An evaluation of environmental factors affecting species distributions

Many different models can be built to explain the distributions of species. Often there is no single model that is clearly better than the alternatives, and this leads to uncertainty over which environmental factors are limiting species’ distributions. We investigated the support for different environmental factors by determining the drop in model performance when selected predictors were excluded from the model building process. We used a paired t-test over 37 plant species so that an environmental factor was only deemed significant if it consistently improved the results for multiple species. Geology and winter minimum temperatures were found to be the environmental factors with the most support, with a significant drop in model performance when either of these factors was excluded. However, there was less support for summer maximum temperature, as other environmental factors could combine to produce similar model performance. Our method of evaluating environmental factors using multiple species will not be capable of detecting predictors that are only important for one or two species, but it is difficult to distinguish these from spurious correlations. The strength of the method is that it increases inference for factors that consistently affect the distributions of many species. We discourage the assessment of models against predefined benchmarks, such as an area under the curve (AUC) of more than 0.7, as many alternative models for the same species produce similar results. Therefore, the benchmarks do not provide any indication of how the performance of the selected model compares to alternative models, and they provide weak inference to accept any selected model.     

Effects of life‐history requirements on the distribution of a threatened reptile

Survival and reproduction are the two primary life‐history traits essential for species’ persistence; however, the environmental conditions that support each of these traits may not be the same. Despite this, reproductive requirements are seldom considered when estimating species’ potential distributions. We sought to examine potentially limiting environmental factors influencing the distribution of an oviparous reptile of conservation concern with respect to the species’ survival and reproduction and to assess the implications of the species’ predicted climatic constraints on current conservation practices. We used ecological niche modeling to predict the probability of environmental suitability for the alligator snapping turtle (Macrochelys temminckii). We built an annual climate model to examine survival and a nesting climate model to examine reproduction. We combined incubation temperature requirements, products of modeled soil temperature data, and our estimated distributions to determine whether embryonic development constrained the northern distribution of the species. Low annual precipitation constrained the western distribution of alligator snapping turtles, whereas the northern distribution was constrained by thermal requirements during embryonic development. Only a portion of the geographic range predicted to have a high probability of suitability for alligator snapping turtle survival was estimated to be capable of supporting successful embryonic development. Historic occurrence records suggest adult alligator snapping turtles can survive in regions with colder climes than those associated with consistent and successful production of offspring. Estimated egg‐incubation requirements indicated that current reintroductions at the northern edge of the species’ range are within reproductively viable environmental conditions. Our results highlight the importance of considering survival and reproduction when estimating species’ ecological niches, implicating conservation plans, and benefits of incorporating physiological data when evaluating species’ distributions.     

Science or Slaughter: Need for Lethal Sampling of Sharks

Abstract: General consensus among scientists, commercial interests, and the public regarding the status of shark populations is leading to an increasing need for the scientific community to provide information to help guide effective management and conservation actions. Experience from other marine vertebrate taxa suggests that public, political, and media pressures will play an increasingly important part in setting research, management, and conservation priorities. We examined the potential implications of nonscientific influences on shark research. In particular, we considered whether lethal research sampling of sharks is justified. Although lethal sampling comes at a cost to a population, especially for threatened species, the conservation benefits from well‐designed studies provide essential data that cannot be collected currently in any other way. Methods that enable nonlethal collection of life‐history data on sharks are being developed (e.g., use of blood samples to detect maturity), but in the near future they will not provide widespread or significant benefits. Development of these techniques needs to continue, as does the way in which scientists coordinate their use of material collected during lethal sampling. For almost half of the known shark species there are insufficient data to determine their population status; thus, there is an ongoing need for further collection of scientific data to ensure all shark populations have a future. Shark populations will benefit most when decisions about the use of lethal sampling are made on the basis of scientific evidence that is free from individual, political, public, and media pressures.     

