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Rodent population control is a global problem, complicated by evolved non-responsiveness to rodenticide treatment. Contraceptives could help mitigate this challenge, but questions remain about their efficacy, especially for rodenticide-resistant populations. We used an age-dependent demographic model to generate two hypotheses: Fertility reduction applied early in female lifetimes (1) is more effective in controlling rodent populations than when applied later in female lifetimes, and (2) is effective in controlling rodent populations that are expanding. Compared to controls, fertility reduction applied early, in mid-life, and late in female lifetimes, decreased, matched, and accelerated, respectively, the rates of population growth. Fertility reduction was effective in reducing population size only when sustained over multiple generations and was ineffective when application was episodic. Substituting classic Rattus norvegicus and Microtus agrestis life history data into our simulation framework confirmed that early fertility reduction was effective in controlling population growth, including expanding populations in both species. These simulations generated two additional hypotheses for field applications of fertility control: Over treatment durations, (3) the fraction of the population consisting of juveniles, and (4) the overall population size, will both decrease. We tested these predictions using a 12-month contraceptive bait application on rats in two urban US locations (Washington, DC) where rodenticides were already deployed. Consistent with our predictions, these populations showed marked decreases in the proportion of juvenile to adult rats, and in the total number of rats observed in camera traps over the study period. Our results support fertility control as an effective method for managing rodent populations.

Evolution has been viewed as occurring primarily through selection among individuals. We present a framework based on multilevel selection for evaluating evolutionary change from individuals to communities, with supporting empirical evidence. Essential to this evaluation is the role that interspecific indirect genetic effects play in shaping community organization, in generating variation among community phenotypes, and in creating community heritability. If communities vary in phenotype, and those phenotypes are heritable and subject to selection at multiple levels, then a community view of evolution must be merged with mainstream evolutionary theory. Rapid environmental change during the Anthropocene will require a better understanding of these evolutionary processes, especially selection acting at the community level, which has the potential to eliminate whole communities while favoring others.

A key issue in evolutionary biology is whether selection acting at levels higher than the individual can cause evolutionary change. If it can, then conceptual and empirical studies must consider how selection operates at multiple levels of biological organization. Here, we test the hypothesis that estimates of broad-sense community heritability

Several studies have demonstrated the ecological consequences of genetic variation within a single plant species. For example, these studies show that individual plant genotypes support unique composition of the plants' associated arthropod community. By contrast, fewer studies have explored how plant genetic variation may influence evolutionary dynamics in the plant's associated species. Here, we examine how aphids respond evolutionarily to genetic variation in their host plant. We conducted two experiments to examine local adaptation and rapid evolution of the free‐feeding aphid Chaitophorus populicola across genetic variants of its host plant, Populus angustifolia. To test for local adaptation, we collected tree cuttings and aphid colonies from three sites along an elevation/climate gradient and conducted a reciprocal transplant experiment. In general, home aphids (aphids transplanted onto trees from the same site) produced 1.7–3.4 times as many offspring as foreign aphids (aphids transplanted onto trees from different sites). To test for rapid evolution, we used 4 clonally replicated aphid genotypes and transplanted each onto 5 clonally replicated P. angustifolia genotypes. Each tree genotype started with the same aphid genotype composition. After 21 days (~two aphid generations), aphid genotype composition changed (i.e., aphids evolved) and some tree genotypes supported unique evolutionary trajectories of aphids. These results suggest that plant evolution in response to human perturbation, such as climate change and invasive species, will also result in evolutionary responses in strongly interacting species that could cascade to affect whole communities., We conducted two experiments to examine local adaptation and rapid evolution of the free‐feeding aphid Chaitophorus populicola across a genetic gradient of its host plant, Populus angustifolia. We found evidence of local adaptation, such that local aphids produced 1.7–3.4 times as many offspring as foreign aphids. Further, in just 21 days aphids rapidly and differentially evolved on different genotypes of their host plant.

The North American beaver (Castor canadensis Kuhl) and cottonwoods (Populus spp.) are foundation species, the interactions of which define a much larger community and affect a threatened riparian habitat type. Few studies have tested the effect of these interactions on plant chemistry and a diverse arthropod community. We experimentally examined the impact of beaver foraging on riparian communities by first investigating beaver food preferences for one cottonwood species, Fremont cottonwood (P. fremontii S. Watson), compared to other locally available woody species. We next examined the impact of beaver foraging on twig chemistry and arthropod communities in paired samples of felled and unfelled cottonwood species in northern Arizona (P. fremontii) and southwestern Colorado (narrowleaf cottonwood, P. angustifolia James, and Eastern cottonwood, P. deltoides W. Bartram ex Marshall). Four major patterns emerged: (1) In a cafeteria experiment, beavers chose P. fremontii six times more often than other woody native and exotic species. (2) With two cottonwood species, we found that the nitrogen and salicortin concentrations were up to 45% greater and lignin concentration 14% lower in the juvenile resprout growth of felled trees than the juvenile growth on unfelled trees (six of seven analyses were significant for P. fremontii and four of six were significant for P. angustifolia). (3) With two cottonwood species, arthropod community composition on juvenile branches differed significantly between felled and unfelled trees, with up to 38% greater species richness, 114% greater relative abundance and 1282% greater species diversity on felled trees (six of seven analyses with P. fremontii and four of six analyses with P. angustifolia were significant). The above findings indicate that the highest arthropod diversity is achieved in the heterogenous stands of mixed felled and unfelled trees than in stands of cottonwoods, where beavers are not present. These results also indicate that beaver herbivory changes the chemical composition in 10 out of 13 chemical traits in the juvenile growth of two of the three cottonwood species to potentially allow better defense against future beaver herbivory. (4) With P. deltoides, only one of five analyses in chemistry was significant, and none of the four arthropod community analyses were significant, suggesting that this species and its arthropod community responds differently to beaver. Potential reasons for these differences are unknown. Overall, our findings suggest that in addition to their impact on riparian vegetation, other mammals, birds, and aquatic organisms, beavers also may define the arthropod communities of two of three foundation tree species in these riparian ecosystems.

