Environmentally and behaviourally mediated co‐occurrence of functional traits in bird communities of tropical forest fragments

Two major theories of community assembly – based on the assumption of ‘limiting similarity’ or ‘habitat filtering’, respectively – predict contrasting patterns in the spatial arrangement of functional traits. Previous analyses have made progress in testing these predictions and identifying underlying processes, but have also pointed to theoretical as well as methodological shortcomings. Here we applied a recently developed methodology for spatially explicit analysis of phylogenetic meta‐community structure to study the pattern of co‐occurrence of functional traits in Afrotropical and Neotropical bird species inhabiting forest fragments. Focusing separately on locomotory, dietary, and dispersal traits, we tested whether environmental filtering causes spatial clustering, or competition leads to spatial segregation as predicted by limiting similarity theory. We detected significant segregation of species co‐occurrences in African fragments, but not in the Neotropical ones. Interspecific competition had a higher impact on trait co‐occurrence than filter effects, yet no single functional trait was able to explain the observed degree of spatial segregation among species. Despite high regional variability spanning from spatial segregation to aggregation, we found a consistent tendency for a clustered spatial patterning of functional traits among communities in fragmented landscapes, particularly in non‐territorial species. Overall, we show that behavioural effects, such as territoriality, and environmental effects, such as the area of forest remnants or properties of the landscape matrix in which they are embedded, can strongly affect the pattern of trait co‐occurrence. Our findings suggest that trait‐based analyses of community structure should include behavioural and environmental covariates, and we here provide an appropriate method for linking functional traits, species ecology and environmental conditions to clarify the drivers underlying spatial patterns of species co‐occurrence.     

Liana co‐occurrence patterns in a temperate rainforest

Questions: Are liana–host interactions structured at the community level? Do liana–host interactions differ between species, growth form guilds or habitats? Location: Otari‐Wilton's Bush, on the southern tip of North Island, New Zealand. The forest contains 75 ha of mature and regenerating conifer–broadleaf forest. Methods: Nine liana species were quantified among 217 trees to test for negative co‐occurrence patterns. We also conducted additional analyses within and among compartments embedded in the community‐level matrix. Liana and host abundance distributions were assessed across two contrasting habitats. Results: Community‐level analyses revealed negative co‐occurrence patterns. Positive, neutral and negative co‐occurrence patterns were found among compartments within the community‐level matrix. Host species compartments were consistent with randomized expectations, while positive co‐occurrence patterns were found within the host species matrix. Negative co‐occurrence patterns were found inconsistently among lianas that share the same region of host space, and those that do not. Conclusions: Overall, results indicate the liana community is structured non‐randomly. Liana–host interactions appear to follow an opportunistic growth strategy and interactions are due mostly to habitat partitioning.     

Predictor species: Improving assessments of rare species occurrence by modeling environmental co‐responses

Designing an effective conservation strategy requires understanding where rare species are located. Because rare species can be difficult to find, ecologists often identify other species called conservation surrogates that can help inform the distribution of rare species. Species distribution models typically rely on environmental data when predicting the occurrence of species, neglecting the effect of species' co‐occurrences and biotic interactions. Here, we present a new approach that uses Bayesian networks to improve predictions by modeling environmental co‐responses among species. For species from a European peat bog community, our approach consistently performs better than single‐species models and better than conventional multi‐species approaches that include the presence of nontarget species as additional independent variables in regression models. Our approach performs particularly well with rare species and when calibration data are limited. Furthermore, we identify a group of “predictor species” that are relatively common, insensitive to the presence of other species, and can be used to improve occurrence predictions of rare species. Predictor species are distinct from other categories of conservation surrogates such as umbrella or indicator species, which motivates focused data collection of predictor species to enhance conservation practices.     

