Temperature sensitivity of pest reproductive numbers in age-structured PDE models, with a focus on the invasive spotted lanternfly.

Invasive pest establishment is a pervasive threat to global ecosystems, agriculture, and public health. The recent establishment of the invasive spotted lanternfly in the northeastern United States has proven devastating to farms and vineyards, necessitating urgent development of population dynamical models and effective control practices. In this paper, we propose a stage-age-structured system of PDEs to model insect pest populations, in which underlying dynamics are dictated by ambient temperature through rates of development, fecundity, and mortality. The model incorporates diapause and non-diapause pathways, and is calibrated to experimental and field data on the spotted lanternfly. We develop a novel moving mesh method for capturing age-advection accurately, even for coarse discretization parameters. We define a one-year reproductive number ([Formula: see text]) from the spectrum of a one-year solution operator, and study its sensitivity to variations in the mean and amplitude of the annual temperature profile. We quantify assumptions sufficient to give rise to the low-rank structure of the solution operator characteristic of part of the parameter domain. We discuss establishment potential as it results from the pairing of a favorable [Formula: see text] value and transient population survival, and address implications for pest control strategies.

Disturbed habitats locally reduce the signal of deep evolutionary history in functional traits of plants.

The functioning of present ecosystems reflects deep evolutionary history of locally cooccurring species if their functional traits show high phylogenetic signal (PS). However, we do not understand what drives local PS. We hypothesize that local PS is high in undisturbed and stressful habitats, either due to ongoing local assembly of species that maintained ancestral traits, or to past evolutionary maintenance of ancestral traits within habitat species-pools, or to both. We quantified PS and diversity of 10 traits within 6704 local plant communities across 38 Dutch habitat types differing in disturbance or stress. Mean local PS varied 50-fold among habitat types, often independently of phylogenetic or trait diversity. Mean local PS decreased with disturbance but showed no consistent relationship to stress. Mean local PS exceeded species-pool PS, reflecting nonrandom subsampling from the pool. Disturbance or stress related more strongly to mean local than to species-pool PS. Disturbed habitats harbour species with evolutionary divergent trait values, probably driven by ongoing, local assembly of species: environmental fluctuations might maintain different trait values within lineages through an evolutionary storage effect. If functional traits do not reflect phylogeny, ecosystem functioning might not be contingent on the presence of particular lineages, and lineages might establish evolutionarily novel interactions.

Disentangling the effects of plant species invasion and urban development on arthropod community composition.

Urban development and species invasion are two major global threats to biodiversity. These threats often co-occur, as developed areas are more prone to species invasion. However, few empirical studies have tested if both factors affect biodiversity in similar ways. Here we study the individual and combined effects of urban development and plant invasion on the composition of arthropod communities. We assessed 36 paired invaded and non-invaded sample plots, invaded by the plant Antigonon leptopus, with half of these pairs located in natural and the other half in developed land-use types on the Caribbean island of St. Eustatius. We used several taxonomic and functional variables to describe community composition and diversity. Our results show that both urban development and A. leptopus invasion affected community composition, albeit in different ways. Development significantly increased species richness and exponential Shannon diversity, while invasion had no effect on these variables. However, invasion significantly increased arthropod abundance and caused biotic homogenization. Specifically, uninvaded arthropod communities were distinctly different in species composition between developed and natural sites, while they became undistinguishable after A. leptopus invasion. Moreover, functional variables were significantly affected by species invasion, but not by urban development. Invaded communities had higher community-weighted mean body size and the feeding guild composition of invaded arthropod communities was characterized by the exceptional numbers of nectarivores, herbivores, and detritivores. With the exception of species richness and exponential Shannon diversity, invasion influenced four out of six response variables to a greater degree than urban development did. Hence, we can conclude that species invasion is not just a passenger of urban development but also a driver of change.

Island biogeography of the Anthropocene.

For centuries, biogeographers have examined the factors that produce patterns of biodiversity across regions. The study of islands has proved particularly fruitful and has led to the theory that geographic area and isolation influence species colonization, extinction and speciation such that larger islands have more species and isolated islands have fewer species (that is, positive species-area and negative species-isolation relationships). However, experimental tests of this theory have been limited, owing to the difficulty in experimental manipulation of islands at the scales at which speciation and long-distance colonization are relevant. Here we have used the human-aided transport of exotic anole lizards among Caribbean islands as such a test at an appropriate scale. In accord with theory, as anole colonizations have increased, islands impoverished in native species have gained the most exotic species, the past influence of speciation on island biogeography has been obscured, and the species-area relationship has strengthened while the species-isolation relationship has weakened. Moreover, anole biogeography increasingly reflects anthropogenic rather than geographic processes. Unlike the island biogeography of the past that was determined by geographic area and isolation, in the Anthropocene--an epoch proposed for the present time interval--island biogeography is dominated by the economic isolation of human populations.

