THE IMPORTANCE OF DIATOM CELL SIZE IN COMMUNITY ANALYSIS1

The large variation in size and shape in diatoms is shown by morphometric measurements of 515 benthic and pelagic diatom species from the Baltic Sea area. The largest mean cell dimension (mostly the apical axis) varied between 4.2 and 653 μm, cell surface area between 55 and 344,000 μm², and cell volume between 21 and 14.2 × 10⁶μm³. The shape‐related index, length to width ratio, was between 1.0 and 63.3 and the shape‐ and size‐related index, surface area to volume ratio, was between 0.02 and 3.13. Diatom community analysis by multivariate statistics is usually based on counts of a fixed number of diatom valves with species scores irrespective of cell size. This procedure underestimates the large species for two reasons. First, the importance of a species with higher cell volume is usually larger in a community. Second, larger species usually have lower abundances and their occurrence in the diatom counts is stochastic. This article shows that co‐occurring small and large diatom species can respond very differently to environmental constraints. Large epiphytic diatoms responded most to macroalgal host species and small epiphytic diatoms most to environmental conditions at the sampling site. Large epilithic diatoms responded strongly to salinity, whereas small epilithic diatoms did so less clearly. The conclusion is that different scale‐dependent responses are possible within one data set. The results from the test data also show that important ecological information from diatom data can be missed when the large species are neglected or underestimated.     

Environmental controls of diatom species in northeast Pacific sediments

Non-linear, unimodal techniques are suitable for estimation of environmental properties in the northeast (NE) Pacific based on fossil diatom taxa (species and/or species groups) found in modern (core-top) sediments. Canonical Correspondence Analysis (CCA), a constrained ordination technique, discerned best-fit relationships among two multivariate datasets (the floral and the environmental) and thus yielded insight into the environmental variables that best explain the species variance within diatom populations. Based on these insights, we developed predictive functions for annual Primary Productivity (PP) and seasonal range of sea Sea-Surface Temperature using unimodal models and cross validation techniques. Estimates of annual PP (r²_jack = 0.92; RMSEP = 91.94 gC/m²/y) explained the highest percentage of variance in the core-top diatom record (22.1%).     

Diatom (Bacillariophyceae) assemblages in salt marshes of south‐central Chile: Relations with tidal inundation time and salinity

To reconstruct sea‐level history from changes in tidal environments using diatom assemblages, we need to better understand the relations among brackish diatom assemblages and changing environments along elevational gradients from diverse coastal sites. Our statistical analysis reveals relations between environmental variables and brackish benthic diatom assemblages in the little studied region of south‐central Chile. Along four transects across salt marshes at two sites, we identified 224 diatom taxa in 112 samples. Detrended canonical correspondence analysis showed that tidal exposure time index and salinity were appropriately regressed against the abundance of diatom species using unimodal‐based methods. Our tests of classical and inverse regressions of weighted average and weighted averaging partial least squares (WA‐PLS) showed that WA‐PLS resulted in the highest coefficient of determination and the lowest root‐mean square of the error of prediction. Our regression will be useful in reconstructing environmental variables from fossil diatom assemblages in Chile.     

Laurentian Great Lakes Phytoplankton and Their Water Quality Characteristics,Including a Diatom-Based Model for Paleoreconstruction of Phosphorus

Recent shifts in water quality and food web characteristics driven by anthropogenic impacts on the Laurentian Great Lakes warranted an examination of pelagic primary producers as tracers of environmental change. The distributions of the 263 common phytoplankton taxa were related to water quality variables to determine taxon-specific responses that may be useful in indicator models. A detailed checklist of taxa and their environmental optima are provided. Multivariate analyses indicated a strong relationship between total phosphorus (TP) and patterns in the diatom assemblages across the Great Lakes. Of the 118 common diatom taxa, 90 (76%) had a directional response along the TP gradient. We further evaluated a diatom-based transfer function for TP based on the weighted-average abundance of taxa, assuming unimodal distributions along the TP gradient. The r² between observed and inferred TP in the training dataset was 0.79. Substantial spatial and environmental autocorrelation within the training set of samples justified the need for further model validation. A randomization procedure indicated that the actual transfer function consistently performed better than functions based on reshuffled environmental data. Further, TP was minimally confounded by other environmental variables, as indicated by the relatively large amount of unique variance in the diatoms explained by TP. We demonstrated the effectiveness of the transfer function by hindcasting TP concentrations using fossil diatom assemblages in a Lake Superior sediment core. Passive, multivariate analysis of the fossil samples against the training set indicated that phosphorus was a strong determinant of historical diatom assemblages, verifying that the transfer function was suited to reconstruct past TP in Lake Superior. Collectively, these results showed that phytoplankton coefficients for water quality can be robust indicators of Great Lakes pelagic condition. The diatom-based transfer function can be used in lake management when retrospective data are needed for tracking long-term degradation, remediation and trajectories.     

