to carbon degradation, carbon fixation, methane metabolism, nitrogen cycling, phosphorus cycling, and sulfur cycling in 38 sediments along the water depth ranging from 0 to 90 m in Lugu Lake, China, using Quantitative Microbial Ecology Chip (QMEC) and then explored their water-depth diversity pattern and abiotic and biotic drivers. Functional gene diversity showed a hump-shaped pattern along the depth and peaked at around 50 m which is the low boundary of thermocline layer. There were specific environmental preferences among functional gene subgroups such as nitrogen and sulfur cycling genes preferring to deeper and shallow waters, respectively. The dissimilarity of total functional genes increased with water depth distance indicating a distance-decay relationship. There was a congruence between functional and taxonomic composition by showing the positive correlations between the compositions of functional genes and bacteria or archaea. This phenomenon is consistently observed for the six functional gene subgroups, and the congruence strength was highest for nitrogen cycling while lowest for sulfur cycling. Compared to abiotic factors, biotic factors were more relevant to the functional gene diversity and composition. Biotic factors explained 28.5 % of the variance of functional gene diversity, while water depth and other environmental factors such as water total phosphorus and sediment carbon explained 4.2 % and 3.8 %, respectively. For functional gene composition, biotic factors accounted for 25.2 % of the variance, whereas water depth and other environmental factors contributed to 0.7 % and 4.4 %, respectively. Among all explanatory variables for function genes, bacterial composition had the highest contribution, which was further supported by showing its direct effects of 1.45 and 0.86 on functional diversity and composition, respectively. This study for the first time quantified the microbial functional genes along water depth in deep lakes and provided a more comprehensive understanding of the functional dynamics in microbial communities within aquatic ecosystems.

Robust hydraulic traits correlation in woody species despite large trait variation along natural and experimental environmental gradients

Abstract Hydraulic traits are major determinants of plant fitness, thus exerting control over vegetation structure, function and distribution. Yet it remains unclear whether and how hydraulic traits respond to environmental stimuli (i.e. phenotypic variation of hydraulic traits; PVHT) and if the coordination between different hydraulic traits and the trait–climate relationship are affected by PVHT. Here, we synthesized data of PVHT (maximum hydraulic conductivity and water potential inducing 50% loss of hydraulic conductivity) and potentially related morphological and anatomical traits (e.g. sapwood density, branch Huber value, mean and hydraulic weighted conduit diameter). We analysed the magnitude, direction and source of variation of the plastic response, as well as the influence of environmental factors on trait coordination. Additionally, we compared the intra‐ and inter‐specific variation between key hydraulic traits and climate metrics (mean annual precipitation and mean annual temperature) at the site of growth, as well as across the population range. PVHT was highly variable in both magnitude and direction, which was contingent on the environmental factor. The variation in PVHT mainly occurred at high taxonomic levels (i.e. family and genus), whereas phenology explained little variation for PVHT. Despite the high variability, trait correlation remained robust in the presence of environmental stimuli. Moreover, trait–climate relationships differed at inter‐specific and intra‐specific levels. The intra‐specific variation of hydraulic traits in most species showed no correlation with climate metrics compared with the high correlation of hydraulic traits with climate metrics across species. Our findings suggest that the high variability of PVHT does not affect the trait correlation which may be valuable in predicting vegetation dynamics under varying environments. The distinct trait–climate relationships highlight the need to unravel the driving force of PVHT, as well as the adaptive strategy across populations. Read the free Plain Language Summary for this article on the Journal blog.

Improving the performance of macroinvertebrate based multi-metric indices by incorporating functional traits and an index performance-driven approach

Human-driven multiple pressures impact freshwater ecosystems worldwide, reducing biodiversity, and impacting ecosystem functioning and services provided to human societies. Multi-metric indices (MMIs) are suitable tools for tracking the effects of anthropogenic pressures on freshwater ecosystems because they incorporate various biological metrics responding to multiple pressures at different levels of biological organization. However, the performance and applicability of MMIs depend on their metrics' selection and their calibration against natural environmental gradients. In this study, we aimed to unravel i) how incorporating functional trait-based metrics affects the performance of MMIs, ii) how disentangling the natural environmental gradients from anthropogenic pressures effects affects the performance of MMIs, and iii) how the performance of MMIs developed using a metric performance-driven approach compares with MMIs developed using an index performance-driven approach. We carried out a field survey measuring abiotic and biotic variables at 53 sites in the Karun River basin (Iran) in 2018. For functional trait-based metrics, we used 15 macroinvertebrate traits and calculated community-weighted mean trait values and functional diversity indices. We used random forest modeling to account for the effect of natural environmental gradients on each metric. Based on our results, incorporating functional traits increased the MMI performance significantly and facilitated ecological interpretation of MMIs. Both taxonomic and functional components of macroinvertebrate assemblages co-varied strongly with natural environmental gradients, and accounting for these covariations improved the performance of MMIs. Finally, we found that index performance-driven MMIs performed better in terms of precision, bias, sensitivity, and responsiveness than metric performance-driven MMIs.