Evolutionary time drives global tetrapod diversity.

Global variation in species richness is widely recognized, but the explanation for what drives it continues to be debated. Previous efforts have focused on a subset of potential drivers, including evolutionary rate, evolutionary time (maximum clade age of species restricted to a region), dispersal (migration from one region to another), ecological factors and climatic stability. However, no study has evaluated these competing hypotheses simultaneously at a broad spatial scale. Here, we examine their relative contribution in determining the richness of the most comprehensive dataset of tetrapods to our knowledge (84% of the described species), distinguishing between the direct influences of evolutionary rate, evolutionary time and dispersal, and the indirect influences of ecological factors and climatic stability through their effect on direct factors. We found that evolutionary time exerted a primary influence on species richness, with evolutionary rate being of secondary importance. By contrast, dispersal did not significantly affect richness patterns. Ecological and climatic stability factors influenced species richness indirectly by modifying evolutionary time (i.e. persistence time) and rate. Overall, our findings suggest that global heterogeneity in tetrapod richness is explained primarily by the length of time species have had to diversify.

Human land use promotes the abundance and diversity of exotic species on Caribbean islands.

Human land use causes major changes in species abundance and composition, yet native and exotic species can exhibit different responses to land use change. Native populations generally decline in human-impacted habitats while exotic species often benefit. In this study, we assessed the effects of human land use on exotic and native reptile diversity, including functional diversity, which relates to the range of habitat use strategies in biotic communities. We surveyed 114 reptile communities from localities that varied in habitat structure and human impact level on two Caribbean islands, and calculated species richness, overall abundance, and evenness for every plot. Functional diversity indices were calculated using published trait data, which enabled us to detect signs of trait filtering associated with impacted habitats. Our results show that environmental variation among sampling plots was explained by two Principal Component Analysis (PCA) ordination axes related to habitat structure (i.e., forest or nonforest) and human impact level (i.e., addition of man-made constructions such as roads and buildings). Several diversity indices were significantly correlated with the two PCA axes, but exotic and native species showed opposing responses. Native species reached the highest abundance in forests, while exotic species were absent in this habitat. Human impact was associated with an increase in exotic abundance and species richness, while native species showed no significant associations. Functional diversity was highest in nonforested environments on both islands, and further increased on St. Martin with the establishment of functionally unique exotic species in nonforested habitat. Habitat structure, rather than human impact, proved to be an important agent for environmental filtering of traits, causing divergent functional trait values across forested and nonforested environments. Our results illustrate the importance of considering various elements of land use when studying its impact on species diversity and the establishment and spread of exotic species.

Benefits and limitations of three-dimensional printing technology for ecological research.

BACKGROUND: Ecological research often involves sampling and manipulating non-model organisms that reside in heterogeneous environments. As such, ecologists often adapt techniques and ideas from industry and other scientific fields to design and build equipment, tools, and experimental contraptions custom-made for the ecological systems under study. Three-dimensional (3D) printing provides a way to rapidly produce identical and novel objects that could be used in ecological studies, yet ecologists have been slow to adopt this new technology. Here, we provide ecologists with an introduction to 3D printing. RESULTS: First, we give an overview of the ecological research areas in which 3D printing is predicted to be the most impactful and review current studies that have already used 3D printed objects. We then outline a methodological workflow for integrating 3D printing into an ecological research program and give a detailed example of a successful implementation of our 3D printing workflow for 3D printed models of the brown anole, Anolis sagrei, for a field predation study. After testing two print media in the field, we show that the models printed from the less expensive and more sustainable material (blend of 70% plastic and 30% recycled wood fiber) were just as durable and had equal predator attack rates as the more expensive material (100% virgin plastic). CONCLUSIONS: Overall, 3D printing can provide time and cost savings to ecologists, and with recent advances in less toxic, biodegradable, and recyclable print materials, ecologists can choose to minimize social and environmental impacts associated with 3D printing. The main hurdles for implementing 3D printing-availability of resources like printers, scanners, and software, as well as reaching proficiency in using 3D image software-may be easier to overcome at institutions with digital imaging centers run by knowledgeable staff. As with any new technology, the benefits of 3D printing are specific to a particular project, and ecologists must consider the investments of developing usable 3D materials for research versus other methods of generating those materials.