Contrasting trait assembly patterns in plant and bird communities along environmental and human‐induced land‐use gradients

Human‐driven environmental changes can induce marked shifts in the functional structure of biological communities with possible repercussion on important ecosystem functions and services. At the same time it remains unclear to which extent these changes may differently affect various types of organisms. We investigated species richness and community functional structure of species assemblages at the landscape scale (1 km² plots) for two contrasting model taxa, i.e. plants (producers and sessile organisms) and birds (consumers and mobile organisms), along topography, climate, landscape heterogeneity, and land‐use (agriculture and urbanization) gradients in a densely populated region of Switzerland. Our study revealed that agricultural and urban land uses drove marked shifts in the functional structure of biological communities compared to changes along climate and topography gradients, especially for plants, while for birds these changes were comparable. Agricultural and urban land uses enhanced divergence in traits related to resource use for birds (diet and nesting), growth forms, dispersal, and reproductive traits for plants, while it induced convergence in vegetative plant traits (plant height and leaf dry matter content). These results suggest that contrasting assembly patterns may arise within and across taxonomic groups along the same environmental gradients as result of distinct underlying processes and ‘organism‐specific’ environmental perceptions. Our results further suggest a potential homogenization of biological communities, as well as low functional diversity and redundancy levels of bird assemblages in our human‐dominated study region. This might potentially compromise the maintenance of key ecological processes under future environmental changes.     

Accounting for environmental variation in co‐occurrence modelling reveals the importance of positive interactions in root‐associated fungal communities

Understanding the role of interspecific interactions in shaping ecological communities is one of the central goals in community ecology. In fungal communities, measuring interspecific interactions directly is challenging because these communities are composed of large numbers of species, many of which are unculturable. An indirect way of assessing the role of interspecific interactions in determining community structure is to identify the species co‐occurrences that are not constrained by environmental conditions. In this study, we investigated co‐occurrences among root‐associated fungi, asking whether fungi co‐occur more or less strongly than expected based on the environmental conditions and the host plant species examined. We generated molecular data on root‐associated fungi of five plant species evenly sampled along an elevational gradient at a high arctic site. We analysed the data using a joint species distribution modelling approach that allowed us to identify those co‐occurrences that could be explained by the environmental conditions and the host plant species, as well as those co‐occurrences that remained unexplained and thus more probably reflect interactive associations. Our results indicate that not only negative but also positive interactions play an important role in shaping microbial communities in arctic plant roots. In particular, we found that mycorrhizal fungi are especially prone to positively co‐occur with other fungal species. Our results bring new understanding to the structure of arctic interaction networks by suggesting that interactions among root‐associated fungi are predominantly positive.     

Species interactions weakly modify climate‐induced tree co‐occurrence patterns

Aims

Species distributions are hypothesized to be underlain by a complex association of processes that span multiple spatial scales including biotic interactions, dispersal limitation, fine‐scale resource gradients and climate. Species disequilibrium with climate may reflect the effects of non‐climatic processes on species distributions, yet distribution models have rarely directly considered non‐climatic processes. Here, we use a Joint Species Distribution Model (JSDM) to investigate the influence of non‐climatic factors on species co‐occurrence patterns and to directly quantify the relative influences of climate and alternative processes that may generate correlated responses in species distributions, such as species interactions, on tree co‐occurrence patterns.

Location

US Rocky Mountains.

Methods

We apply a Bayesian JSDM to simultaneously model the co‐occurrence patterns of ten dominant tree species across the Rocky Mountains, and evaluate climatic and residual correlations from the fitted model to determine the relative contribution of each component to observed co‐occurrence patterns. We also evaluate predictions generated from the fitted model relative to a single‐species modelling approach.

Results

For most species, correlation due to climate covariates exceeded residual correlation, indicating an overriding influence of broad‐scale climate on co‐occurrence patterns. Accounting for covariance among species did not significantly improve predictions relative to a single‐species approach, providing limited evidence for a strong independent influence of species interactions on distribution patterns.

Conclusions

Overall, our findings indicate that climate is an important driver of regional biodiversity patterns and that interactions between dominant tree species contribute little to explain species co‐occurrence patterns among Rocky Mountain trees.     