Variation in Pesticide Tolerance of Tadpoles among and within Species of Ranidae and Patterns of Amphibian Decline

Abstract: There is significant variation among and within amphibian species with respect to reports of population decline; declining species are often found in environments that are physiograpically similar to environments where the same species is thriving. Because variability exists among organisms in their sensitivity to environmental stressors, it is important to determine the degree of this variation when undertaking conservation efforts. We conducted both lethal (time-to-death) and sublethal (activity change) assays to determine the degree of variation in the sensitivity of tadpoles to a pesticide, carbaryl, at three hierarchical levels: among ranid species, among several populations of a single ranid species (   Rana sphenocephala ), and within populations of R. sphenocephala . We observed significant variation in time to death among the nine ranid species and among the 10 R. sphenocephala populations we tested. Four out of eight R. sphenocephala populations exhibited significantly different times to death among families. The magnitude of the activity change in response to exposure to sublethal carbaryl levels was significantly different among species and within R. sphenocephala populations. Chemical contamination, at lethal or sublethal levels, can alter natural regulatory processes such as juvenile recruitment in amphibian populations and should be considered a contributing cause of declines in amphibian populations.     

The evaluation strip: A new and robust method for plotting predicted responses from species distribution models

Increasing use is being made in conservation management of statistical models that couple extensive collections of species and environmental data to make predictions of the geographic distributions of species. While the relationships fitted between a species and its environment are relatively transparent for many of these modeling techniques, others are more ‘black box’ in character, only producing geographic predictions and providing minimal or untraditional summaries of the fitted relationships on which these predictions are based. This in turn prevents robust evaluation of the ecological sensibility of such models, a necessary process if model predictions are to be treated with confidence. Here we propose a new but simple method for visualizing modeled responses that can be implemented with any modeling method, and demonstrate its application using five common methods applied to the prediction of an Australian tree species. This is achieved by insetting an “evaluation strip” into the spatial data layers, which, after predictions have been made, can be clipped out and used for creating plots of the modelled responses. We present findings of the application strip for algorithms GLMs, GAMs, CLIM, DOMAIN and MARS. Evaluation strips can be constructed to investigate either uni-variate responses, or the simultaneous variation in predicted values in relation to two variables. The latter option is particularly useful for evaluating responses in models that allow the fitting of complex interaction terms.     

Provenance, life span, and phylogeny do not affect grass species' responses to nitrogen and phosphorus

Successful conservation management requires an understanding of how species respond to intervention. Native and exotic species may respond differently to management interventions due to differences arising directly from their origin (i.e., provenance) or indirectly due to biased representations of different life history types (e.g., annual vs. perennial life span) or phylogenetic lineages among provenance (i.e., native or exotic origin) groups. Thus, selection of a successful management regime requires knowledge of the life history and provenance-bias in the local flora and an understanding of the interplay between species characteristics across existing environmental gradients in the landscape. Here we tested whether provenance, phylogeny, and life span interact to determine species distributions along natural gradients of soil chemistry (e.g., soil nitrogen and phosphorus) in 10 upland prairie sites along a 600-km latitudinal transect running from southern Vancouver Island in British Columbia, Canada, to the Willamette Valley in Oregon, USA. We found that soil nitrate, phosphorus, and pH exerted strong control over community composition. However, species distributions along environmental gradients were unrelated to provenance, life span, or phylogenetic groupings. We then used a greenhouse experiment to more precisely measure the response of common grass species to nitrogen and phosphorus supply. As with the field data, species responses to nutrient additions did not vary as a function of provenance, life span, or phylogeny. Native and exotic species differed strongly in the relationship between greenhouse-measured tolerance of low nutrients and field abundance. Native species with the greatest ability to maintain biomass production at low nutrient supply rates were most abundant in field surveys, as predicted by resource competition theory. In contrast, there was no relationship between exotic-species biomass at low nutrient levels and field abundance. The implications of these findings for management of invasive species are substantial in that they overturn a general belief that reduction of nutrient supplies favors native species. The idiosyncratic nature of species response to nutrients in this study suggests that manipulation of nutrient supplies is unlikely to alter the overall balance between native and exotic species, although it may well be useful to control specific exotic species.     

Multiple stressors and amphibian declines: dual impacts of pesticides and fish on yellow-legged frogs.