Consistent abundance distributions of marine fishes in an old,climatically buffered,infertile seascape

Aim Macroecological theory predicts that along direct physiological gradients there will be unimodal abundance distributions of species and consistent rates of assemblage turnover. However, the majority of marine studies that have investigated the realized distribution of species along latitudinal or temperature gradients have generally found unimodal distributions to be rare. We assess fish distributions along a temperature gradient in a stable oligotrophic seascape and suggest that unimodal distributions will be more common. Location Nearshore demersal fish habitat extending 1500 km along the coast of south‐western Australia. Methods The relative abundances of demersal fish species were sampled off the coast of south‐western Australia along a temperature gradient. The confounding influence of other environmental variables was tested, and the assemblage was found to be highly correlated with temperature. For the 20 most abundant species, quantile regression spline models were used to construct a model within which 95% of their abundance was expected to fall. We compared the results from this study with the proportion of unimodal species abundance distributions observed in other studies. Results Of the 20 most abundant species, 19 displayed patterns that indicated temperature was an important factor influencing their range and relative abundance; with 15 species exhibiting unimodal abundance distributions, four having ramped distribution to one end of the sampled range and one showing no consistent pattern. Main conclusions The high diversity and percentage of endemic species in terrestrial and marine habitats of south‐western Australia is likely to be due to the stable geological and oceanographic history of the region. In comparison, studies of abundance distribution in other marine systems have been conducted in relatively heterogeneous and productive environments. The old, climatically buffered, oligotrophic seascape of south‐western Australia has provided a simple system in which the consistent influence of physiological gradients on the abundance distribution of fish species can be observed.     

Structure and diversity of intertidal benthic diatom assemblages in contrasting shores: a case study from the Tagus estuary1

The structure of intertidal benthic diatoms assemblages in the Tagus estuary was investigated during a 2‐year survey, carried out in six stations with different sediment texture. Nonparametric multivariate analyses were used to characterize spatial and temporal patterns of the assemblages and to link them to the measured environmental variables. In addition, diversity and other features related to community physiognomy, such as size‐class or life‐form distributions, were used to describe the diatom assemblages. A total of 183 diatom taxa were identified during cell counts and their biovolume was determined. Differences between stations (analysis of similarity (ANOSIM), R = 0.932) were more evident than temporal patterns (R = 0.308) and mud content alone was the environmental variable most correlated to the biotic data (BEST, ρ = 0.863). Mudflat stations were typically colonized by low diversity diatom assemblages (H′ ~ 1.9), mainly composed of medium‐sized motile epipelic species (250–1,000 μm³), that showed species‐specific seasonal blooms (e.g., Navicula gregaria Donkin). Sandy stations had more complex and diverse diatom assemblages (H′ ~ 3.2). They were mostly composed by a large set of minute epipsammic species (<250 μm³) that, generally, did not show temporal patterns. The structure of intertidal diatom assemblages was largely defined by the interplay between epipelon and epipsammon, and its diversity was explained within the framework of the Intermediate Disturbance Hypothesis. However, the spatial distribution of epipelic and epipsammic life‐forms showed that the definition of both functional groups should not be over‐simplified.     

The evolutionary species pool hypothesis and patterns of freshwater diatom diversity along a pH gradient

