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Hymenopterans make up about 20% of all animal species, but most are poorly known and lack high-quality genomic resources. One group of important, yet understudied hymenopterans are parasitoid wasps in the family Braconidae. Among this understudied group is the genus Cotesia, a clade of ~1,000 species routinely used in studies of physiology, ecology, biological control, and genetics. However, our ability to understand these organisms has been hindered by a lack of genomic resources. We helped bridge this gap by generating a high-quality genome assembly for the parasitoid wasp, Cotesia glomerata (Braconidae; Microgastrinae). We generated this assembly using multiple sequencing technologies, including Oxford Nanopore, whole-genome shotgun sequencing, and 3D chromatin contact information (HiC). Our assembly is one of the most contiguous, complete, and publicly available hymenopteran genomes, represented by 3,355 scaffolds with a scaffold N50 of ~28 Mb and a BUSCO score of ~99%. Given the genome sizes found in closely related species, our genome assembly was ~50% larger than expected, which was apparently induced by runaway amplification of 3 types of repetitive elements: simple repeats, long terminal repeats, and long interspersed nuclear elements. This assembly is another step forward for genomics across this hyperdiverse, yet understudied order of insects. The assembled genomic data and metadata files are publicly available via Figshare (https://doi.org/10.6084/m9.figshare.13010549).

Hymenopteran species in which sex is determined through a haplo-diploid mechanism known as complementary sex determination (CSD) are vulnerable to a unique form of inbreeding depression. Diploids heterozygous at one or more CSD loci develop into females but diploids homozygous at all loci develop into diploid males, which are generally sterile or inviable. Species with multiple polymorphic CSD loci (ml-CSD) may have lower rates of diploid male production than species with a single CSD locus (sl-CSD), but it is not clear if polymorphism is consistently maintained at all loci. Here, we assess the rate of diploid male production in a population of Cotesia rubecula, a two-locus CSD parasitoid wasp species, approximately 20 years after the population was introduced for biological control. We show that diploid male production dropped from 8-13% in 2005 and 2006 to 3-4% by 2015. We also show from experimental crosses that the population maintained polymorphism at both CSD loci in 2015. We use theory and simulations to show that balancing selection on all CSD alleles promotes polymorphism at several loci in ml-CSD populations. Our study supports the hypothesis that ml-CSD populations have lower diploid male production and are more likely to persist than comparable sl-CSD populations.

Background: The genetic diversity within populations has been shown to affect ecosystem functions, including productivity and invasion resistance. To date most experiments have focused on manipulation of genotypic richness and have ignored other measures of genetic diversity.Aims: In the present study we aimed to establish whether manipulated genotypic richness and genomic dissimilarity of Andropogon gerardii affect productivity and invasion resistance.Methods: We created experimental mesocosms with three levels of genotypic richness: one-, three-, and nine-genotypes. In the three-genotype treatment, we manipulated a range of genomic dissimilarity values (genetic relatedness among individuals). At the end of one growing season we measured above-ground, below-ground and total biomass of the mesocosms, and invasion resistance to Andropogon bladhii.Results: Overall, we found no significant effect of genotypic richness on any measure of ecosystem function, although there tended to be more root biomass (due to...

Contemporary insights from evolutionary ecology suggest that population divergence in ecologically important traits within predators can generate diversifying ecological selection on local community structure. Many studies acknowledging these effects of intraspecific variation assume that local populations are situated in communities that are unconnected to similar communities within a shared region. Recent work from metacommunity ecology suggests that species dispersal among communities can also influence species diversity and composition but can depend upon the relative importance of the local environment. Here, we study the relative effects of intraspecific phenotypic variation in a fish predator and spatial processes related to plankton species dispersal on multitrophic lake plankton metacommunity structure. Intraspecific diversification in foraging traits and residence time of the planktivorous fish alewife (Alosa pseudoharengus) among coastal lakes yields lake metacommunities supporting three lake types which differ in the phenotype and incidence of alewife: lakes with anadromous, landlocked, or no alewives. In coastal lakes, plankton community composition was attributed to dispersal versus local environmental predictors, including intraspecific variation in alewives. Local and beta diversity of zooplankton and phytoplankton was additionally measured in response to intraspecific variation in alewives. Zooplankton communities were structured by species sorting, with a strong influence of intraspecific variation in A. pseudoharengus. Intraspecific variation altered zooplankton species richness and beta diversity, where lake communities with landlocked alewives exhibited intermediate richness between lakes with anadromous alewives and without alewives, and greater community similarity. Phytoplankton diversity, in contrast, was highest in lakes with landlocked alewives. The results indicate that plankton dispersal in the region supplied a migrant pool that was strongly structured by intraspecific variation in alewives. This is one of the first studies to demonstrate that intraspecific phenotypic variation in a predator can maintain contrasting patterns of multitrophic diversity in metacommunities.