More than 40% of Earth's 5700+ amphibian species have undergone recent declines. Despite the likely involvement of multiple factors in driving these declines, most studies continue to focus on single stressors. In California (USA), separate studies have implicated either introduced fish or pesticides as causal agents. To date, however, no study has simultaneously evaluated the respective roles of these two potential stressors nor attempted to assess their relative importance, information critical for the development of effective conservation efforts and environmental policies. We examined the role and relative effect of fish and pesticides on the mountain yellow-legged frog (Rana muscosa) using unusually detailed data sets for a large portion of R. muscosa's historic range in California's Sierra Nevada. Habitat characteristics and presence/absence of R. muscosa and fish were quantified at each of 6831 sites during field surveys. Pesticide use upwind of each site was calculated from pesticide application records and predominant wind directions. Using generalized additive models, we found that, after accounting for habitat effects, the probability of R. muscosa presence was significantly reduced by both fish and pesticides, with the landscape-scale effect of pesticides much stronger than that of fish. The degree to which a site was sheltered from the predominant wind (and associated pesticides) was also a significant predictor of R. muscosa presence. Taken together, these results represent the strongest evidence to date that windborne pesticides are contributing to amphibian declines in pristine locations. Our results suggest that amphibian declines may have complex multi-factorial causes, and caution that single-factor studies that demonstrate the importance of one factor should not be used as evidence against the importance of other factors.     

Patterns of growth,energy utilization and reproduction in some meso- and bathypelagic fishes off Southern California

We have studied growth, energy use and reproduction in 4 mesopelagic fishes and 5 bathypelagic fishes living off Southern California (USA). All of the mesopelagic species underwent diurnal vertical migrations, while none of the bathypelagic species did so. The life histories of these pelagic fishes were compared among themselves and with epipelagic sardines and anchovies studied by others. The epipelagic species had the highest growth rates (estimated from otoliths, expressed in standard length or kilocalories), the mesopelagic species had the lowest growth rates and the bathypelagic species had intermediate growth rates. The relatively rapid growth rates of the bathypelagic fishes were achieved by high relative growth efficiencies made possible by low metabolic rates. Of the species studied, the lifespans of the epipelagic and bathypelagic species ranged from 4 to 8 yr and the lifespans of mesopelagic species from 5 to 8 yr. Data on egg diameters suggest that the mesopelagic species first reproduce in their 3rd yr, while the bathypelagic species do so in their last year. Epipelagic fishes generally have a large size, rapid growth, long life and early, repeated reproduction. Mesopelagic fishes are characterized by small size, slow growth, long life and early, repeated reproduction. Bathypelagic fishes generally have large size, rapid growth, somewhat shorter lives and late reproduction, which is possible a single event. The latter pattern is evidently feasible only in a rather stable environment where juvenile survivorship would always display relatively low variability. Many unusual characteristics of deep-living animals have possibly been selected by factors peculiar to the environment; however, such characteristics are just as likely to have been selected by factors equally present in many other environments, but not expressed there due to masking selective forces. In particular, we have in mind the darkness, stability and homogeneity of the bathypelagic realm as phenomena which represent the effective absence of many selective forces.The first two authors contributed equally and their names are listed in alphabetical order     

Mating system,optimal number of matings,and sperm transfer in the Argentine ant Iridomyrmex humilis

Summary In ants, because males have a finite sperm supply and females mate only at the beginning of their reproductive lives, it is possible to infer which is the limiting sex from a few parameters: the amount of sperm produced by males, the amount of sperm stored by females, and the numerical sex ratio. In the Argentine ant Iridomyrmex humilis mating takes place in the nest. Laboratory experiments and field data showed that the numerical sex ratio is heavily male-biased (10.1:1) and that the maximum number of sperm a female can store is similar to the number of sperm a male possesses. Thus females are the limiting sex in this species. In a set of mating experiments, one queen was presented with 1–20 males. The highest proportion of successful matings occurred when females were presented with two males. There was a significant negative correlation between the amount of sperm queens stored and the number of males present in the mating arena. This relationship most likely resulted from male interference during the copulation process. When several males were present in the arena, the mating pair was frequently disturbed by other males trying to copulate. Newly mated queens collected from the field stored 172,000 ± 76,000 sperm, a quantity most similar to that measured in laboratory mating experiments with a ratio of 5 or 10 males per queen. Because the operational sex ratio in I. humilis is highly male-biased, male interference may also decrease the amount of sperm queens store in the field. In many ants, fewer sperm stored by queens should decrease their reproductive success because they would run out of sperm earlier in their reproductive lifetimes. However, comparison of the amount of sperm present in young and old I. humilis queens collected in the field suggests that most use only a small proportion of their sperm supply during their lifetimes. Males mate once and discharge all their sperm during a single mating. Females may mate with several males but dissection of these males indicated that in most cases only one of them had empty seminal vesicles thus suggesting that a single male is responsible for most of the sperm transfer. Thus caution should be exercised in inferring multiple inseminations, as is frequently done in eusocial insects, from the observation of multiple copulations. Correspondence to: L. Keller     