Aim To interpret the unimodal relationship between diatom species richness and lake pH within the context of the evolutionary species pool hypothesis (SPH). We test the following primary prediction arising from the SPH: the size of the potential species pool (PSP) will increase along a gradient representing the historical commonness of different pH environments (pH commonness). To do this we assume that the present‐day spatial dominance of near‐neutral pH conditions compared with acidic and alkaline conditions reliably mimics the relative spatial availabilities of historical pH conditions among freshwater lakes. We also determine whether local richness represents a constant proportion of PSP size along the pH commonness gradient. Location Two hundred and thirty‐four lakes distributed over a 405,000 km² region of the north‐eastern United States of America. Methods Sediment diatom morphospecies lists and pH data were acquired from the US Environmental Protection Agency's Environmental Monitoring and Assessment Program (EMAP) website. Using 248 morphospecies that occurred in at least 10 of the 234 lakes, four different measures of PSPs were calculated along the pH gradient. Local species richness was equated with the number of species occurring within the lake. Alpha diversity was equated with the average species richness of lakes with similar pH values. A combination of statistical methods were employed, including correlations, quadratic regression and piecewise regression. Results PSP size increased significantly with pH commonness for all four measures of PSP size, thus supporting the primary prediction of the evolutionary SPH. Local richness comprised a larger proportion of the PSP within acidic lakes than within circumneutral lakes. Alpha diversity and lake species richness both increased significantly with pH commonness, but the former did so in a two‐step fashion. We test and reject several alternative contemporary time‐scale explanations for our findings. Main Conclusions Our findings are consistent with the hypothesis that diatom taxonomic richness is presently lower within acidic and highly alkaline lakes than in circumneutral lakes owing to the limited opportunity in space and/or time for the evolution of suitably adapted species. Whereas ecological processes can explain why certain species are excluded from particular habitats, e.g. acidic lakes, they cannot account for why so few species are adapted to those habitats in the first place.     

Freshwater diatom biogeography in the Canadian Arctic Archipelago

Aim Document the biogeographical distributions of diatoms in the Canadian Arctic in relation to environmental factors. Location The Canadian Arctic Archipelago. Methods Diatoms were extracted from lake sediments and treated using standard methods. Rarefaction‐estimated species richness, diatom concentrations (valves cc^?1), and diatom abundance were mapped using a Geographic Information System. The physical and chemical parameters of the lakes were measured. Results A total of 326 taxa from 63 genera were found in 62 lakes of the study area. Up to 85 and as low as eight taxa were identified in any one lake, and rarefaction‐estimated species richness correlated with lake size. Nearby lakes could have greatly different diatom assemblages. Many taxa showed limited distributions. Response surfaces and ordination techniques indicate that a large number of taxa prefer colder regions of the Arctic while the abundance of others was influenced by lake water chemical parameters such as TKN and SiO₂. Main conclusions Although many taxa were common and found across the study area, diatom assemblages showed regional differences in the Arctic. Some taxa were not found in either the northern or southern parts of the Archipelago and others were restricted to particular regions. Newly delineated genera showed interpretable geographical patterns and could be related to environmental factors, suggesting that this more natural grouping may enhance our understanding of diatom ecology. Geographical, physical, and chemical factors are needed to explain diatom distributions in the Arctic.     

How many freshwater diatoms are pH specialists? A response to Pither & Aarssen (2005)

Pither & Aarssen (2005) propose a null model approach to assess the proportion of niche specialist taxa along ecological gradients. They apply this methodology to a large data set of lacustrine diatom assemblages and conclude that a majority of the taxa are generalists on a pH gradient. This conflicts with previous work, which shows that many diatom taxa have a statistically significant relationship with pH. We demonstrate the methods used by Pither & Aarssen (2005) have a high Type II error for rare taxa, and that this problem is compounded by the non-uniform sampling of the pH gradient which effectively precludes acid-lake specialist diatoms from being recognized as such. We re-analyse the data used by Pither & Aarssen (2005) and show that most of the diatoms have a statistically significant relationship with pH, and we thus refute their conclusions that few diatom species are specialists.     

Development of diatom-based salinity models for paleoclimatic research from lakes in British Columbia (Canada)

Diatoms were identified and enumerated from the surface sediments of 65 lakes located on the Cariboo and Chilcotin Plateaux (British Columbia, Canada). These lakes span a large gradient in lakewater ionic concentration (fresh through hypersaline) and composition, as well as other physical/chemical variables. Almost all of the study lakes had higher salinities in the late-summer than in the spring. The lakes with spring salinities >8 g l^–1 showed the largest seasonal increases in salinity. Ionic composition was similar in the spring and late-summer for most lakes. Both ionic concentration (i.e. salinity) and composition were important environmental variables that could account for the different diatom floras in the lakes. Diatom assemblages characteristic of carbonate-dominated and sulfate-dominated waters were identified. Other variables such as water depth and phosphorus concentration were also important.The majority (87%) of diatom taxa had estimated salinity optima < 3 g l^–1 Halophilic diatom taxa had broader tolerances to salinity when compared to the fresh water taxa, however taxa with narrow and broad tolerances could be identified across the salinity gradient. Species diversity was weakly but significantly correlated to lakewater salinity (r ² = 0.18 to 0.3, P < 0.05).Salinity inference models were developed based on the relationship between the diatom assemblages and the spring, late-summer and average salinity. The correlations between the measured and diatominferred salinity, based on the spring (r = 0.95), late-summer (r = 0.94) and average (r = 0.95) salinity data, are high because there was an extremely strong correlation (r = 0.98) between the log transformed spring and late-summer measured salinities. These salinity reconstruction models provide a tool that can be used to infer past climatic changes as part of paleolimnological studies from appropriate closed-basin lakes in British Columbia.     