Prairie dogs (Cynomys spp.) produce an alarm call when a predator appears. Gunnison's prairie dogs (Cynomys gunnisoni) produce calls that vary in spectral structure according to predator type and specific characteristics unique to the individual predator, such as color and shape. These calls vary depending on geographic location and have been characterized as dialects. Black-tailed prairie dogs (Cynomys ludovicianus) also encode information about terrestrial predators in their alarm calls. However, nothing is known about whether there are dialects in these alarm calls, and if so, why these dialects might have arisen. We studied the alarm calls and habitat characteristics of eight different black-tailed prairie dog colonies within the Southwestern USA: Maxwell National Wildlife Refuge, NM, Vermejo Park Ranch, NM, Kiowa National Grasslands, NM, Rita Blanca National Grasslands, TX, and Comanche National Grasslands, CO. An analysis of time and frequency changes in alarm calls showed that colony sites that were located within 30 km each other did not have significant differences in alarm call structure, but sites that were more distant were significantly different from the other sites, consistent with the existence of dialects as well as with genetic models of isolation by distance. The acoustic structure of the alarm calls was not explained by differences in habitat characteristics at each site. A comparison of Squared Euclidian Distances (D2) of alarm calls versus linear distances between sites showed a significant positive correlation, implying that alarm call dialect differences increase the farther one site is from another. This relationship is consistent with a cultural diffusion model of dialects, and is similar to that observed in human dialects.

The functional advantages of arsenic (As) hyperaccumulation by plants are poorly understood. One proposed benefit, termed elemental allelopathy, occurs when hyperaccumulated As is cycled from the plant back into the top layer of soil, allowing As hyperaccumulators to gain an advantage over intolerant species by increasing soil As concentrations ([As]) underneath their canopy. To date, there are no studies that detail the presence of increased soil [As] associated with As hyperaccumulators. In this study, we documented variation in the soil [As] associated with the Chinese brake fern, Pteris vittata L. and also compared the effects of environmentally relevant soil and solution [As] on competitor plant growth. Four populations of P. vittata were identified in central Florida, USA. P. vittata tissue samples and soil samples were collected at the base of and at 3 m away from ferns in each population (n = 36). Five sample locations were randomly selected from each site, and soils from the base and 3 m away from each fern were collected to examine the effects of naturally occurring soil [As] on the germination and growth of a potential competitor plant (Oxalis stricta). Solutions with increasing [As] were also used to examine the threshold for negative effects of [As] on O. stricta growth. [As] were measured using inductively coupled plasma mass spectrometry (ICP-MS). Overall, soil [As] from the base of ferns was nearly twice that of soil 3 m away indicating that ferns hyperaccumulate As. However, ferns and their associated soil, contained different [As] depending on their collection site, indicating that these populations accumulate and use [As] differently. O. stricta growth decreased and germination was delayed as solution and soil [As] increased. However, the relative distance from the fern that the soil was collected from did not affect growth, which would be expected with elemental allelopathy. Our results show that P. vittata is associated with higher soil [As] and these concentrations are sufficient to inhibit growth of competitors. However, the absence of a strong inhibitory relationship associated with proximity to the fern across all locations suggests that the possible functional advantages of elemental allelopathy may depend on site specific characteristics.

A new leech species Helobdella blinni sp. n., is described from Montezuma Well, an isolated travertine spring mound located in central Arizona, USA. In its native habitat, Helobdella blinni had been previously identified as Helobdella stagnalis (Linnaeus, 1758), which was later reclassified to Helobdella modesta (Verrill, 1872). Similar to the European Helobdella stagnalis and North American Helobdella modesta, Helobdella blinni has six pairs of testisacs, five pairs of smooth crop caecae, one lobed pair of posteriorly-directed crop caecae, one pair of eyes, a nuchal scute, and diffuse salivary glands. However, the pigmentation of this new species ranges from light to dark brown, unlike Helobdella modesta which tends to be light grey in color. Also, Helobdella modesta produces a clutch of 12-–35 pink eggs, whereas Helobdella blinni produces smaller clutches of white eggs (7–14, 0.5 ± 0.15 mm, N = 7) and consequently broods fewer young (1–14, 7 ± 3.3 mm, N = 97). Helobdella blinni are also able to breed year-round due to the constant warm water conditions in Montezuma Well. Their breeding season is not restricted by seasonal temperatures. These species are morphologically similar, however, comparing the COI mtDNA sequences of Helobdella blinni with sequences from nearby populations of Helobdella modesta and other Helobdella species from GenBank indicate that Helobdella blinni is genetically distinct from these other Helobdella populations.