In environments characterized by regional heterogeneity among patches, competitor diversity can enhance ecosystem functions such as biomass production. Studies that have addressed the strength of diversity effects in heterogeneous environments have primarily considered a patchy distribution of resources. However, in many systems, top–down effects influence competitor productivity and composition. We use a three-trophic level consumer–resource model to ask how differential responses to predation influence consumer diversity effects at two scales; 1) in patches with and without predator populations, and 2) at a ‘regional’ scale, consisting of one patch with- and one patch without a predator population. At the local scale, the strength and direction of consumer diversity effects depended on the strength of the differential response to predation. Positive or negative influences of consumer richness on equilibrium consumer biomass were the result of a selection effect of diversity. At the regional scale, we observed transgressive overyielding driven by a positive complementarity effect for parameters that define a strong differential response to predation. Given the prevalence of spatially and temporally heterogeneous top–down effects on competitor composition in many ecosystems and trophic levels, we advocate consideration of differential predation as an important step towards incorporating realistic trophic complexity into diversity–function studies.

Predation has important cascading impacts on primary producer biomass and community composition in many ecosystems. While most studies have focused on the consequences of interspecific or density differences in predators, it is recognized that phenotypic variation within species can have strong and cascading community and ecosystem consequences at lower trophic levels. In coastal New England lakes, both the presence and life history form of the zooplanktivorous fish alewife, Alosa pseudoharengus, have strong influence on the biomass, size structure and community composition of crustacean zooplankton communities. Here we test the hypothesis that alewife presence and life history will have cascading impacts on phytoplankton biomass and community composition in a mesocosm experiment that previously reported strong biomass and compositional differences of crustacean zooplankton communities among alewife treatments. We show that alewife life history led to small but statistically significant differences in phytoplankton community composition among treatments. This compositional difference was driven primarily by an increase in the density of two edible phytoplankton genera associated with lower zooplankton biomass in the anadromous alewife treatment. Our results show that intraspecific variation in a predator can have cascading effects on primary producer communities. However we did not observe significant differences in total algal biomass.

Coarse woody habitat (CWH) in the littoral zone is an important habitat feature in freshwater systems and has been suggested to influence nest density and nest site selection by black basses Micropterus spp. (e.g., largemouth bass M. salmoides and smallmouth bass M. dolomieu). To test for a relationship between nest site selection or nest density and the abundance of littoral CWH, we monitored largemouth bass nest site selection in the littoral zones of two small, northern Wisconsin lakes (comprising a total of three separated basins) for three consecutive spawning seasons. Our study sites varied in natural and manipulated abundances of CWH; spawning seasons before and after a whole-basin CWH reduction or a whole-basin CWH addition were examined. Within-basin analysis provided some evidence that local variation in CWH abundance influenced local nest density; however, this relationship was only significant for one basin in a single season. Among basins and across seasons, we observed a positive bu...

One of the oldest questions in ecology is how species diversity in any given trophic level is related to the availability of essential resources that limit biomass (e.g. water, nutrients, light or prey). Researchers have tried to understand this relationship by focusing either on how diversity is influenced by the availability of resources, or alternatively, how resource abundance is influenced by species diversity. These contrasting perspectives have led to a seeming paradox '... is species diversity the cause or the consequence of resources that limit community biomass?' Here we present results of an experiment that show it is possible for species diversity and resource density to exhibit reciprocal causal relationships in the same ecological system. Using a guild of ladybeetle predators and their aphid prey, we manipulated the number of predator species in field enclosures to examine how predator diversity impacts prey population size. At the same time, we manipulated the abundance of aphid prey in discrete habitat patches within each enclosure to determine how smaller-scale spatial variation in resource abundance affects the number of co-occurring predator species. We found that the number of ladybeetle species added to enclosures had a significant impact on aphid population dynamics because interference competition among the predators reduced per capita rates of predation and, in turn, the overall efficiency of the predator guild. At the same time, spatial variation in aphid abundance among smaller habitat patches generated variation in the observed richness of ladybeetles because more species occurred in patches where predators aggregated in response to high aphid density. The results of our experiment demonstrate that it is possible for species diversity to simultaneously be a cause and a consequence of resource density in the same ecological system, and they shed light on how this might occur for groups of mobile consumers that exhibit rapid responses to spatial and temporal variation in their prey.

Accelerating rates of species extinction have prompted a growing number of researchers to manipulate the richness of various groups of organisms and examine how this aspect of diversity impacts ecological processes that control the functioning of ecosystems. We summarize the results of 44 experiments that have manipulated the richness of plants to examine how plant diversity affects the production of biomass. We show that mixtures of species produce an average of 1.7 times more biomass than species monocultures and are more productive than the average monoculture in 79% of all experiments. However, in only 12% of all experiments do diverse polycultures achieve greater biomass than their single most productive species. Previously, a positive net effect of diversity that is no greater than the most productive species has been interpreted as evidence for selection effects, which occur when diversity maximizes the chance that highly productive species will be included in and ultimately dominate the biomass of polycultures. Contrary to this, we show that although productive species do indeed contribute to diversity effects, these contributions are equaled or exceeded by species complementarity, where biomass is augmented by biological processes that involve multiple species. Importantly, both the net effect of diversity and the probability of polycultures being more productive than their most productive species increases through time, because the magnitude of complementarity increases as experiments are run longer. Our results suggest that experiments to date have, if anything, underestimated the impacts of species extinction on the productivity of ecosystems.