Species co‐occurrence patterns and dietary resource competition in primates

Diamond (Assembly of species communities. In: Cody ML, Diamond JM, editors. Ecology and evolution of communities. Cambridge: Belknap. p 342–444 ( 1975 )) argued that interspecific competition between species occupying similar niches results in a nonrandom pattern of species distributions. In particular, some species pairs may never be found in the same community due to competitive exclusion. Rigorous analytical methods have been developed to investigate the possible role that interspecific competition has on the evolution of communities. Many studies that have implemented these methods have shown support for Diamond's assembly rules, yet there are numerous exceptions. We build on this previous research by examining the co‐occurrence patterns of primate species in 109 communities from across the world. We used EcoSim to calculate a checkerboard (C) score for each region. The C score provides a measure of the proportion of species pairs that do not co‐occur in a set of communities. High C scores indicate that species are nonrandomly distributed throughout a region, and interspecific competition may be driving patterns of competitive exclusion. We conducted two sets of analyses. One included all primate species per region, and the second analysis assigned each species to one of four dietary guilds: frugivores, folivores, insectivores, and frugivore‐insectivores. Using all species per region, we found significantly high C scores in 9 of 10 regions examined. For frugivores, we found significantly high‐C scores in more than 50% of regions. In contrast, only 23% of regions exhibited significantly high‐C scores for folivores. Our results suggest that communities are nonrandomly structured and may be the result of greater levels of interspecific competition between frugivores compared to folivores. Am J Phys Anthropol, 2010. © 2010 Wiley‐Liss, Inc.     

Examining the link between competition and negative co‐occurrence patterns

Negative species co‐occurrence patterns have long intrigued ecologists because of their potential link to competition. Although manipulative field experiments have consistently revealed evidence of competition in natural communities, there is little evidence that this competition produces negative co‐occurrence patterns. Evidence does suggest that abiotic variation, dispersal limitation and herbivory can contribute to patterns of negative co‐occurrence among species; it is possible these influences have obscured a link with competition. Here, we test for a connection between negative co‐occurrence and competition by examining a small‐scale, relatively homogeneous old‐field plant community where the influence of abiotic variation was likely to be minimal and we accounted for the impact of herbivory with an herbivore exclosure treatment. Using three years of data (two biennial periods), we tested whether negatively co‐occurring pairs of species, when occasionally found together, experienced asymmetric abundance decline more frequently than positively co‐occurring pairs, for which there is no such expectation. We found no evidence that negatively co‐occurring pairs consistently suffered asymmetric abundance decline more frequently than positively co‐occurring pairs, providing no evidence that competition is a primary driver of negative co‐occurrence patterns in this community. Our results were consistent across control and herbivore exclosure treatments, suggesting that herbivores are not driving patterns of negative species co‐occurrence in this community. Any influence of competition or herbivory on co‐occurrence patterns is small enough that it is obscured by other factors such as substrate heterogeneity, dispersal and differential species responses to climatic variation through time. We interpret our results as providing evidence that competition is not responsible for producing negative co‐occurrence patterns in our study community and suggest that this may be the case more broadly.     

Using co‐occurrence network topology in assessing ecological stress in benthic macroinvertebrate communities