Alternative mating tactics appear to evolve when sexual selection is strong. Because such conditions are usually observed in species with polygynous or polyandrous mating systems, alternative mating tactics in monogamous mating systems are seldom documented and are poorly understood. In prairie voles, Microtus ochrogaster, a species widely known for forming monogamous pair-bonds, the expression of territoriality within each sex is dimorphic, and includes non-territorial “wanderers” as well as territorial “residents.” To explore the variance in fitness, measured in offspring numbers, among breeding individuals expressing these alternative mating tactics, we compiled parentage data over three years for male and female prairie voles from natural populations in Indiana and Kansas, USA. We found that: (1) the average fitnesses of males and females within each population were identical when adjusted by the sex ratio; (2) the variance in fitness in male and female prairie voles was comparable to that of many highly polygynous species; (3) The average fitnesses of male tactics, and of female tactics, were equivalent within and among years within each location; (4) consistent with negative frequency-dependent selection acting on mating phenotypes, the between-tactic variance in fitness for male and female mating tactics decreased with increasing study duration; (5) consistent with negative assortative mating, resident males and wanderer females produced offspring primarily in monogamous partnerships, whereas wanderer males and resident females produced offspring primarily in polygamous partnerships. Our results show that the conditions necessary for the persistence of alternative mating tactics are indistinguishable from those for phenotypically less flexible alternative mating strategies, and that alternative mating tactics can evolve in both sexes in monogamous species when fitness variance within each sex is large.

We examined the hypothesis that ecological niche models (ENMs) more accurately predict species distributions when they incorporate information on population genetic structure, and concomitantly, local adaptation. Local adaptation is common in species that span a range of environmental gradients (e.g., soils and climate). Moreover, common garden studies have demonstrated a covariance between neutral markers and functional traits associated with a species' ability to adapt to environmental change. We therefore predicted that genetically distinct populations would respond differently to climate change, resulting in predicted distributions with little overlap. To test whether genetic information improves our ability to predict a species' niche space, we created genetically informed ecological niche models (gENMs) using Populus fremontii (Salicaceae), a widespread tree species in which prior common garden experiments demonstrate strong evidence for local adaptation. Four major findings emerged: (i) gENMs predicted population occurrences with up to 12-fold greater accuracy than models without genetic information; (ii) tests of niche similarity revealed that three ecotypes, identified on the basis of neutral genetic markers and locally adapted populations, are associated with differences in climate; (iii) our forecasts indicate that ongoing climate change will likely shift these ecotypes further apart in geographic space, resulting in greater niche divergence; (iv) ecotypes that currently exhibit the largest geographic distribution and niche breadth appear to be buffered the most from climate change. As diverse agents of selection shape genetic variability and structure within species, we argue that gENMs will lead to more accurate predictions of species distributions under climate change.

Although genetics in a single species is known to impact whole communities, little is known about how genetic variation influences species interaction networks in complex ecosystems. Here, we examine the interactions in a community of arthropod species on replicated genotypes (clones) of a foundation tree species, Populus angustifolia James (narrowleaf cottonwood), in a long-term, common garden experiment using a bipartite "genotype-species" network perspective. We combine this empirical work with a simulation experiment designed to further investigate how variation among individual tree genotypes can impact network structure. Three findings emerged: (1) the empirical "genotype-species network" exhibited significant network structure with modularity being greater than the highly conservative null model; (2) as would be expected given a modular network structure, the empirical network displayed significant positive arthropod co-occurrence patterns; and (3) furthermore, the simulations of "genotype-species" networks displayed variation in network structure, with modularity in particular clearly increasing, as genotypic variation increased. These results support the conclusion that genetic variation in a single species contributes to the structure of ecological interaction networks, which could influence eco-ogical dynamics (e.g., assembly and stability) and evolution in a community context.