The Evolutionary Legacy of Diversification Predicts Ecosystem Function.

Theory suggests that the structure of evolutionary history represented in a species community may affect its functioning, but phylogenetic diversity metrics do not allow for the identification of major differences in this structure. Here we propose a new metric, ELDERness (for Evolutionary Legacy of DivERsity) to estimate evolutionary branching patterns within communities by fitting a polynomial function to lineage-through-time (LTT) plots. We illustrate how real and simulated community branching patterns can be more correctly described by ELDERness and can successfully predict ecosystem functioning. In particular, the evolutionary history of branching patterns can be encapsulated by the parameters of third-order polynomial functions and further measured through only two parameters, the "ELDERness surfaces." These parameters captured variation in productivity of a grassland community better than existing phylogenetic diversity or diversification metrics and independent of species richness or presence of nitrogen fixers. Specifically, communities with small ELDERness surfaces (constant accumulation of lineages through time in LTT plots) were more productive, consistent with increased productivity resulting from complementary lineages combined with niche filling within lineages. Overall, while existing phylogenetic diversity metrics remain useful in many contexts, we suggest that our ELDERness approach better enables testing hypotheses that relate complex patterns of macroevolutionary history represented in local communities to ecosystem functioning.

pez: phylogenetics for the environmental sciences.

UNLABELLED: pez is an R package that permits measurement, modelling and simulation of phylogenetic structure in ecological data. pez contains the first implementation of many methods in R, and aggregates existing data structures and methods into a single, coherent package. AVAILABILITY AND IMPLEMENTATION: pez is released under the GPL v3 open-source license, available on the Internet from CRAN (http://cran.r-project.org). The package is under active development, and the authors welcome contributions (see http://github.com/willpearse/pez). CONTACT: will.pearse@gmail.com.

Synchronous dynamics of zooplankton competitors prevail in temperate lake ecosystems.

Although competing species are expected to exhibit compensatory dynamics (negative temporal covariation), empirical work has demonstrated that competitive communities often exhibit synchronous dynamics (positive temporal covariation). This has led to the suggestion that environmental forcing dominates species dynamics; however, synchronous and compensatory dynamics may appear at different length scales and/or at different times, making it challenging to identify their relative importance. We compiled 58 long-term datasets of zooplankton abundance in north-temperate and sub-tropical lakes and used wavelet analysis to quantify general patterns in the times and scales at which synchronous/compensatory dynamics dominated zooplankton communities in different regions and across the entire dataset. Synchronous dynamics were far more prevalent at all scales and times and were ubiquitous at the annual scale. Although we found compensatory dynamics in approximately 14% of all combinations of time period/scale/lake, there were no consistent scales or time periods during which compensatory dynamics were apparent across different regions. Our results suggest that the processes driving compensatory dynamics may be local in their extent, while those generating synchronous dynamics operate at much larger scales. This highlights an important gap in our understanding of the interaction between environmental and biotic forces that structure communities.

Subsidies to predators, apparent competition and the phylogenetic structure of prey communities.

Ecosystems are fragmented by natural and anthropogenic processes that affect organism movement and ecosystem dynamics. When a fragmentation restricts predator but not prey movement, then the prey produced on one side of an ecosystem edge can subsidize predators on the other side. When prey flux is high, predator density on the receiving side increases above that possible by in situ prey productivity, and when low, the formerly subsidized predators can impose strong top-down control of in situ prey--in situ prey experience apparent competition from the subsidy. If predators feed on some evolutionary clades of in situ prey over others, then subsidy-derived apparent competition will induce phylogenetic structure in prey composition. Dams fragment the serial nature of river ecosystems by prohibiting movement of organisms and restricting flowing water. In the river tailwater just below a large central Mexican dam, fish density was high and fish gorged on reservoir-derived zooplankton. When the dam was closed, water flow and the zooplankton subsidy ceased, densely packed pools of fish formed, fish switched to feed on in situ prey, and the tailwater macroinvertebrate community was phylogenetic structured. We derived expectations of structure from trait-based community assembly models based on macroinvertebrate body size, tolerance to anthropogenic disturbance, and fish-diet selectivity. The diet-selectivity model best fit the observed tailwater phylogenetic structure. Thus, apparent competition from subsidies phylogenetically structures prey communities, and serial variation in phylogenetic community structure can be indicative of fragmentation in formerly continuous ecosystems.

Relevance of evolutionary history for food web structure.