Regional and fine-scale local adaptation in salinity tolerance in Daphnia inhabiting contrasting clusters of inland saline waters.

Understanding the spatial scales at which organisms can adapt to strong natural and human-induced environmental gradients is important. Salinization is a key threat to biodiversity, ecosystem functioning and the provision of ecosystem services of freshwater systems. Clusters of naturally saline habitats represent ideal test cases to study the extent and scale of local adaptation to salinization. We studied local adaptation of the water flea Daphnia magna, a key component of pond food webs, to salinity in two contrasting landscapes-a dense cluster of sodic bomb crater ponds and a larger-scale cluster of soda pans. We show regional differentiation in salinity tolerance reflecting the higher salinity levels of soda pans versus bomb crater ponds. We found local adaptation to differences in salinity levels at the scale of tens of metres among bomb crater pond populations but not among geographically more distant soda pan populations. More saline bomb crater ponds showed an upward shift of the minimum salt tolerance observed across clones and a consequent gradual loss of less tolerant clones in a nested pattern. Our results show evolutionary adaptation to salinity gradients at different spatial scales, including fine-tuned local adaptation in neighbouring habitat patches in a natural landscape.

A new method for bioassessment of ecosystems with complex communities and environmental gradients

Bioassessment of complex and heterogeneous ecosystems is a challenge when there are multiple, strong, natural environmental gradients; unknown, or spatially varying, mixtures of stressors; and large numbers of taxa with unknown responses to both the environmental gradients and the stressors. Current methods of bioassessment are not designed for use under this set of constraints. To address this gap, we have developed an assessment method appropriate for well-sampled, heterogeneous systems with many taxa. In the bioassessment described below, we model taxa occurrence as a function of natural environmental gradients, then use residual covariance patterns between all pairs of taxa to estimate the impact of human disturbance across sites as a latent construct. The derivation of the method from an underlying causal model allows the metric value at each site and the associated taxa responses to be partitioned into contributions from a set of putative stressors. We apply this method as a case study to the subtidal benthic invertebrate community of Puget Sound, WA (USA) and demonstrate a partial decomposition of the metric values to a set of stressors including sediment organic carbon, nitrogen, metals, and organic pollutants. While this method provides new opportunities to estimate, communicate, and understand the ecological condition of complex, heterogeneous ecosystems, due to the requirement for broad, detailed data to inform its estimates, it will likely be most appropriate for monitoring programs.

Evaluating the effectiveness of M-AMBI with other biotic indexes in a temperate estuary

Need for a scalable and widely applicable index has been increasingly important. This study evaluates the applicability of the M-AMBI, a potential comprehensive index, at small spatial scales. M-AMBI was compared to regional indices (EMAP-E and GOM B-IBI), assessing response to natural environmental gradients and low oxygen stress. Results indicate poor agreement between indices with M-AMBI and GOM B-IBI showing positive correlation but significant disagreement in habitat condition. EMAP-E had no agreement. Indices showed similar patterns of better habitat scores in higher salinities. M-AMBI also showed a negative relationship with sediment organic matter and total nitrogen. DO influenced all indices with M-AMBI the most sensitive. However, mismatches between DO and index score were observed further calibration may be needed before adoption into programs. Overall, the M-AMBI demonstrates potential at smaller, local scales, but additional studies are needed to validate its performance in different coastal environments and under different conditions.