Introduced species are known to have large impacts on native ecosystems, by extirpating species,altering species interactions and changing community composition. The mechanisms by which intro-duced species impact native communities often include both direct and indirect effects. Here, wecombined observational field surveys, an exclosure experiment, network analysis and structural equationmodeling to examine the direct and indirect effects of an introduced species on native communities.Specifically, we investigated how introduced elk indirectly impact the composition and co-occurrencepatterns within a community of plant-associated arthropods by directly altering the phenotype of anative plant, Solidago velutina. Surveying the arthropods associated with the plant's inflorescence, twomain patterns emerged. i) Using field observations across 500 km 2 and an exclosure experiment,S. velutina growing in the presence of elk had 67e90% fewer flowering ramets,15e85% lower percentageof flowering ramets, 33e45% fewer florets per inflorescence and 25e45% lower sexual ramet biomass. ii)Using the same exclosures, the arthropod community on S. velutina in the presence of elk, had 45% fewerspecies, a 70% reduction in abundance and a significant change to the species composition and co-occurrence network structure. The results from the network analysis suggested that introduced spe-cies' impacts on communities can be morethan changes in richness or abundance, but include changes tospecies interactions. Structural equation modeling showed that elk caused a decrease in inflorescencesize of S. velutina, which affected the arthropod community, suggesting that communities can changewithout extirpation of their host plant species. These results highlight the importance of changingintraspecific variation as a mechanism by which invasive species alter community composition andspecies interactions.© 2015 Elsevier Masson SAS. All rights reserved.

Summary 1. Community genetics studies frequently focus on individual communities associated with individual plant genotypes, but little is known about the genetically based relationships among taxonomically and spatially disparate communities. We integrate studies of a wide range of communities living on the same plant genotypes to understand how the ecological and evolutionary dynamics of one community may be constrained or modulated by its underlying genetic connections to another community. 2. We use pre-existing data sets collected from Populus angustifolia (narrowleaf cottonwood) growing in a common garden to test the hypothesis that the composition of pairs of distinct communities (e.g. endophytes, pathogens, lichens, arthropods, soil microbes) covary across tree genotypes, such that individual plant genotypes that support a unique composition of one community are more likely to support a unique composition of another community. We then evaluate the hypotheses that physical proximity, taxonomic similarity, time between sampling (time attenuation), and interacting foundation species within communities explain the strength of correlations. 3. Three main results emerged. First, Mantel tests between communities revealed moderate to strong (q = 0.25‐0.85) community‐genetic correlations in almost half of the comparisons; correlations among phyllosphere endophyte, pathogen and arthropod communities were the most robust. Secondly, physical proximity determined the strength of community‐genetic correlations, supporting a physical proximity hypothesis. Thirdly, consistent with the interacting foundation species hypothesis, the most abundant species drove many of the stronger correlations. Other hypotheses were not supported. 4. Synthesis. The field of community genetics demonstrates that the structure of communities varies among plant genotypes; our results add to this field by showing that disparate communities covary among plant genotypes. Eco-evolutionary dynamics between plants and their associated organisms may therefore be mediated by the shared connections of different communities to plant genotype, indicating that the organization of biodiversity in this system is genetically based and non-neutral.

Because introduced species may strongly interact with native species and thus affect their fitness, it is important to examine how these interactions can cascade to have ecological and evolutionary consequences for whole communities. Here, we examine the interactions among introduced Rocky Mountain elk, Cervus canadensis nelsoni, a common native plant, Solidago velutina, and the diverse plant-associated community of arthropods. While introduced species are recognized as one of the biggest threats to native ecosystems, relatively few studies have investigated an evolutionary mechanism by which introduced species alter native communities. Here, we use a common garden design that addresses and supports two hypotheses. First, native S. velutina has rapidly evolved in the presence of introduced elk. We found that plants originating from sites with introduced elk flowered nearly 3 weeks before plants originating from sites without elk. Second, evolution of S. velutina results in a change to the plant-associated arthropod community. We found that plants originating from sites with introduced elk supported an arthropod community that had ~35 % fewer total individuals and a different species composition. Our results show that the impacts of introduced species can have both ecological and evolutionary consequences for strongly interacting species that subsequently cascade to affect a much larger community. Such evolutionary consequences are likely to be long-term and difficult to remediate.

Author(s): Grady, Kevin C; Wood, Troy E; Kolb, Thomas E; Hersch-Green, Erika; Shuster, Stephen M; Gehring, Catherine A; Hart, Stephen C; Allan, Gerard J; Whitham, Thomas G

Cuticular pigmentation is highly variable in Paracerceis sculpta Holmes, 1904, a Gulf of California isopod. Individuals expressing L2r (= Laterals-2 red) have red pigmentation on the lateral margins of their 6-7th body segments. Individuals expressing 3rs (= Three red stripes) have three red pigmentation zones running the length of their bodies. Individuals expressing Cd (= Cephalon dark) have black head capsules. In 12 years of field collections, L2r and 3rs represented less than 0.3% of all individuals (N=5491) with the frequency of each sex proportional to population frequencies. Of all individuals scored as Cd (N=178), 98% were females. We crossed marked and unmarked parents in all possible combinations. Progeny ratios for L2r and 3rs met Mendelian expectations within families as did adult expression of 3rs despite observed biases in family sex ratio. In three generations, Cd females crossed to unmarked males produced families with 1:1 sex ratios, 100% Cd daughters and no Cd sons. Sons from these families never produced Cd daughters. Our results suggest that L2r and 3rs are controlled by dominant, autosomal alleles. The sex-limited expression of Cd corroborates previous results suggesting female heterogamety in P. sculpta and in other flabelliferan isopods. The appearance of rare phenotypes controlled by dominant alleles is paradoxical given the hypothesis that allelic dominance evolves in response to positive selection. However, this combination might persist if apostatic selection imposed by visual predators occurs in this species’ structured populations, thereby favoring dominance modifiers that suppress fitness losses by heterozygotes. This species with its large number of cuticular markers could provide tests of this hypothesis.