Explaining the structure of ecosystems is one of the great challenges of ecology. Simple models for food web structure aim at disentangling the complexity of ecological interaction networks and detect the main forces that are responsible for their shape. Trophic interactions are influenced by species traits, which in turn are largely determined by evolutionary history. Closely related species are more likely to share similar traits, such as body size, feeding mode and habitat preference than distant ones. Here, we present a theoretical framework for analysing whether evolutionary history--represented by taxonomic classification--provides valuable information on food web structure. In doing so, we measure which taxonomic ranks better explain species interactions. Our analysis is based on partitioning of the species into taxonomic units. For each partition, we compute the likelihood that a probabilistic model for food web structure reproduces the data using this information. We find that taxonomic partitions produce significantly higher likelihoods than expected at random. Marginal likelihoods (Bayes factors) are used to perform model selection among taxonomic ranks. We show that food webs are best explained by the coarser taxonomic ranks (kingdom to class). Our methods provide a way to explicitly include evolutionary history in models for food web structure.

Paninvasion severity assessment of a U.S. grape pest to disrupt the global wine market.

Economic impacts from plant pests are often felt at the regional scale, yet some impacts expand to the global scale through the alignment of a pest's invasion potentials. Such globally invasive species (i.e., paninvasives) are like the human pathogens that cause pandemics. Like pandemics, assessing paninvasion risk for an emerging regional pest is key for stakeholders to take early actions that avoid market disruption. Here, we develop the paninvasion severity assessment framework and use it to assess a rapidly spreading regional U.S. grape pest, the spotted lanternfly planthopper (Lycorma delicatula; SLF), to spread and disrupt the global wine market. We found that SLF invasion potentials are aligned globally because important viticultural regions with suitable environments for SLF establishment also heavily trade with invaded U.S. states. If the U.S. acts as an invasive bridgehead, Italy, France, Spain, and other important wine exporters are likely to experience the next SLF introductions. Risk to the global wine market is high unless stakeholders work to reduce SLF invasion potentials in the U.S. and globally.

Picante: R tools for integrating phylogenies and ecology.

SUMMARY: Picante is a software package that provides a comprehensive set of tools for analyzing the phylogenetic and trait diversity of ecological communities. The package calculates phylogenetic diversity metrics, performs trait comparative analyses, manipulates phenotypic and phylogenetic data, and performs tests for phylogenetic signal in trait distributions, community structure and species interactions. AVAILABILITY: Picante is a package for the R statistical language and environment written in R and C, released under a GPL v2 open-source license, and freely available on the web (http://picante.r-forge.r-project.org) and from CRAN (http://cran.r-project.org).

Communities contain closely related species during ecosystem disturbance.

Predicting community and species responses to disturbance is complicated by incomplete knowledge about species traits. A phylogenetic framework should partially solve this problem, as trait similarity is generally correlated with species relatedness, closely related species should have similar sensitivities to disturbance. Disturbance should thus result in community assemblages of closely related species. We tested this hypothesis with 18 disturbed and 16 reference whole-lake, long-term zooplankton data sets. Regardless of disturbance type, communities generally contained more closely related species when disturbed. This effect was independent of species richness, evenness, and abundance. Communities already under stress (i.e., those in acidic lakes) changed most when disturbed. Species sensitivities to specific disturbances were phylogenetically conserved, were independent of body size, and could be predicted by the sensitivities of close relatives within the same community. Phylogenetic relatedness can effectively act as a proxy for missing trait information when predicting community and species responses to disturbance.

Phylogenetic metrics of community similarity.

We derive a new metric of community similarity that takes into account the phylogenetic relatedness among species. This metric, phylogenetic community dissimilarity (PCD), can be partitioned into two components, a nonphylogenetic component that reflects shared species between communities (analogous to Sørensen' s similarity metric) and a phylogenetic component that reflects the evolutionary relationships among nonshared species. Therefore, even if a species is not shared between two communities, it will increase the similarity of the two communities if it is phylogenetically related to species in the other community. We illustrate PCD with data on fish and aquatic macrophyte communities from 59 temperate lakes. Dissimilarity between fish communities associated with environmental differences between lakes often has a phylogenetic component, whereas this is not the case for macrophyte communities. With simulations, we then compare PCD with two other metrics of phylogenetic community similarity, II(ST) and UniFrac. Of the three metrics, PCD was best at identifying environmental drivers of community dissimilarity, showing lower variability and greater statistical power. Thus, PCD is a statistically powerful metric that separates the effects of environmental drivers on compositional versus phylogenetic components of community structure.

Species diversity as a surrogate for conservation of phylogenetic and functional diversity in terrestrial vertebrates across the Americas.