Environmental and genetic variation in an asexual plant

Species may respond to variation in environmental conditions by modifying their phenotype to match local levels of resource availability. This phenotypic response can be driven by plastic physiological change, or by adaptive genetic change. Here we use Lemna minor (lesser duckweed), a small aquatic macrophyte that is increasingly used as a model in ecology and evolution, to investigate the source and maintenance of phenotypic variation in natural environments. We found substantial phenotypic variation in L. minor in the field, with its frond area and root length changing predictably over natural environmental gradients of resource availability. Separating environmental and genetic variation in these traits in a common garden, we attribute the majority of phenotypic variation we observed in the field to phenotypic plasticity. Despite this, there was substantial within-site genetic variation. We found evidence of strong purifying selection in the field, that is necessarily balanced by mutation and migration. Using measures of environmental and genetic variation in phenotype and fitness, we estimate the rates of dispersal and evolution of fitness necessary to sustain the observed levels of genetic variation.

Context-dependence of fungal community responses to dominant tree mycorrhizal types in Northern hardwood forests

Dominant tree community mycorrhizal associations can influence soil biogeochemistry and nutrient cycling, suggesting a prominent role of mycorrhizas in shaping belowground microbial community composition and function. The degree to which the mycorrhizal type of dominant trees interacts with natural environmental gradients to influence belowground microbial communities is, however, unclear. Likewise, it is unknown if community-level mycorrhizal associations can influence the local microbial community encountered by an individual tree through spillover effects. To address these questions, we studied fungal communities from soil, roots, and leaf litter surrounding individual arbuscular (AM) and ectomycorrhizal (ECM) trees embedded in gradients of tree mycorrhizal dominance from three climatically distinct locations in the Adirondack Mountains, NY, USA. We found that dominant tree mycorrhizal types interact with site location to explain more variation in fungal community composition, richness, and function than specific soil properties, such as pH. This finding was consistent for all three sample types, but soil-associated fungi demonstrated the largest amount of explainable variation compared to root- and leaf litter-associated fungi. The relative abundance of plant pathogens was especially responsive to tree mycorrhizal dominance, increasing with AM dominance around individual AM trees but not around ECM trees in the same forests. These “mycorrhizal-spillover” effects on AM trees were also strongest in our warmest, driest site and weakest in our coolest, wettest site, indicating that the strength of mycorrhizal spillover is context-dependent in mixed-mycorrhizal forests.

Functional diversity of macrofaunal assemblages as indicators to assess heavy metal pollution in the Bohai Sea, China.

Functional diversity of macrofaunal assemblages can reflect the composition and differences of functional traits, indicating their response to various contaminants, especially heavy metal pollution. We explored the effects of environment variables over gradients of heavy metal pollution on macrofaunal assemblages, using biological traits analysis, generalized linear model (GLM), AZTI marine biotic index (AMBI), and various biodiversity indexes. The RLQ (co-inertia analysis) and fourth-corner approaches were used to investigate the specific response of functional traits to heavy metal pollution. Most sites were environmentally degraded by heavy metal pollution and macrofaunal body size had a miniaturization trend. There was a significant correlation between functional diversity indexes and AMBI. The RLQ and fourth-corner analysis and GLM models showed that heavy metal and natural environmental gradients had a profound effect on functional diversity. The functional divergence and dispersion indexes, along with the abundance of some specific species, were appropriate indexes for heavy metal pollution.

Salinity Induces Allometric Accumulation of Sulfur in Plants and Decouples Plant Nitrogen‐Sulfur Correlation in Alpine and Arid Wetlands

AbstractSalinization alters the elemental balance of wetlands and induces variations in plant survival strategies. Sulfur (S) plays vital roles in serving regulatory and catalytic functions in stress resistance of plants. Yet, how plant S and its relationships with nitrogen (N) vary across natural environmental gradients are not well documented. We collected 1,366 plant samples and 230 water and sediment samples from 230 wetlands in Tibetan Plateau and adjacent arid regions of western China, to analyze the effects of environmental variables on plant S accumulation and N‐S correlations. We found that plant S correlated with N in unimodal patterns. Salinity, rather than temperature or nutrient supply, promoted disproportionate accumulation of S but limited N uptake, inducing decoupling of N‐S correlation in plants. Toward high salinity, the faster increasing rates of total S than that of glutathione, the most abundant organic‐S compound in plant resistance, provided potential evidences explaining the decoupled plant N‐S correlation. A salinity of 3.9‰ was calculated to be a threshold at which substantial changes in plant N‐S correlation occurred. We designed a conceptual model to illustrate the mechanisms driving variations of N‐S correlation in plants and environments along salinity gradient. Furthermore, high salinity filtered out the salt‐sensitive species and reassembled the communities. In conclusion, increased salinity affected wetland plants by inducing S accumulation in plants and selecting salt‐tolerant species with high S concentrations at community level, providing evidences for plant adaptive mechanisms to salinity in arid regions.