Contrasting patterns in elevational diversity between microorganisms and macroorganisms

Aim Data and analyses of elevational gradients in diversity have been central to the development and evaluation of a range of general theories of biodiversity. Elevational diversity patterns have, however, been severely understudied for microbes, which often represent decomposer subsystems. Consequently, generalities in the patterns of elevational diversity across different trophic levels remain poorly understood. Our aim was to examine elevational gradients in the diversity of macroinvertebrates, diatoms and bacteria along a stony stream that covered a large elevational gradient. Location Laojun Mountain, Yunnan province, China. Methods The sampling scheme included 26 sites spaced at elevational intervals of 89 m from 1820 to 4050 m elevation along a stony stream. Macroinvertebrate and diatom richness were determined based on the morphology of the specimens. Taxonomic richness for bacteria was quantified using a molecular fingerprinting method. Over 50 environmental variables were measured at each site to quantify environmental variables that could correlate with the patterns of diversity. We used eigenvector‐based spatial filters with multiple regressions to account for spatial autocorrelation. Results The bacterial richness followed an unexpected monotonic increase with elevation. Diatoms decreased monotonically, and macroinvertebrate richness showed a clear unimodal pattern with elevation. The unimodal richness pattern for macroinvertebrates was best explained by the mid‐domain effect (r² = 0.72). The diatom richness was best explained by the variation in nutrient supply, and the increase in bacterial richness with elevation may be related to an increased carbon supply. Main conclusions We found contrasting patterns in elevational diversity among the three studied multi‐trophic groups comprising unicellular and multicellular aquatic taxa. We also found that there may be fundamental differences in the mechanisms underlying these species diversity patterns.     

Declines in littoral species richness across both spatial and temporal nutrient gradients: a palaeolimnological study of two taxonomic groups

1. Using a palaeolimnological approach in shallow lakes, we quantified the species richness responses of diatoms and Cladocera to phosphorus enrichment. We also examined differences in species richness responses between littoral and pelagic assemblages of our focal communities. To address both spatial and temporal relationships, our study includes an analysis of both surface sediments from 40 lakes and of a lake sediment record spanning c. 120 years. The objective of our study was to determine whether similar species richness patterns occurred across trophic levels, as well as along spatial and temporal gradients. 2. We found that both diatom and Cladocera species richness estimates significantly declined with increasing phosphorus across space and through time. When the assemblages were subdivided according to known habitat preferences, littoral biodiversity maintained a negative trend, whereas pelagic species richness tended to show no relationship with phosphorus. 3. Negative productivity–diversity patterns have been observed across almost all palaeolimnological studies that span large productivity gradients. This congruence in patterns is most likely due to the similarity in data collection methods and in focal communities studied. The contrasting responses between littoral and pelagic assemblages may be explained by the differences in physical habitat and the pool of taxa in each of these environments. Consistent with the literature, we found statistical support for the idea that littoral diversity declines could be explained by an interaction between macrophytes and nutrients along strong trophic gradients. The general lack of a diversity response in our pelagic assemblages could be attributable to the limited pool of subfossil taxa. The response of the pelagic diatom could also be related to their broad range of nutrient tolerances. 4. The observed negative response of species richness to phosphorus enrichment, particularly in the littoral assemblages, has implications for ecosystems functioning because communities with reduced biodiversity often are less resilient to anthropogenic change.     

Is salinity an obstacle for biological invasions?