Ecological monitoring of streams has often focused on assessing the biotic integrity of individual benthic macroinvertebrate (BMI) communities through local measures of diversity, such as taxonomic or functional richness. However, as individual BMI communities are frequently linked by a variety of ecological processes at a regional scale, there is a need to assess biotic integrity of groups of communities at the scale of watersheds. Using 4,619 sampled communities of streambed BMIs, we investigate this question using co‐occurrence networks generated from groups of communities selected within California watersheds under different levels of stress due to upstream land use. Building on a number of arguments in theoretical ecology and network theory, we propose a framework for the assessment of the biotic integrity of watershed‐scale groupings of BMI communities using measures of their co‐occurrence network topology. We found significant correlations between stress, as described by a mean measure of upstream land use within a watershed, and topological measures of co‐occurrence networks such as network size (r = ?.81, p < 10^–4), connectance (r = .31, p < 10^–4), mean co‐occurrence strength (r = .25, p < 10^–4), degree heterogeneity (r = ?.10, p < 10^–4), and modularity (r = .11, p < 10^–4). Using these five topological measures, we constructed a linear model of biotic integrity, here a composite of taxonomic and functional diversity known as the California Stream Condition Index, of groups of BMI communities within a watershed. This model can account for 66% of among‐watershed variation in the mean biotic integrity of communities. These observations imply a role for co‐occurrence networks in assessing the current status of biotic integrity for BMI communities, as well as their potential use in assessing other ecological communities.     

Fire disturbance disrupts co‐occurrence patterns of terrestrial vertebrates in Mediterranean woodlands

Aim This paper uses null model analysis to explore the pattern of species co‐occurrence of terrestrial vertebrate fauna in fire‐prone, mixed evergreen oak woodlands. Location The Erico–Quercion ilicis of the Mediterranean belt (50–800 m a.s.l.) in the Madonie mountain range, a regional park in northern Sicily (37°50′ N, 14°05′ E), Italy. Methods The stratified sampling of vertebrates in a secondary succession of recent burned areas (BA, 1–2 years old), intermediate burned areas (INT, 4–10 years old) and ancient burned areas (CNB, > 50 years old), plus forest fragments left within burned areas (FF, 1–2 years old) permitted the comparison of patterns of species co‐occurrence using a set of separate presence/absence matrices. First, the breeding avifauna derived from standardized point counts was analysed using Stone & Roberts’C‐score, and by a null model algorithm (fixed/equiprobable). Secondly, the analysis was repeated using all vertebrate species recorded in the succession. Results Sixty‐five species were recorded in the 2‐year study period in the four sample treatments. Birds were found to make up the largest component (63%) of the recorded assemblage. The BA treatment had the lowest species richness, followed in order by the small, medium and large FFs, and then by the CNBs. For both analyses (birds and total vertebrates), the C‐scores were quite small and not significantly different from those that could be expected by chance in the BA and INT burned areas; this indicates a random co‐occurrence among vertebrates of those assemblages. Contrariwise, for both analyses in the CNBs, the C‐scores were large and significantly different from the simulated indices, thereby indicating a non‐random co‐occurrence pattern (segregation) of vertebrates in the undisturbed woodlands. In addition, C‐score values for the surviving FFs show a significant aggregation of species. Main conclusions The null model analyses highlighted a new aspect of fire disturbance in Mediterranean woodland ecosystems: the disruption in patterns of co‐occurrence in the terrestrial vertebrate community. Wildfire alters community organization, inducing, for at least 10 years, a random aggregate of species. Communities re‐assemble themselves, showing the occurrence of species segregation at least 50 years after fire.     

Multisite and multispecies measures of overlap,co‐occurrence,and co‐diversity

Several indices measure the association or segregation between two species and the similarity or differentiation between two sets of species. These indices are based on the overlap in the distribution of species (measured with the number of co‐occurrences) or on the overlap in species composition of sites (measured with the number of species that are shared between two sites). This paper shows that when evaluating more than two species the number of overlaps and the number of pairwise co‐occurrences are not equal, as it is the case for two species. Equivalently, when comparing more than two species assemblages, the number of overlaps differ from the number of instances of species sharing by pairs of sites (the ‘co‐diversities’). Considering this distinction, two different types of multispecies and multisite indices can be derived: indices of general overlap and indices of co‐occurrence or co‐diversity. Here I present a complete series of the two types of indices that correspond to the popular Jaccard, Sørensen, and Simpson two‐species or two‐site indices. Indices of general overlap are defined by three parameters (the total number of species, the total number of sites, and the total number of occurrences), whereas indices of co‐occurrence or co‐diversity depend on those parameters plus an additional one that is defined by the values of species richness or range size. Consequently, the two types of indices respond differently to null models, depending on the parameters that are fixed or randomized. All indices correlate well with the mean of the traditional indices calculated pair by pair, and the correspondence is extremely close for the new indices of co‐occurrence and co‐diversity. These properties should be useful in clarifying some of the confusion that exists in the current discussion about the advantages and disadvantages of pairwise vs community‐wide approaches in the analysis of diversity.     