Site‐Occupancy Distribution Modeling to Correct Population‐Trend Estimates Derived from Opportunistic Observations

Abstract: Species’ assessments must frequently be derived from opportunistic observations made by volunteers (i.e., citizen scientists). Interpretation of the resulting data to estimate population trends is plagued with problems, including teasing apart genuine population trends from variations in observation effort. We devised a way to correct for annual variation in effort when estimating trends in occupancy (species distribution) from faunal or floral databases of opportunistic observations. First, for all surveyed sites, detection histories (i.e., strings of detection–nondetection records) are generated. Within‐season replicate surveys provide information on the detectability of an occupied site. Detectability directly represents observation effort; hence, estimating detectablity means correcting for observation effort. Second, site‐occupancy models are applied directly to the detection‐history data set (i.e., without aggregation by site and year) to estimate detectability and species distribution (occupancy, i.e., the true proportion of sites where a species occurs). Site‐occupancy models also provide unbiased estimators of components of distributional change (i.e., colonization and extinction rates). We illustrate our method with data from a large citizen‐science project in Switzerland in which field ornithologists record opportunistic observations. We analyzed data collected on four species: the widespread Kingfisher (Alcedo atthis) and Sparrowhawk (Accipiter nisus) and the scarce Rock Thrush (Monticola saxatilis) and Wallcreeper (Tichodroma muraria). Our method requires that all observed species are recorded. Detectability was <1 and varied over the years. Simulations suggested some robustness, but we advocate recording complete species lists (checklists), rather than recording individual records of single species. The representation of observation effort with its effect on detectability provides a solution to the problem of differences in effort encountered when extracting trend information from haphazard observations. We expect our method is widely applicable for global biodiversity monitoring and modeling of species distributions.     

Generalists and specialists along a latitudinal transect: patterns of thermal adaptation in six species of damselflies

Tropical organisms colonizing temperate environments face reduced average temperatures and dramatic thermal fluctuations. Theoretical models postulate that thermal specialization should be favored either when little environmental variation is experienced within generations or when among-generation variation is small relative to within-generation variation. To test these predictions, we studied six temperate species of damselflies differing in latitudinal distribution. We developed a computer model simulating how organisms experience environmental variation (accounting for diapause and voltinism) and performed a laboratory experiment assaying thermal sensitivities of growth rates. The computer model showed opposing latitudinal trends in among- and within-generation thermal variability: within-generation thermal variability decreased toward higher latitudes, whereas relative levels of among-generation thermal variability peaked at midlatitudes (where a shift in voltinism occurred). The growth experiment showed that low-latitude species were more thermally generalized than mid- and high-latitude species, supporting the prediction that generalists are favored under high levels of within-generation variation. Northern species had steeper, near-exponential reaction norms suggestive of thermal specialization. However, they had strikingly high thermal optima and grew very slowly over most of the thermal range they are expected to experience in the field. This observation is at present difficult to explain. These results highlight the importance of considering interactions between life history and environmental variation when deriving expectations of thermal adaptation.     

Inbreeding Depression Accumulation across Life‐History Stages of the Endangered Takahe

Abstract: Studies evaluating the impact of inbreeding depression on population viability of threatened species tend to focus on the effects of inbreeding at a single life‐history stage (e.g., juvenile survival). We examined the effects of inbreeding across the full life‐history continuum, from survival up to adulthood, to subsequent reproductive success, and to the recruitment of second‐generation offspring, in wild Takahe ( Porphyrio hochstetteri ) by analyzing pedigree and fitness data collected over 21 breeding seasons. Although the effect size of inbreeding at individual life‐history stages was small, inbreeding depression accumulated across multiple life‐history stages and ultimately reduced long‐term fitness (i.e., successful recruitment of second‐generation offspring). The estimated total lethal equivalents (2B) summed across all life‐history stages were substantial (16.05, 95% CI 0.08–90.8) and equivalent to an 88% reduction in recruitment of second‐generation offspring for closely related pairs (e.g., sib–sib pairings) relative to unrelated pairs (according to the pedigree). A history of small population size in the Takahe could have contributed to partial purging of the genetic load and the low level of inbreeding depression detected at each single life‐history stage. Nevertheless, our results indicate that such “purged” populations can still exhibit substantial inbreeding depression, especially when small but negative fitness effects accumulate across the species’ life history. Because inbreeding depression can ultimately affect population viability of small, isolated populations, our results illustrate the importance of measuring the effects of inbreeding across the full life‐history continuum.     