Invasions of freshwater habitats by marine and brackish species have become more frequent in recent years with many of those species originating from the Ponto‐Caspian region. Populations of Ponto‐Caspian species have successfully established in the North and Baltic Seas and their adjoining rivers, as well as in the Great Lakes–St. Lawrence River region. To determine if Ponto‐Caspian taxa more readily acclimatize to and colonize diverse salinity habitats than taxa from other regions, we conducted laboratory experiments on 22 populations of eight gammarid species native to the Ponto‐Caspian, Northern European and Great Lakes–St. Lawrence River regions. In addition, we conducted a literature search to survey salinity ranges of these species worldwide. Finally, to explore evolutionary relationships among examined species and their populations, we sequenced the mitochondrial cytochrome c oxidase subunit I gene (COI) from individuals used for our experiments. Our study revealed that all tested populations tolerate wide ranges of salinity, however, different patterns arose among species from different regions. Ponto‐Caspian taxa showed lower mortality in fresh water, while Northern European taxa showed lower mortality in fully marine conditions. Genetic analyses showed evolutionary divergence among species from different regions. Due to the geological history of the two regions, as well as high tolerance of Ponto‐Caspian species to fresh water, whereas Northern European species are more tolerant of fully marine conditions, we suggest that species originating from the Ponto‐Caspian and Northern European regions may be adapted to freshwater and marine environments, respectively. Consequently, the perception that Ponto‐Caspian species are more successful colonizers might be biased by the fact that areas with highest introduction frequency of NIS (i.e., shipping ports) are environmentally variable habitats which often include freshwater conditions that cannot be tolerated by euryhaline taxa of marine origin.     

DIATOM ASSOCIATIONS IN YAQUINA ESTUARY,OREGON: A MULTIVARIATE ANALYSIS1

A cluster analysis and a principal components analysis were performed to examine the degree to which the diatom flora of Yaquina estuary can be partitioned into discrete associations and to relate the composition of the flora to selected physical properties of the estuary. The species composition of the diatom flora during the winter was more closely associated with salinity and intertidal exposure them in the summer. A cluster of freshwater taxa obtained for the winter data was the most conspicuous diatom association defined by the cluster analysis. Although taxa within a cluster had more similar distributions than taxa from different clusters, there was considerable variation in the distributions of marine and brackish water taxa within a multispecies cluster.     

Quantifying species indicator values for trophic diatom indices: a comparison of approaches

This study compares two approaches for constructing diatom-based indices for monitoring river eutrophication. The first approach is based on weighted averaging of species indicator values with the underlying assumption that species have symmetrical unimodal distributions along the nutrient gradient, and their distributions are sufficiently described by a single indicator value per species. The second approach uses multiple indicator values for individual taxa and is based on the possibility that species have complex asymmetrical response curves. Multiple indicator values represent relative probabilities that a species would be found within certain ranges of nutrient concentration. We used 155 benthic diatom samples collected from rivers in the Northern Piedmont ecoregion (Northeastern U.S.A.) to construct two datasets: one used for developing models and indices, and another for testing them. To characterize the shape of species response curves we analyzed changes in the relative abundance of 118 diatom taxa common in this dataset along the total phosphorus (TP) gradient by fitting parametric and non-parametric regression models. We found that only 34 diatoms had symmetrical unimodal response to TP. Among several indices that use a single indicator value for each species, the best was the weighted averaging partial least square (WA-PLS) inference model. The correlation coefficient between observed and inferred TP in the test dataset was 0.67. The best index that employed multiple indicator values for each species had approximately the same predictive power as the WA-PLS based index, but in addition, this index provided a sample-specific measure of uncertainty for the TP estimation.     

Consistent patterns in diatom assemblages and diversity measures across water‐depth gradients from eight Boreal lakes from north‐western Ontario (Canada)

1. Until recently, the distribution of diatom species assemblages and their attributes (e.g. species richness and evenness) in relation to water depth have been identified but not quantified, especially across several lakes in a region. Here, we examined diatom assemblages in the surface sediment across a water‐depth gradient in eight small, boreal lakes in north‐western Ontario, minimally disturbed by human activities. 2. Surface‐sediment diatom assemblages were collected within each lake along a gentle slope from near‐shore to the centre deep basin of the lake, at a resolution of ～1 m water depth. Analysis of sedimentary samples provided an integrated view of assemblages that were living in the lake over several years and enabled a high‐resolution analysis of many lakes. The study lakes ranged in water chemistry, morphology and size and are located along an east–west transect approximately 250 km long in north‐western Ontario (Canada). 3. The majority of diatom species were distributed along a continuum of depth, with those taxa having similar habitat requirements forming distinct, though overlapping, assemblages. Three major zones of diatom assemblages in each lake were consistently identified: (i) a near‐shore assemblage of Achnanthes (sensu lato), Nitzschia, Cymbella (sensu lato) and other benthic species; (ii) a mid‐depth assemblage of small Fragilaria (sensu lato)/small Aulacoseira and various Navicula taxa; and (iii) a deep‐water assemblage of planktonic origin (mainly Discotella spp.). 4. The depth of the transition between assemblage zones varied between the eight lakes. The boundary between the deep‐water planktonic zone and the mid‐depth benthic zone varied according to water chemistry and was probably related to light attenuation. The boundary was deeper in lakes with the lower dissolved organic carbon and total phosphorus (TP) (i.e. less light attenuation) and vice versa. 5. Generally, species richness, species evenness and turnover rate of species as a function of depth were significantly lower in the planktonic assemblage zone in comparison with the two zones nearer the shore. Reproducibility of species and assemblage distributions across the depth gradient of the lakes illustrated that, despite potential for sediment transport, detailed ecological characterisation of diatom species can be gleaned from sedimentary data. Such data are often lacking, particularly for near‐shore benthic species.     