Environmental factors affecting patterns of distribution and co‐occurrence of two competing raptor species

Habitat selection is a complex process, that is affected by several factors, including habitat characteristics, environmental conditions, and both intra‐ and interspecific interactions. We analysed habitat preferences of two top avian predators, Peregrine Falcon Falco peregrinus, a medium‐sized diurnal raptor, and Eagle Owl Bubo bubo, a large nocturnal raptor. These two species are known to compete for preferred nest‐sites, and proximity to cliffs with Eagle Owls may reduce Peregrine breeding output through predation of young Falcons. We investigated the environmental factors affecting occurrence and coexistence of the two species and the potential role of habitat suitability in favouring co‐occurrence in 3519 km² of the central pre‐Alps of Italy, where the two species breed on cliffs and sometimes co‐occur on the same cliff. Peregrines settled on long, steep and favourably orientated cliffs in woodland landscapes close to urban areas. Eagle Owls settled on topographically similar cliffs, but in lower rainfall areas compared with cliffs occupied by Peregrines and cliffs unoccupied by either species. Sites where the two species co‐occurred were characterized by more horizontally extended cliffs compared with sites of exclusive occurrence of each species. An analysis of relative habitat suitability revealed that sites where the two species co‐occurred had the highest predicted probability of occupancy for both species, suggesting that those sites should be regarded as high‐quality sites. Breeding productivity of Eagle Owls was negatively affected by the co‐occurrence of Peregrines, whereas the effect of Eagle Owl proximity on Peregrine productivity varied according to cliff suitability for the Peregrines. Habitat selection had fitness consequences for Eagle Owls because breeding productivity increased with cliff length. Environmental conditions, particularly climatic factors, could allow the widespread coexistence of these competing raptors at the landscape scale, whereas at the local scale co‐occurrence could take place only on larger cliffs. These were preferred sites for both species, presumably because breeding at such sites offsets the costs of settling close to the competitor species.     

Specific plasmid patterns and high rates of bacterial co‐occurrence within the coral holobiont

Despite the importance of coral microbiomes for holobiont persistence, the interactions among these are not well understood. In particular, knowledge of the co‐occurrence and taxonomic importance of specific members of the microbial core, as well as patterns of specific mobile genetic elements (MGEs), is lacking. We used seawater and mucus samples collected from Mussismilia hispida colonies on two reefs located in Bahia, Brazil, to disentangle their associated bacterial communities, intertaxa correlations, and plasmid patterns. Proxies for two broad‐host‐range (BHR) plasmid groups, IncP‐1β and PromA, were screened. Both groups were significantly (up to 252 and 100%, respectively) more abundant in coral mucus than in seawater. Notably, the PromA plasmid group was detected only in coral mucus samples. The core bacteriome of M. hispida mucus was composed primarily of members of the Proteobacteria, followed by those of Firmicutes. Significant host specificity and co‐occurrences among different groups of the dominant phyla (e.g., Bacillaceae and Pseudoalteromonadaceae and the genera Pseudomonas, Bacillus, and Vibrio) were detected. These relationships were observed for both the most abundant phyla and the bacteriome core, in which most of the operational taxonomic units showed intertaxa correlations. The observed evidence of host‐specific bacteriome and co‐occurrence (and potential symbioses or niche space co‐dominance) among the most dominant members indicates a taxonomic selection of members of the stable bacterial community. In parallel, host‐specific plasmid patterns could also be, independently, related to the assembly of members of the coral microbiome.     