Model based grouping of species across environmental gradients

We present a novel approach to the statistical analysis and prediction of multispecies data. The approach allows the simultaneous grouping and quantification of multiple species’ responses to environmental gradients. The underlying statistical model is a finite mixture model, where mixing is performed over the individual species’ responses to environmental gradients. Species with similar responses are grouped with minimal information loss. We term these groups species archetypes. Each species archetype has an associated GLM that can be used to predict distributions with appropriate measures of uncertainty. Initially, we illustrate the concept and method using artificial data and then with application to real data comprising 200 species from the Great Barrier Reef (GBR) lagoon on 13 oceanographic and geological gradients from 12°S to 24°S. The 200 species from the GBR are well represented by 15 species archetypes. The model is interpreted through maps of the probability of presence for a fine scale set of locations throughout the study area. Maps of uncertainty are also produced to provide statistical context. The presence of each species archetype was strongly influenced by oceanographic gradients, principally temperature, oxygen and salinity. The number of species in each group ranged from 4 to 34. The method has potential application to the analysis of multispecies distribution patterns and for multispecies management.     

建立室内实验程序用于量化鸟类对种子消耗量的杀虫剂处理效应

种衣剂中使用的杀虫剂会影响鸟类对种子的消耗量,从而影响鸟类的实际暴露风险。虽然量化这种对消耗量的影响目前不被考虑在环境风险评估中,但这是一个潜在的选择,例如在比较不同农药的暴露风险时应予以考虑。这种方法可以突出用于防止鸟类服用致死或亚致死剂量的杀虫剂。为了制定一个标准,我们在已建立的笔试方法的基础上开发了一个室内测试程序,包括饥饿期的“两选择”和“无选择”阶段。测试期间采用动物福利的最高标准。利用统计方法确定最合适的重复次数并分析最终结果。杀虫剂对种子消耗量的影响用处理后种子与未处理种子消耗的比率表示。该消耗因子可作为种子处理的鸟类风险评估中的回避因子。 作为案例,我们提出了一个获得绿雀（Carduelis chloris）和日本鹌鹑（Coturnix japonica）对未处理和使用杀真菌剂处理的油菜籽（Brassica napus）的种子和特定物种消耗量因子的试验程序。总体来说,两种鸟类的体质都没有受到试验程序的负面影响。试验程序适用于显示预期消耗模式的差异,例如在“两选择”试验中比在“无选择”试验中对经过杀真菌剂处理的种子有更高的避免性。而种子消耗量在不同物种和杀真菌剂处理组有差别,这使得我们可以对不同杀真菌剂的规避效应进行排序。使用本文所提出的标准程序可以针对更多鸟类物种得到具有可比性的农药和物种特定的消费因子。     

Evidence for life history changes in high-altitude populations of three perennial forbs

Relatively little is known about how the life histories of perennial forb species, and especially their lifetime patterns of growth, vary across environmental gradients. We used a post hoc approach (herb-chronology) to determine plant age and previous growth (width of successive annual rings in roots) in three species of perennial forb (two long-lived species [Penstemon venustus, Lupinus laxiflorus] and one short-lived [Rudbeckia occidentalis]) along a 1000-m altitudinal gradient in the Wallowa Mountains (northeast Oregon, USA). Plants from the highest altitude tended to be considerably older and produced up to five times as many flowering shoots as lowland plants. In addition, mean ring widths of high-altitude plants were about half those of lowland plants. In plants from low and intermediate altitudes, ring width either decreased linearly or varied inconsistently during the life of the plant. In contrast, ring widths of high-altitude plants increased at first and later decreased, resulting in curvilinear growth trajectories that were highly consistent among species. Together, these data for three ecologically distinct forb species provide evidence of a consistent shift toward more conservative and strongly constrained life histories at higher altitudes. More generally, the results indicate the possible importance of changes in selection pressures across strong environmental gradients on life history strategies within a single species.     