Over the past decade an increasing number of studies have experimentally manipulated the number of species in a community and examined how this alters the aggregate production of species biomass. Many of these studies have shown that the effects of richness on biomass change through time, but we have limited understanding of the mechanisms that produce these dynamic trends. Here we report the results of an experiment in which we manipulated the richness of freshwater algae in laboratory microcosms. We used two experimental designs (additive and substitutive) that make different assumptions about how patches are initially colonized, and then tracked the development of community biomass from the point of initial colonization through a period of 6-12 generations of the focal species. We found that the effect of initial species richness on biomass production qualitatively shifted twice over the course of the experiment. The first shift occurred as species transitioned from density-independent to dependent phases of population growth. At this time, intraspecific competition caused monocultures to approach their respective carrying capacities more slowly than polycultures. As a consequence, species tended to over-yield for a brief time, generating a positive, but transient effect of diversity on community biomass. The second shift occurred as communities approached carrying capacity. At this time, strong interspecific interactions caused biomass to be dominated by the competitively superior species in polycultures. As this species had the lowest carrying capacity, a negative effect of diversity on biomass resulted in late succession. Although these two shifts produced dynamics that appeared complex, we show that the patterns can be fit to a simple Lotka-Volterra model of competition. Our results suggest that the effects of algal diversity on primary production change in a predictable sequence through successional time.

1. One of the oldest questions in ecology is how species diversity in any given trophic level is related to the availability of essential resources that limit biomass (e.g. water, nutrients, light or prey). Researchers have tried to understand this relationship by focusing either on how diversity is influenced by the availability of resources, or alternatively, how resource abundance is influenced by species diversity. These contrasting perspectives have led to a seeming paradox '... is species diversity the cause or the consequence of resources that limit community biomass?' 2. Here we present results of an experiment that show it is possible for species diversity and resource density to exhibit reciprocal causal relationships in the same ecological system. Using a guild of ladybeetle predators and their aphid prey, we manipulated the number of predator species in field enclosures to examine how predator diversity impacts prey population size. At the same time, we manipulated the abundance of aphid prey in discrete habitat patches within each enclosure to determine how smaller-scale spatial variation in resource abundance affects the number of co-occurring predator species. 3. We found that the number of ladybeetle species added to enclosures had a significant impact on aphid population dynamics because interference competition among the predators reduced per capita rates of predation and, in turn, the overall efficiency of the predator guild. At the same time, spatial variation in aphid abundance among smaller habitat patches generated variation in the observed richness of ladybeetles because more species occurred in patches where predators aggregated in response to high aphid density. 4. The results of our experiment demonstrate that it is possible for species diversity to simultaneously be a cause and a consequence of resource density in the same ecological system, and they shed light on how this might occur for groups of mobile consumers that exhibit rapid responses to spatial and temporal variation in their prey.

Over the past decade, accelerating rates of species extinction have prompted an increasing number of studies to reduce the number of species experimentally in a variety of ecosystems and examine how this aspect of diversity alters the efficiency by which communities capture biologically essential resources and convert them into new tissue. Here we summarize the results of 164 experiments (reported in 84 publications) that have manipulated the richness of primary producers, herbivores, detritivores, or predators in a variety of terrestrial and aquatic ecosystems and examined how this impacts (1) the standing stock abundance or biomass of the focal trophic group, (2) the abundance or biomass of that trophic group's primary resource(s), and/or (3) the extent to which that trophic group depletes its resource(s). Our summary includes studies that have focused on the top-down effects of diversity, whereby researchers have examined how the richness of trophic group t impacts the consumption of a shared resource, and also studies that have focused on the bottom-up effects of diversity, whereby researchers have examined how the richness of trophic group t impacts consumption of t by the next highest trophic level. The first portion of the data set provides information about the source of data and relevant aspects of the experimental design, including the spatial and temporal scales at which the work was performed. The second portion gives the magnitude of each response variable, the standard deviation, and the level of replication at each level of species richness manipulated. The third portion of the data set summarizes the magnitude of diversity effects in two ways. First, log ratios are used to compare the response variable in the most diverse polyculture to either the mean of all monocultures or the species having the highest/lowest value in monoculture. Second, data from each level of species richness are fit to three nonlinear functions (log, power, and hyperbolic) to assess which best characterizes the shape of diversity effects. The final portion of the data set summarizes any information that helps parse diversity effects into that attributable to species richness vs. that attributable to changes in species composition across levels of richness. The complete data sets corresponding to abstracts published in the Data Papers section of the journal are published electronically in Ecological Archives at 〈http://esapubs.org/archive〉. (The accession number for each Data Paper is given directly beneath the title.)