Living and dead bivalves are congruent surrogates for whole benthic macroinvertebrate communities in Puget Sound

To integrate paleoecological data with the “whole fauna” data used in biological monitoring, analyses usually must focus on the subset of taxa that are inherently preservable, for example by virtue of biomineralized hardparts, and those skeletal remains must also be identifiable in fragmentary or otherwise imperfect condition, thus perhaps coarsening analytical resolution to the genus or family level. Here we evaluate the ability of readily preserved bivalves to reflect patterns of compositional variation from the entire infaunal macroinvertebrate fauna as typically sampled by agencies in ocean monitoring, using data from ten long-established subtidal stations in Puget Sound, Washington State. Similarity in compositional variation among these stations was assessed for five taxonomic subsets (the whole fauna, polychaetes, malacostracans, living bivalves, dead bivalves) at four levels of taxonomic resolution (species, genera, families, orders) evaluated under four numerical transformations of the original count data (proportional abundance, square root- and fourth root-transformation, presence-absence). Using the original matrix of species-level proportional abundances of the whole fauna as a benchmark of “compositional variation,” we find that living and dead bivalves had nearly identical potential to serve as surrogates of the whole fauna; they were further offset from the whole fauna than was the polychaete subset (which dominates the whole fauna), but were far superior as surrogates than malacostracans. Genus- and family-level data were consistently strong surrogates of species-level data for most taxonomic subsets, and correlations declined for all subsets with increasing severity of data transformation, although this effect lessened for subsets with high community evenness. The strong congruence of death assemblages with living bivalves, which are themselves effective surrogates of compositional variation in the whole fauna, is encouraging for using bivalve dead-shell assemblages to complement conventional monitoring data, notwithstanding strong natural environmental gradients with potential to bias shell preservation.

Integrating natural gradients, experiments, and statistical modeling in a distributed network experiment: An example from the WaRM Network.

A growing body of work examines the direct and indirect effects of climate change on ecosystems, typically by using manipulative experiments at a single site or performing meta‐analyses across many independent experiments. However, results from single‐site studies tend to have limited generality. Although meta‐analytic approaches can help overcome this by exploring trends across sites, the inherent limitations in combining disparate datasets from independent approaches remain a major challenge. In this paper, we present a globally distributed experimental network that can be used to disentangle the direct and indirect effects of climate change. We discuss how natural gradients, experimental approaches, and statistical techniques can be combined to best inform predictions about responses to climate change, and we present a globally distributed experiment that utilizes natural environmental gradients to better understand long‐term community and ecosystem responses to environmental change. The warming and (species) removal in mountains (WaRM) network employs experimental warming and plant species removals at high‐ and low‐elevation sites in a factorial design to examine the combined and relative effects of climatic warming and the loss of dominant species on community structure and ecosystem function, both above‐ and belowground. The experimental design of the network allows for increasingly common statistical approaches to further elucidate the direct and indirect effects of warming. We argue that combining ecological observations and experiments along gradients is a powerful approach to make stronger predictions of how ecosystems will function in a warming world as species are lost, or gained, in local communities.

Spatial variation in the direct and indirect effects of plant diversity on soil respiration in an arid region

Soil respiration (Rs) is a key ecosystem function that controls the terrestrial energy balance and element cycling. Quantifying how plant diversity affects Rs is critical for predicting the impact of global plant diversity loss on ecosystem function. However, it is unclear how soil factors and plant diversity are spatially interrelated under natural environmental gradients, and how plant diversity affects the spatial variation of Rs on the basis of soil factors. Our objectives were to quantify the direction and magnitude of direct and indirect effects of plant diversity on Rs. We assessed spatial variability in the relationships between soil factors (soil chemistry and microclimate), multiple dimensions of plant diversity (taxonomic, functional, and phylogenetic diversity), and Rs using a new combination of geographically weighted regression and structural equation modeling based on survey data from three plots (river bank, transitional zone, and desert margin plots) along an environmental gradient in northwest China. Reduced plant diversity had a negative impact on Rs. However, when the effects of soil factors on Rs and spatial non-stationarity were considered simultaneously, the relationship between plant diversity and Rs changed in magnitude and direction. From the river bank to the desert margin, the positive effect of taxonomic and functional diversity on Rs gradually weakened, while the effect of phylogenetic diversity on Rs changed from a negative to a positive effect. Plant diversity and soil factors together explained 55%–75% of the spatial variation in Rs. Among them, the contribution of plant diversity to the spatial variation of Rs gradually decreased from the river bank to the desert margin, while the contribution of soil factors gradually increased. Functional diversity (17.39%–18.93%) and phylogenetic diversity (10.75%–19.53%) better explained the spatial variability of Rs compared with taxonomic diversity (2.67%–6.01%). The results provide strong evidence that plant diversity, especially functional and phylogenetic diversity, is a key driver of Rs, expanding our understanding of the relationship between plant diversity and Rs. These findings more accurately reveal the changing characteristics of carbon dynamics in desert ecosystems, and provide a methodological framework for studying the spatial variability of the relationship between plant diversity and ecosystem function.

