Scientists often need to know whether pairs of entities tend to occur together or independently. Standard approaches to this issue use co-occurrence indices such as Jaccard, Sørensen-Dice, and Simpson. We show that these indices are sensitive to the prevalences of the entities they describe and that this invalidates their interpretability. We propose an index, α, that is insensitive to prevalences. Published datasets reanalyzed with both α and Jaccard’s index ( J ) yield profoundly different biological inferences. For example, a published analysis using J contradicted predictions of the island biogeography theory finding that community stability increased with increasing physical isolation. Reanalysis of the same dataset with the estimator α ˆ reversed that result and supported theoretical predictions. We found similarly marked effects in reanalyses of antibiotic cross-resistance and human disease biomarkers. Our index α is not merely an improvement; its use changes data interpretation in fundamental ways.

Drawing on a long history in macroecology, correlation analysis of microbiome datasets is becoming a common practice for identifying relationships or shared ecological niches among bacterial taxa. However, many of the statistical issues that plague such analyses in macroscale communities remain unresolved for microbial communities. Here, we discuss problems in the analysis of microbial species correlations based on presence-absence data. We focus on presence-absence data because this information is more readily obtainable from sequencing studies, especially for whole-genome sequencing, where abundance estimation is still in its infancy. First, we show how Pearson’s correlation coefficient (r) and Jaccard’s index (J)–two of the most common metrics for correlation analysis of presence-absence data–can contradict each other when applied to a typical microbiome dataset. In our dataset, for example, 14% of species-pairs predicted to be significantly correlated by r were not predicted to be significantly correlated using J, while 37.4% of species-pairs predicted to be significantly correlated by J were not predicted to be significantly correlated using r. Mismatch was particularly common among species-pairs with at least one rare species (<10% prevalence), explaining why r and J might differ more strongly in microbiome datasets, where there are large numbers of rare taxa. Indeed 74% of all species-pairs in our study had at least one rare species. Next, we show how Pearson’s correlation coefficient can result in artificial inflation of positive taxon relationships and how this is a particular problem for microbiome studies. We then illustrate how Jaccard’s index of similarity (J) can yield improvements over Pearson’s correlation coefficient. However, the standard null model for Jaccard’s index is flawed, and thus introduces its own set of spurious conclusions. We thus identify a better null model based on a hypergeometric distribution, which appropriately corrects for species prevalence. This model is available from recent statistics literature, and can be used for evaluating the significance of any value of an empirically observed Jaccard’s index. The resulting simple, yet effective method for handling correlation analysis of microbial presence-absence datasets provides a robust means of testing and finding relationships and/or shared environmental responses among microbial taxa.

Alpine treelines are expected to shift upward due to recent climate change. However, interpretation of changes in montane systems has been problematic because effects of climate change are frequently confounded with those of land use changes. The eastern Himalaya, particularly Langtang National Park, Central Nepal, has been relatively undisturbed for centuries and thus presents an opportunity for studying climate change impacts on alpine treeline uncontaminated by potential confounding factors. We studied two dominant species, Abies spectabilis (AS) and Rhododendron campanulatum (RC), above and below the treeline on two mountains. We constructed 13 transects, each spanning up to 400 m in elevation, in which we recorded height and state (dead or alive) of all trees, as well as slope, aspect, canopy density, and measures of anthropogenic and animal disturbance. All size classes of RC plants had lower mortality above treeline than below it, and young RC plants (<2 m tall) were at higher density above treeline than below. AS shows little evidence of a position change from the historic treeline, with a sudden extreme drop in density above treeline compared to below. Recruitment, as measured by size–class distribution, was greater above treeline than below for both species but AS is confined to ~25 m above treeline whereas RC is luxuriantly growing up to 200 m above treeline. Synthesis. Evidence suggests that the elevational limits of RC have shifted upward both because (a) young plants above treeline benefited from facilitation of recruitment by surrounding vegetation, allowing upward expansion of recruitment, and (b) temperature amelioration to mature plants increased adult survival. We predict that the current pure stand of RC growing above treeline will be colonized by AS that will, in turn, outshade and eventually relegate RC to be a minor component of the community, as is the current situation below the treeline.