Effects of coral transplantation in sites of varying distances and environmental conditions

Several scleractinian coral species with different growth forms and life history strategies were studied in terms of colony growth (expressed as projected linear increment) and survivorship over a range of distances and environmental conditions in the Philippines. The experimental design consisted of 1 m² plots grouped within a reef site, to several sites within reef systems separated by a distance of about 340 km. There were distinct differences among species, with submassive and massive forms displaying slower growth but better survival, confirming results of other studies. They probably play the role of framework builders of the reef. In contrast, the delicate branching and foliose species had higher growth rates but poor survivorship. This observation, plus their ease of fragmentation, suggests they act more as fillers of the reef matrix. There was high variability in colony increment of a species among the square meter plots, but not among sites within a reef system. Thus, more regular pattern could be observed at this level. In contrast to growth, survivorship differed significantly among sites, being lowest in the site which harbored the greatest amount of dead coral. Growth and survival, however, are not sufficient performance measures to evaluate the success of coral transplantation. Reproduction and subsequent recruitment must also be taken into account. It is recommended that coral transplant and restoration studies consider the broad environmental context of restoration and seek to develop assembly rules that will allow practitioners to match coral types and sequence of interventions to each unique context.     

Maximum entropy modeling of species geographic distributions

The availability of detailed environmental data, together with inexpensive and powerful computers, has fueled a rapid increase in predictive modeling of species environmental requirements and geographic distributions. For some species, detailed presence/absence occurrence data are available, allowing the use of a variety of standard statistical techniques. However, absence data are not available for most species. In this paper, we introduce the use of the maximum entropy method (Maxent) for modeling species geographic distributions with presence-only data. Maxent is a general-purpose machine learning method with a simple and precise mathematical formulation, and it has a number of aspects that make it well-suited for species distribution modeling. In order to investigate the efficacy of the method, here we perform a continental-scale case study using two Neotropical mammals: a lowland species of sloth, Bradypus variegatus, and a small montane murid rodent, Microryzomys minutus. We compared Maxent predictions with those of a commonly used presence-only modeling method, the Genetic Algorithm for Rule-Set Prediction (GARP). We made predictions on 10 random subsets of the occurrence records for both species, and then used the remaining localities for testing. Both algorithms provided reasonable estimates of the species’ range, far superior to the shaded outline maps available in field guides. All models were significantly better than random in both binomial tests of omission and receiver operating characteristic (ROC) analyses. The area under the ROC curve (AUC) was almost always higher for Maxent, indicating better discrimination of suitable versus unsuitable areas for the species. The Maxent modeling approach can be used in its present form for many applications with presence-only datasets, and merits further research and development.     

GIS-based niche modeling for mapping species' habitat

Ecological "niche modeling" using presence-only locality data and large-scale environmental variables provides a powerful tool for identifying and mapping suitable habitat for species over large spatial extents. We describe a niche modeling approach that identifies a minimum (rather than an optimum) set of basic habitat requirements for a species, based on the assumption that constant environmental relationships in a species' distribution (i.e., variables that maintain a consistent value where the species occurs) are most likely to be associated with limiting factors. Environmental variables that take on a wide range of values where a species occurs are less informative because they do not limit a species' distribution, at least over the range of variation sampled. This approach is operationalized by partitioning Mahalanobis D2 (standardized difference between values of a set of environmental variables for any point and mean values for those same variables calculated from all points at which a species was detected) into independent components. The smallest of these components represents the linear combination of variables with minimum variance; increasingly larger components represent larger variances and are increasingly less limiting. We illustrate this approach using the California Gnatcatcher (Polioptila californica Brewster) and provide SAS code to implement it.     

Algorithms and biplots for double constrained correspondence analysis

Correspondence analysis with linear external constraints on both the rows and the columns has been mentioned in the ecological literature, but lacks full mathematical treatment and easily available algorithms and software. This paper fills this gap by defining the method as maximizing the fourth-corner correlation between linear combinations, by providing novel algorithms, which demonstrate relationships with related methods, and by making a detailed study of possible biplots and associated approximations. The method is illustrated using ecological data on the abundances of species in sites and where the species are characterized by traits and sites by environmental variables. The trait data and environment data form the external constraints and the question is which traits and environmental variables are associated, how these associations drive species abundances and how they can be displayed in biplots. With microbiome data becoming widely available, these and related multivariate methods deserve more study as they might be routinely used in the future.