We illustrate a method for delaying and possibly eliminating the evolution of non-responsiveness to the treatments now used to control pest populations. Using simulations and estimates of the variance in relative fitness, i.e., the opportunity for selection, in a rat-like mammal, we show that the selection responsible for the evolution of non-responsiveness to pesticides and sterility-inducers, is similar in its action to sexual selection, and for this reason can be orders of magnitude stronger than that which exists for untreated populations. In contrast, we show that when contraceptives are used to reduce the fertility of a pest species, with non-responders embedded within such populations, the opportunity for selection favoring non-responsiveness is reduced to that which is expected by chance alone. In pest species with separate sexes, we show that efforts to control pest populations or to mitigate selection favoring non-responsiveness, are likely to be ineffective when members of one sex are sterilized or killed. We also show that while mating preferences can impede the rate at which resistance evolves, they are more likely to accelerate this process, arguing against the use of sterile male approaches for controlling pests. Our results suggest that contraceptives are more effective at controlling pest populations and slowing the evolution of non-responsiveness than treatments that cause sterilization or death in target species. Furthermore, our results indicate that contraceptives that work differentially on each sex will be most effective in mitigating selection favoring non-responders. Our results have significant implications for the development and application of treatments to manage pests, now and into the future.

Climate change, particularly increased aridity, poses a significant threat to plants and the biotic communities they support. Dioecious species may be especially vulnerable to climate change given that they often exhibit spatial segregation of the sexes, reinforced by physiological and morphological specialization of each sex to different microhabitats. In dimorphic species, the overexpression of a trait by one gender versus the other may become suppressed in future climates. Data suggest that males will generally be less sensitive to increased aridity than co-occurring females and, consequently, extreme male-biased sex ratios are possible in a significant number of populations. The effects of male-biased sex ratios are likely to cascade to dependent community members, especially those that are specialized on one sex.

Although genetics in a single species is known to impact whole communities, little is known about how genetic variation influences species interaction networks in complex ecosystems. Here, we examine the interactions in a community of arthropod species on replicated genotypes (clones) of a foundation tree species, Populus angustifolia James (narrowleaf cottonwood), in a long-term, common garden experiment using a bipartite "genotype-species" network perspective. We combine this empirical work with a simulation experiment designed to further investigate how variation among individual tree genotypes can impact network structure. Three findings emerged: (1) the empirical "genotype-species network" exhibited significant network structure with modularity being greater than the highly conservative null model; (2) as would be expected given a modular network structure, the empirical network displayed significant positive arthropod co-occurrence patterns; and (3) furthermore, the simulations of "genotype-species" networks displayed variation in network structure, with modularity in particular clearly increasing, as genotypic variation increased. These results support the conclusion that genetic variation in a single species contributes to the structure of ecological interaction networks, which could influence eco-ogical dynamics (e.g., assembly and stability) and evolution in a community context.

The Operational Sex Ratio, or OSR, first defined as ‘the ratio of receptive females to potential mating males at any one time,’ was designed to measure the level of competition for mates in animal populations. The apparent utility of the OSR as a proxy for sexual selection intensity established it as a standard metric for animal mating system research. However, the relationships between OSR and actual measures of sexual selection have proven inconsistent. Thus, while useful for characterizing known experimental populations, estimates of OSR are not equivalent to those for sexual selection and are unlikely account for observed patterns of evolutionary change.

Background and aims Soils can act as agents of natural selection, causing differential fitness among genotypes and/or families of the same plant species, especially when soils have extreme physical or chemical properties. More subtle changes in soils, such as variation in microbial communities, may also act as agents of selection. We hypothesized that variation in soil properties within a single river drainage can be a selective gradient, driving local adaptation in plants. Methods Using seeds collected from individual genotypes of Populus angustifolia James and soils collected from underneath the same trees, we use a reciprocal transplant design to test whether seedlings would be locally adapted to their parental soil type. Results We found three patterns: 1. Soils from beneath individual genotypes varied in pH, soil texture, nutrient content, microbial biomass and the physiological status of microorganisms. 2. Seedlings grown in local soils experienced 2.5-fold greater survival than seedlings planted in non-local soils. 3. Using a composite of height, number of leaves and leaf area to measure plant growth, seedlings grew ∼17.5% larger in their local soil than in non-local soil. Conclusions These data support the hypothesis that variation in soils across subtle gradients can act as an important selective agent, causing differential fitness and local adaptation in plants.