Human microbiome research is rife with studies attempting to deduce microbial correlation networks from sequencing data. Standard correlation and/or network analyses may be misleading when taken as an indication of taxon interactions because “correlation is neither necessary nor sufficient to establish causation”; environmental filtering can lead to correlation between non-interacting taxa. Unfortunately, microbial ecologists have generally used correlation as a proxy for causality although there is a general consensus about what constitutes a causal relationship: causes both precede and predict effects. We apply one of the first causal models for detecting interactions in human microbiome samples. Specifically, we analyze a long duration, high resolution time series of the human microbiome to decipher the networks of correlation and causation of human-associated microbial genera. We show that correlation is not a good proxy for biological interaction; we observed a weak negative relationship between correlation and causality. Strong interspecific interactions are disproportionately positive, whereas almost all strong intraspecific interactions are negative. Interestingly, intraspecific interactions also appear to act at a short time-scale causing vast majority of the effects within 1–3 days. We report how different taxa are involved in causal relationships with others, and show that strong interspecific interactions are rarely conserved across two body sites whereas strong intraspecific interactions are much more conserved, ranging from 33% between the gut and right-hand to 70% between the two hands. Therefore, in the absence of guiding assumptions about ecological interactions, Granger causality and related techniques may be particularly helpful for understanding the driving factors governing microbiome composition and structure. The human microbiome comprises thousands of microbial taxa, each of them potentially having interactions with many other taxa. Elucidating these interactions can be helpful for human health and beyond. The most conclusive approach for understanding microbial interactions would be myriads of controlled experiments in laboratory settings, each designed to test how taxa interact with one another. As this is a formidable challenge, an alternative strategy for constructing such an interaction network is a statistical framework employing formal models. Using conditional Granger causality, we demonstrate that correlation measures, which are very popular statistics in microbiome studies, are unreliable as proxies of microbial interactions. Overall, we found that strong interactions within a genus tended to be negative (e.g., competition), and they tended to occur on very short timescales (1–2 days). Many of such interactions were common across body sites. By contrast, strong interactions between genera tended to be positive (e.g., mutualism/facilitation) and were more evenly distributed over a range of timescales, up to the 20-day window that we considered. Very few of these interactions were conserved across body sites. We conclude that models of causality can be particularly useful in elucidating microbial interaction networks when laboratory investigations of interactions are impractical.

Species range maps based on expert opinion are a critical resource for conservation planning. Expert maps are usually accompanied by species descriptions that specify sources of internal range heterogeneity, such as habitat associations, but these are rarely considered when using expert maps for analysis. Here, we develop a quantitative metric (“Expert Score”) to evaluate the agreement between an expert map and a habitat probability surface obtained from a species distribution model (SDM). This method rewards both the avoidance of unsuitable sites and the inclusion of suitable sites in the expert map. We obtained expert maps of 330 butterfly species from each of two widely used North American sources (Glassberg and Scott) and computed species‐wise Expert Scores for each. Overall, the Glassberg maps secured higher scores than Scott (Expert Score = 0.61 vs 0.41) but this was due to the specific rules they used to include or exclude areas from ranges. The predictive performance of expert maps was generally hampered by the inclusion of unsuitable sites. Surprisingly, map topology was the primary predictor of expert performance rather than any factor related to species characteristics, such as mobility. Given the heterogeneity and discontinuity of suitable landscape, expert maps that are drawn with more detail are more likely to agree with SDMs and thus minimize both commission and omission errors. Article impact assessment: Detailed expert maps are more likely to agree with species distribution models and thus minimize commission and omission errors. This article is protected by copyright. All rights reserved

Understanding ecological patterns and response to climate change requires unbiased data on species distribution. This can be challenging, especially in biodiverse but extreme environments like the Himalaya. This study presents the results of the first ever application of Unmanned Aerial Systems (UAS) imagery for species-level mapping of vegetation in the Himalaya following a hierarchical Geographic Object Based Image Analysis (GEOBIA) method. The first level of classification separated green vegetated objects from the rest with overall accuracy of 95%. At the second level, seven cover types were identified (including four woody vegetation species). For this, the suitability of various spectral, shape and textural features were tested for classifying them using an ensemble decision tree algorithm. Spectral features alone yielded ~70% accuracy (kappa 0.66) whereas adding textural and shape features marginally improved the accuracy (73%) but at the cost of a substantial increase in processing time. Contrast in plant morphological traits was the key to distinguishing nearby stands as different species. Hence, broad-leaved versus fine needle leaved vegetation were mapped more accurately than structurally similar classes such as Rhododendron anthopogon versus non-photosynthetic vegetation. Results highlight the potential and limitations of the suggested UAS-GEOBIA approach for detailed mapping of plant communities and suggests future research directions.