Seasonal dynamics and co‐occurrence patterns of honey bee pathogens revealed by high‐throughput RT‐qPCR analysis

The health of the honey bee Apis mellifera is challenged by introduced parasites that interact with its inherent pathogens and cause elevated rates of colony losses. To elucidate co‐occurrence, population dynamics, and synergistic interactions of honey bee pathogens, we established an array of diagnostic assays for a high‐throughput qPCR platform. Assuming that interaction of pathogens requires co‐occurrence within the same individual, single worker bees were analyzed instead of collective samples. Eleven viruses, four parasites, and three pathogenic bacteria were quantified in more than one thousand single bees sampled from sixteen disease‐free apiaries in Southwest Germany. The most abundant viruses were black queen cell virus (84%), Lake Sinai virus 1 (42%), and deformed wing virus B (35%). Forager bees from asymptomatic colonies were infected with two different viruses in average, and simultaneous infection with four to six viruses was common (14%). Also, the intestinal parasites Nosema ceranae (96%) and Crithidia mellificae/Lotmaria passim (52%) occurred very frequently. These results indicate that low‐level infections in honey bees are more common than previously assumed. All viruses showed seasonal variation, while N. ceranae did not. The foulbrood bacteria Paenibacillus larvae and Melissococcus plutonius were regionally distributed. Spearman's correlations and multiple regression analysis indicated possible synergistic interactions between the common pathogens, particularly for black queen cell virus. Beyond its suitability for further studies on honeybees, this targeted approach may be, due to its precision, capacity, and flexibility, a viable alternative to more expensive, sequencing‐based approaches in nonmodel systems.     

Predicting resilience of ecosystem functioning from co‐varying species' responses to environmental change

Understanding how environmental change affects ecosystem function delivery is of primary importance for fundamental and applied ecology. Current approaches focus on single environmental driver effects on communities, mediated by individual response traits. Data limitations present constraints in scaling up this approach to predict the impacts of multivariate environmental change on ecosystem functioning. We present a more holistic approach to determine ecosystem function resilience, using long‐term monitoring data to analyze the aggregate impact of multiple historic environmental drivers on species' population dynamics. By assessing covariation in population dynamics between pairs of species, we identify which species respond most synchronously to environmental change and allocate species into “response guilds.” We then use “production functions” combining trait data to estimate the relative roles of species to ecosystem functions. We quantify the correlation between response guilds and production functions, assessing the resilience of ecosystem functioning to environmental change, with asynchronous dynamics of species in the same functional guild expected to lead to more stable ecosystem functioning. Testing this method using data for butterflies collected over four decades in the United Kingdom, we find three ecosystem functions (resource provisioning, wildflower pollination, and aesthetic cultural value) appear relatively robust, with functionally important species dispersed across response guilds, suggesting more stable ecosystem functioning. Additionally, by relating genetic distances to response guilds we assess the heritability of responses to environmental change. Our results suggest it may be feasible to infer population responses of butterflies to environmental change based on phylogeny—a useful insight for conservation management of rare species with limited population monitoring data. Our approach holds promise for overcoming the impasse in predicting the responses of ecosystem functions to environmental change. Quantifying co‐varying species' responses to multivariate environmental change should enable us to significantly advance our predictions of ecosystem function resilience and enable proactive ecosystem management.     

Statistical challenges of evaluating diversity patterns across environmental gradients in mega‐diverse communities

Ibanez et al. (Journal of Vegetation Science, this issue) applied sample size‐ and coverage‐based rarefaction to analyse the elevational richness pattern in New Caledonian tree communities. We comment on the statistical assumptions behind rarefaction/extrapolation and suggest pooling small plot data to effectively assess/detect the diversity pattern. Broadening the analysis to include abundance‐sensitive diversity measures and phylogenetic information can provide important additional insights.