Evolutionary biologists have developed several indices, such as selection gradients (b) and the opportunity for sexual selection (Is), to quantify the actual and ⁄ or potential strength of sexual selection acting in natural or experimental populations. In a recent paper, Klug et al. (J. Evol. Biol. 23, 2010, 447) contend that selection gradients are the only legitimate metric for quantifying sexual selection. They argue that Is and similar mating-systembased metrics provide unpredictable results, which may be uncorrelated with selection acting on a trait, and should therefore be abandoned. We find this view short-sighted and argue that the choice of metric should be governed by the research question at hand. We describe insights that measures such as the opportunity for selection can provide and also argue that Klug et al. have overstated the problems with this approach while glossing over similar issues with the interpretation of selection gradients. While no metric perfectly characterizes sexual selection in all circumstances, thoughtful application of existing measures has been and continues to be informative in evolutionary studies.

The sea star Aquilonastra corallicola reproduces asexually by splitting across the central disc, a process known as fission. Although photoperiod and body mass are known to influence the rate of reproduction in fissiparous asteroid species, to our knowledge, the effects of population density have not been experimentally measured in asteroids. We tested the effects of three population densities on the rate of fission in laboratory populations maintained for 4 months at low, medium, and high (n = 5, 15, and 30 per treatment replicate, respectively) population densities. Our results show that population density has a significant, inverse effect upon rate of reproduction in this species. This suggests that density-dependent regulation of asexual reproduction is a significant factor in explaining the observed patterns of population structure in marine clonal invertebrates.

Genetic variation in plants is known to influence arthropod assemblages and species interactions. However, these influences may be contingent upon local environmental conditions. Here, we examine how plant genotype-based trophic interactions and patterns of natural selection change across environments. Studying the cottonwood tree, Populus angustifolia, the galling aphid, Pemphigus betae and its avian predators, we used three common gardens across an environmental gradient to examine the effects of plant genotype on gall abundance, gall size, aphid fecundity and predation rate on galls. Three patterns emerged: (i) plant genotype explained variation in gall abundance and predation, (ii) G×E explained variation in aphid fecundity, and environment explained variation in gall abundance and gall size, (iii) natural selection on gall size changed from directional to stabilizing across environments.

In Caribbean Panama, nonreproductive male and female stomatopods are solitary and defend their own coral-rubble cavities. When breeding pairs form, however, males assume all responsibility for cavity defense. To compare success in cavity defense and defensive tactics among paired and unpaired males, and to examine the tendency for paired stomatopods to exchange their present mates for larger (higher quality) individuals, we introduced same-sized and 15% larger male, and same-sized and 15% larger reproductive female intruders to paired and unpaired male residents in a balanced design. Paired males were more successful at cavity defense than unpaired males, evidently because paired males strike intruders more than unpaired males, and because intruders fight less intensely against paired males than against unpaired males. Paired males occasionally attempted extrapair copulations, but showed little tendency to abandon their mates in favor of larger females. Paired females, however, mated readily with intruder males that evicted resident males. Populationwide female breeding synchrony and prolonged female receptivity before oviposition reduce variance in male mating success and may force males to guard the breeding cavity to assure their paternity. Uncertainty about the reproductive condition of intruder females may prevent males from exchanging mates.

In Paracerceis sculpta, a Gulf of California isopod, a-males usually defend aggregations of breeding females within the spongocoels of intertidal sponges. Breeding isopods are also found in habitats other than sponges, but interactions among individuals in these locations are poorly known. To investigate plasticity in male-male interactions associated with mate acquisition in breeding habitats other than sponges, we examined the behavior of pairs of a-males in two distinct environments; those in which breeding sites were absent, but females were present, and those in which both breeding sites and females were absent. We established males as residents in the presence and in the absence of females by allowing them to occupy arenas for 1 h before intruders were introduced. Residents and intruders engaged in significantly more total acts and grappled more in the presence of females than when females were absent. In the presence of females, residents became active first in most cases, and gripped females more than intruders. Latency to activity was significantly longer when receptive females were present than when they were absent. However, residents seldom interacted with intruders when females were lacking. Our results suggest that in locations other than sponges, a-males compete for females and can recognize their status as residents and as intruders. These results also suggest that, as in many vertebrate and invertebrate species, P. sculpta a-males attempt to mate opportunistically, and that contests for mates in atypical breeding habitats may be common in nature. We discuss the importance of distinguishing rare and highly contingent behaviors from those likely to represent adaptive behavioral plasticity, and we propose a simple criterion of commonality for their identification in behavioral analyses.

Pea crabs, Dissodactylus primitivus, inhabit multiple echinoid (heart urchin) hosts. Male and female crabs move among hosts in search for mates, and both sexes mate multiple times, creating opportunities for post-copulatory sexual selection. For such selection to occur, only a fraction of the males who succeed in mating can also succeed in siring progeny. Jossart et al. 2014 used 4 microsatellite loci to document parentage and mating frequencies of both sexes in D. primitivus. From these data we identified the mean and variance in female offspring numbers, as well as the proportions of the female population that were gravid and not bearing offspring. We next identified the proportions of the male population who had (1) mated and sired offspring, (2) mated but failed to sire offspring, and (3) failed to mate altogether. We used these results to estimate the opportunity for selection on males and females in terms of mate numbers and offspring numbers, and estimated the sex difference in the opportunity for selection (i.e., the opportunity for sexual selection) using both forms of data. We then partitioned the total variance in male fitness into pre- and post-copulatory components and identified the fraction of the total opportunity for selection occurring in each context. Our results show that the opportunity for selection on each sex was of similar magnitude (0.69–0.98), consistent with this polyandrogynous mating system. We also found that 37% of the total opportunity for sexual selection on males occurred within the context of post-copulatory sexual selection. However, the fraction of the total opportunity for selection that was due to sexual selection, estimated using both mate numbers and offspring numbers, was 9% and 23% respectively. Thus, we further reduced our estimate of the opportunity for post-copulatory sexual selection in D. primitivus to less than 10% of the total opportunity for selection (0.37 of 0.09 and 0.23 = 0.03 and 0.09). Our results provide the first estimate of the maximum possible strength of post-copulatory sexual selection in crustaceans using this approach.