Tree diversity depending on environmental gradients promotes biomass stability via species asynchrony in China’s forest ecosystems

There is mounting evidence that biodiversity promotes ecological stability in changing environments. However, understanding diversity–stability relationships and their underlying mechanisms across large-scale tree diversity and natural environmental gradients are still controversial and largely lacking. We used thirty-nine 0.12 ha long-term permanent forest plots spanning China’s various forest types to test the effects of multiple abiotic (climate, soil, age and topography) and biotic factors (taxonomic and structural diversity, functional diversity and community-mean traits, and species asynchrony) on biomass stability and its components (mean biomass and biomass variability) over time. We used multiple analytical methods to identify the best explanatory variables and complicated causal relationships for community biomass stability. Our results showed that species richness increased biomass stability by promoting species asynchrony. Structural and functional diversity had a weaker effect on biomass stability. Forest age and structural diversity increased mean biomass and biomass variability significantly and simultaneously. Communities dominated by tree species with high wood density had high biomass stability. Soil nitrogen enhanced biomass stability directly and indirectly through its effects on mean biomass. Soil nitrogen to phosphorus ratio increased biomass stability via increasing species asynchrony. Precipitation indirectly increased biomass stability by affecting tree diversity. Moreover, the direct and indirect effects of soil nutrients on biomass stability were greater than that of climate variables. Our results suggest that species asynchrony is the main mechanism proposed to explain the stabilizing effect of diversity on community biomass, supporting two mechanisms, namely, the biodiversity insurance hypothesis and complementary dynamics. Soil and climate factors also play an important role in shaping diversity–stability relationships. Our results provide a new insight into how tree diversity affects ecosystem stability across diverse community types and large-scale environmental gradients.

Links between leaf anatomy and leaf mass per area of herbaceous species across slope aspects in an eastern Tibetan subalpine meadow.

Leaf anatomy varies with abiotic factors and is an important trait for understanding plant adaptive responses to environmental conditions. Leaf mass per area (LMA) is a key morphological trait and is related to leaf performance, such as light‐saturated photosynthetic rate per leaf mass, leaf mechanical strength, and leaf lifespan. LMA is the multiplicative product of leaf thickness (LT) and leaf density (LD), both of which vary with leaf anatomy. Nevertheless, how LMA, LT, and LD covary with leaf anatomy is largely unexplored along natural environmental gradients. Slope aspect is a topographic factor that underlies variations in solar irradiation, air temperature, humidity, and soil fertility. In the present study, we examined (1) how leaf anatomy varies with different slope aspects and (2) how leaf anatomy is related to LMA, LD, and LT. Leaf anatomy was measured for 30 herbaceous species across three slope aspects (south‐, west‐, and north‐facing slopes; hereafter, SFS, WFS, and NFS, respectively) in an eastern Tibetan subalpine meadow. For 18 of the 30 species, LMA data were available from previous studies. LD was calculated as LMA divided by LT. Among the slope aspects, the dominant species on the SFS exhibited the highest LTs with the thickest spongy mesophyll layers. The thicker spongy mesophyll layer was related to a lower LD via larger intercellular airspaces. In contrast, LD was the highest on NFS among the slope aspects. LMA was not significantly different among the slope aspects because higher LTs on SFS were effectively offset by lower LDs. These results suggest that the relationships between leaf anatomy and LMA were different among the slope aspects. Mechanisms underlying the variations in leaf anatomy may include different solar radiation, air temperatures, soil water, and nutrient availabilities among the slope aspects.

Estuarine fish diversity as indicator of natural environmental gradients

Estuarine fish diversity as indicator of natural environmental gradients