Fifteen important questions in the spatial ecology of diatoms

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DISTRIBUTION OF DIATOMS IN THE PLANKTON OF YAQUINA ESTUARY,OREGON1

The distributional patterns of diatoms in the plankton of the Yaqnina Estuary, Oregon, were, investigated and related to selected climatic and hydrographic factors. Distribution was strongly influenced by seasonal patterns of rainfall resulting in the introduction of a large volume of fresh water into the estuary during fall and winter. Plankton assemblages in spring, summer and fall had fewer diatom species and exhibited a more rapid rate of change in species composition than in winter. Winter assemblages were further characterized by many pennate diatoms, apparently dislodged from the benthos during periods of high freshwater discharge and silt loads. A statistical measure of community difference indicated an increase in taxonomic homogeneity among assemblages throughout the estuary with the onset of the rainy season in late fall and a gradual transition to a more heterogeneous system again during late spring. Canonical correlation ordered 20 prominent diatom taxa along the salinity gradient and identified possible relationships among certain taxa and selected environmental variables, namely visible light energy and temperature. Redundancy in the species data given the environmental data was only 40%, emphasizing the difficulty in demonstrating a quantitive relationship between plankton dynamics in the field and concurrent measurements of chemical and physical variables.     

THE DISTRIBUTION OF EDAPHIC DIATOMS ALONG ENVIRONMENTAL GRADIENTS OF A LOUISIANA SALT MARSH1

Edaphic (sediment-associated) diatoms were collected from five sites along a gradient from slightly brackish to saline marsh on 30 November 1977 and 29 April 1978. Several environmental factors change along this gradient in addition to salinity, including tidal flushing and percent organic content of the soil. A total of 112 taxa were identified, with Amphora exigua Greg., Navicula phyllepta Kütz., N. salinarum Grun., and N. tripunctata (Müll.) Bory being among the five most abundant taxa on both sampling dates. The taxonomic composition of the diatom flora was very similar to those observed for other salt marshes located along the Gulf and Atlantic coasts. Diatom taxa were generally euryhaline and distributed continuously along the gradient from brackish to saline marsh. Species diversity (H¹) and the number of taxa in a sample (S) were also similar to those reported for other North American salt marshes. The number of taxa in a sample was higher in the saline than in the brackish area on both sampling dates while species diversity was greater in the saline area only in April.     

Plant species richness and diversity along an altitudinal gradient in the Sierra Nevada, Mexico

This article presents an analysis of plant species richness and diversity and its association with climatic and soil variables along a 1300‐m elevation gradient on the Cerro Tláloc Mountain in the northern Sierra Nevada in Mexico. Two 1000‐m² tree sampling plots were created at each of 21 selected sampling sites, as well as two 250‐m² plots for shrubs and six 9‐m² plots for herbaceous plants. Species richness and diversity were estimated for each plant life form, and beta diversity between sites was estimated along the gradient. The relationship between species richness and diversity and environmental variables was modelled using simple linear correlation and regression trees. Species richness and diversity showed a unimodal pattern with a bias towards high values in the lower half of the elevation gradient under study. This response was consistent for all three life forms. Beta diversity increased steadily along the elevation gradient, being lower between contiguous sites at intermediate elevations and high – the species replacement rate was nearly 100%– between sites at the extremes of the gradient. Few species were adapted to the full spectrum of environmental variation along the elevation gradient studied. The regression tree suggests that differences in species richness are mainly influenced by elevation (temperature and humidity) and soil variables, namely A₂ permanent wilting point, organic matter and horizon field capacity and A₁ horizon Mg²⁺.