The Socorro isopod, Thermosphaeroma thermophilum, became the focus of a novel conservation strategy when apparent extirpation of the species from the wild prompted con- struction of the Socorro Isopod Propagation Facility (SIPF) near the native spring in Socorro, New Mexico, USA. We subjected captive populations to 4 habitat treatments over 2 consec- utive 50-month experimental trials. Native populations of T. thermophilum remained signifi- cantly more abundant than captive subpopulations during the 100-month study. Population trends among artificial habitats differed slightly between the trials, but the magnitude of this difference was independent of linear flow within either pool series, suggesting that captive isopods responded to habitat quality. Isopod age structure was most heterogeneous in pools with plants suggesting that vertical structural diversity is necessary to maintain long-term via- bility of T. thermophilum in captivity. Observations of 'breeding huddles' in the SIPF pools implied that controlled propagation may affect the social basis of the species' mating system. This could be a behavioral response to selection on body size that may reduce predation risks from larger male cannibals, or to increase fitness of both sexes under spatially altered sex ratios of artificial environments. Our results accentuate the need for continued research on species- specific management practices for geographically restricted invertebrate taxa such as the endangered Socorro isopod.

Can heritable traits in a single species affect an entire ecosystem? Recent studies show that such traits in a common tree have predictable effects on community structure and ecosystem processes. Because these 'community and ecosystem phenotypes' have a genetic basis and are heritable, we can begin to apply the principles of population and quantitative genetics to place the study of complex communities and ecosystems within an evolutionary framework. This framework could allow us to understand, for the first time, the genetic basis of ecosystem processes, and the effect of such phenomena as climate change and introduced transgenic organisms on entire communities.

The endangered Socorro Isopod, Thermosphaeroma thermophilum, is endemic to a single thermal spring in Socorro, NM. This species is cannibalistic, with males more cannibalistic than females, and with females and juveniles more vulnerable than males as prey. In 1990, the New Mexico Department of Game and Fish, created the Socorro Isopod Propagation Facility (SIPF) near the natural habitat, Sedillo Spring (SS), to increase total population size and to examine the effects of habitat heterogeneity on population growth. We report the genetic and morphological effects of this experiment, using the natural population as a control. Captive subpopulations experienced bottlenecks of known intensity and duration, as well as different intensities of cannibalism. Using 57 AFLP markers, we show that in 6 years (1990–1996), captive subpopulations diverged significantly from the natural population. Also during this 6-year period, body lengths of captive isopods diverged nearly 2-fold from the natural population, evidently because cannibalism and thus selection favoring large size was more intense in captive subpopulations than in nature. This hypothesis is supported by the fact that cannibalism and the apparent response to selection on body size became variable among captive subpopulations when physical structure was added to three of the four SIPF pools in April 1997. As expected if cannibalism was the source of selection for large body size, by August 1998 (15 months = 7–8 generations), the rate at which body size increased became inversely proportional to the amount of physical structure within pools. Although we are unable to separate the specific effects of population subdivision and cannibalism, our results show that these conditions in combination caused rapid changes in genetic variation and the external morphology of these captive subpopulations. Our results have important implications for future attempts to manage and propagate endangered species.

A population of the isopod Paracerceis sculpta Holmes, 1904 was found in Port Aransas, Texas in Northern Gulf of Mexico in 2009, the first report of the species in the Gulf of Mexico. The last large isopod survey in the area was conducted in 1981 implying that P. sculpta invaded some time in the last 28 years. Here, we show size frequency distributions of 858 individuals, alpha males (mean length = 5.48 mm ± 0.09 SE), pre-alpha males (3.94 mm ± 0.15), females (2.76 mm ± 0.03) and juveniles (1.24 mm ± 0.02). Beta and gamma males are absent in Texas, a major contrasting difference between the invasive and native populations.

Nymphophilus minckleyi is a hydrobiid snail endemic to the freshwater spring ecosystem of Cuatro Cienegas, Mexico. We used seven allozyme loci to examine the genetic substructure of N. minckleyi from 14 sites (subpopulations) in the basin and to test the hypothesis that spring pools in Cuatro Cienegas are separated into seven hydrologically distinct drainages. Hierarchical F-statistics suggest significant population structure exists among the fourteen populations but not among the seven proposed drainages. Cluster analysis of Nei’s genetic distance did not show populations grouping according to drainages, although it did reveal alternative clusters. We found two distinct morphotypes that were supported as genetically distinct groups by the allozyme data. Genetic studies of vagile species in desert spring ecosystems can be used to reveal hydrologic connections and identify genetically unique sub-populations.