Preserving the evolutionary history and ecological functions that different species embody, in addition to species themselves, is a growing concern for conservation. Recent studies warn that conservation priority regions identified using species diversity differ from those based on phylogenetic or functional diversity. However, spatial mismatches in conservation priority regions need not indicate low surrogacy among these dimensions in conservation planning. Here, we use data for 10,213 terrestrial vertebrate species across the Americas to evaluate surrogacy; that is, the proportion of phylogenetic or functional diversity represented in conservation plans targeting species. We find that most conservation plans targeting species diversity also represent phylogenetic and functional diversity well, despite spatial mismatches in the priority regions identified by each plan. However, not all phylogenetic and functional diversity is represented within species-based plans, with the highest-surrogacy conservation strategy depending on the proportion of land area included in plans. Our results indicate that targeting species diversity could be sufficient to preserve much of the phylogenetic and functional dimensions of biodiversity in terrestrial vertebrates of the Americas. Incorporating phylogenetic and functional data in broad-scale conservation planning may not always be necessary, especially when the cost of doing so is high.

Separating the determinants of phylogenetic community structure.

The role of competition in forbidding similar species from co-occurring has long been debated. A difficulty in identifying this repulsion of similar species is that similar species share similar environmental requirements and hence show an attraction to communities where these requirements are met. To disentangle these opposing patterns, we use phylogenetic relatedness as an objective metric of species similarities. Studying 11 sunfishes (Centrarchidae) from 890 lakes, we first show no phylogenetic pattern in the raw community data. We then regressed sunfish presence/absence against seven environmental variables and show that lakes with similar water clarity and latitude likely contain closely related species. After statistically removing the environmental effects, phylogenetic repulsion was apparent, with closely related sunfishes less likely to co-occur. Thus, both phylogenetic attraction, driven by environmental filtering, and phylogenetic repulsion, possibly caused by competition, simultaneously occur and obscure one another in the overall phylogenetic structure of sunfish communities.

Phylogenetic measures of biodiversity.

We developed a theoretical framework based on phylogenetic comparative methods to integrate phylogeny into three measures of biodiversity: species variability, richness, and evenness. These metrics can be used in conjunction with permutation procedures to test for phylogenetic community structure. As an illustration, we analyzed data on the composition of 58 lake fish communities in Wisconsin. The fish communities showed phylogenetic underdispersion, with communities more likely to contain closely related species. Using information about differences in environmental characteristics among lakes, we demonstrated that phylogenetic underdispersion in fish communities was associated with environmental factors. For example, lakes with low pH were more likely to contain species in the same clade of acid-tolerant species. Our metrics differ from existing metrics used to calculate phylogenetic community structure, such as net relatedness index and Faith's phylogenetic diversity. Our metrics have the advantage of providing an integrated and easy-to-understand package of phylogenetic measures of species variability, richness, and evenness with well-defined statistical properties. Furthermore, they allow the easy evaluation of contributions of individual species to different aspects of the phylogenetic organization of communities. Therefore, these metrics should aid with the incorporation of phylogenetic information into strategies for understanding biodiversity and its conservation.

A guide to phylogenetic metrics for conservation, community ecology and macroecology.

The use of phylogenies in ecology is increasingly common and has broadened our understanding of biological diversity. Ecological sub-disciplines, particularly conservation, community ecology and macroecology, all recognize the value of evolutionary relationships but the resulting development of phylogenetic approaches has led to a proliferation of phylogenetic diversity metrics. The use of many metrics across the sub-disciplines hampers potential meta-analyses, syntheses, and generalizations of existing results. Further, there is no guide for selecting the appropriate metric for a given question, and different metrics are frequently used to address similar questions. To improve the choice, application, and interpretation of phylo-diversity metrics, we organize existing metrics by expanding on a unifying framework for phylogenetic information. Generally, questions about phylogenetic relationships within or between assemblages tend to ask three types of question: how much; how different; or how regular? We show that these questions reflect three dimensions of a phylogenetic tree: richness, divergence, and regularity. We classify 70 existing phylo-diversity metrics based on their mathematical form within these three dimensions and identify 'anchor' representatives: for α-diversity metrics these are PD (Faith's phylogenetic diversity), MPD (mean pairwise distance), and VPD (variation of pairwise distances). By analysing mathematical formulae and using simulations, we use this framework to identify metrics that mix dimensions, and we provide a guide to choosing and using the most appropriate metrics. We show that metric choice requires connecting the research question with the correct dimension of the framework and that there are logical approaches to selecting and interpreting metrics. The guide outlined herein will help researchers navigate the current jungle of indices.