In a verbal model, Trivers and Willard proposed that, whenever there is sexual selection among males, natural selection should favor mothers that produce sons when in good condition but daughters when in poor condition. The predictions of this model have been the subject of recent debate. We present an explicit population genetic model for the evolution of a maternal-effect gene that biases offspring sex ratio. We show that, like local mate competition, sexual selection favors female-biased sex ratios whenever maternal condition affects the reproductive competitive ability of sons. However, Fisherian sex-ratio selection, which favors a balanced sex ratio, is an opposing force. We show that the evolution of maternal sex-ratio biasing by these opposing selection forces requires a positive covariance across environments between the sex-ratio bias toward sons (b) and the mating success of sons (r). This covariance alone is not a sufficient condition for the evolution of maternal sex-ratio biasing; it must be sufficiently positive to outweigh the opposing sex-ratio selection. To identify the necessary and sufficient conditions, we partition total evolutionary change into three components: (1) maternal sex-ratio bias, (2) sexual selection on sons, and (3) sex-ratio selection. Because the magnitude of the first component asymmetrically affects the strength of the second, biasing broods toward females in a poor environment evolves faster than the same degree of bias toward males in a good environment. Consequently, female-biased sex ratios, rather than male-biased sex ratios, are more likely to evolve. We discuss our findings in the context of the primary sex-ratio biases observed in strongly sexually selected species and indicate how this perspective can assist the experimental study of sex ratio evolution.

Methods for collecting mud shrimp are highly variable, and the efficiency of these techniques is poorly known. In October 1999, on a mud flat near Puerto Penasco, Sonora, Mexico, we compared the 'kiwi' method for extracting mud shrimp from their burrows, with a core sampling method designed to accomplish the same goal. The coring method involved pressing a 10 cm × 90 cm PVC pipe into the substratum, creating suction in the pipe, and withdrawing a core of mud and the shrimp within. The kiwi method required 8-10 researchers to march in a circle on the mud flat to gradually liquefy the mud, thereby forcing shrimp to the surface as the circle closed inward. Two transects of 30 cores conducted at high and at low tide yielded similar numbers of adult animals of both sexes. Previous results show that mud shrimp tend to aggregate within burrows when breeding. In this collection, the distribution of mud shrimp in cores did not differ from a Poisson distribution, suggesting that these shrimp were non-reproductive. ...

We present evidence that the heritable genetic variation within individual species, especially dominant and keystone species, has community and ecosystem conse- quences. These consequences represent extended phenotypes, i.e., the effects of genes at levels higher than the population. Using diverse examples from microbes to vertebrates, we demonstrate that the extended phenotype can be traced from the individuals possessing the trait, to the community, and to ecosystem processes such as leaf litter decomposition and N mineralization. In our development of a community genetics perspective, we focus on intraspecific genetic variation because the extended phenotypes of these genes can be passed from one generation to the next, which provides a mechanism for heritability. In support of this view, common-garden experiments using synthetic crosses of a dominant tree show that their progeny tend to support arthropod communities that resemble those of their parents. We also argue that the combined interactions of extended phenotypes con- tribute to the among-community variance in the traits of individuals within communities. The genetic factors underlying this among-community variance in trait expression, partic- ularly those involving genetic interactions among species, constitute community heritability. These findings have diverse implications. (1) They provide a genetic framework for un- derstanding community structure and ecosystem processes. The effects of extended phe- notypes at these higher levels need not be diffuse; they may be direct or may act in relatively few steps, which enhances our ability to detect and predict their effects. (2) From a con- servation perspective, we introduce the concept of the minimum viable interacting popu- lation (MVIP), which represents the size of a population needed to maintain genetic diversity at levels required by other interacting species in the community. (3) Genotype 3 environ- ment interactions in dominant and keystone species can shift extended phenotypes to have unexpected consequences at community and ecosystem levels, an issue that is especially important as it relates to global change. (4) Documenting community heritability justifies a community genetics perspective and is an essential first step in demonstrating community evolution. (5) Community genetics requires and promotes an integrative approach, from genes to ecosystems, that is necessary for the marriage of ecology and genetics. Few studies span from genes to ecosystems, but such integration is probably essential for understanding the natural world.

Two theoretical frameworks guide research on multiple male phenotypes within natural populations. Each scheme recognizes that male polymorphisms vary in the degree to which genotype and environment influence trait expression. Consensus remains elusive, however, on whether average fitnesses must be equivalent and whether genetic differences need exist for polymorphism to persist over time. Schradin and Lindholm address these hypotheses in African four-striped mice with detailed parentage and body size data. Their results and interpretation call each framework's predictions into question, but reveal a common truth. Debate might be resolved if researchers agreed on which parameters to measure and compare.