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In tropical rain forests, harvestmen assemblages are extremely diverse, with richness often exceeding 25 species. In the neotropics, there are published accounts of harvestmen faunas in South America rainforests (especially Amazonia), but relatively little is known about the community ecology of harvestmen in tropical forests of Central America. In this paper, we provide the first insights into the diverse assemblage of harvestmen inhabiting a wet forest at La Selva Biological Station, Costa Rica. Over five field seasons, we recorded 38 species. During our 2009 field season, we examined variation in species abundance, richness, and composition between adjacent successional forests (young secondary, mature secondary, and primary forests) as well as between distinct habitats (ground/litter layer and shrub/tree layer). Based on night samples (but not day), our results indicate that there are only minor differences in species composition and relative abundance between the forest ages, but no differences in richness. The ground/litter layer and shrub/tree layer habitats differed markedly in species composition, species richness, and relative abundance of several species. Our analysis of covariance supports the hypothesis that leg length is related to climbing behavior for several species belonging to Eupnoi and Laniatores.

Th e evolution of increased competitive ability (EICA) hypothesis provides a compelling explanation for the success of invasive species. It contends that because alien plants have escaped their coevolved natural enemies, selection pressures favor a diversion of resources from herbivore defense to traits that confer increased competitive ability. Here, we provide evidence for EICA in the noxious grassland invader Lespedeza cuneata , by comparing the ancestral genotype introduced to North America in 1930 with modern-day invasive (North American) and native (Japanese) genotypes. We found that the invasive genotype was a better competitor than either the native or the ancestral genotype. Further, the invasive genotype exhibited greater induced resistance but lower constitutive resistance than the ancestral and native genotypes. Our results suggest that selection has played a pivotal role in shaping this invasive plant species into a more aggressive, but less constitutively defended competitor. Th e rapid population growth and high densities reached by invasive species provides scientists with many opportunities to investigate the mechanisms that regulate the abundance and distribution of organisms. Such research is a conservation priority since invasive species cause signifi cant environmental and economic damage, and pinpointing the mechanisms that allow invasive species to achieve community dominance

Although development of critical thinking skills has emerged as an important issue in undergraduate education, implementation of pedagogies targeting these skills across different science, technology, engineering, and mathematics disciplines has proved challenging. Our goal was to assess the impact of targeted interventions in 1) an introductory cell and molecular biology course, 2) an intermediate-level evolutionary ecology course, and 3) an upper-level biochemistry course. Each instructor used Web-based videos to flip some aspect of the course in order to implement active-learning exercises during class meetings. Activities included process-oriented guided-inquiry learning, model building, case studies, clicker-based think–pair–share strategies, and targeted critical thinking exercises. The proportion of time spent in active-learning activities relative to lecture varied among the courses, with increased active learning in intermediate/upper-level courses. Critical thinking was assessed via a pre/posttest design using the Critical Thinking Assessment Test. Students also assessed their own learning through a self-reported survey. Students in flipped courses exhibited gains in critical thinking, with the largest objective gains in intermediate and upper-level courses. Results from this study suggest that implementing active-learning strategies in the flipped classroom may benefit critical thinking and provide initial evidence suggesting that underrepresented and first-year students may experience a greater benefit.

The effects of early-season herbivory and subsequent induced plant responses have the potential to affect the diversity of herbivorous insect communities. We investigated the seasonal development of the herbivore fauna on common milkweed (Asclepias syriaca) to understand the effect of early-season herbivory by different species on insect growth, natural colonization, and community composition. First, we showed that damage by an early-season stem-feeding weevil (Rhyssomatus lineaticollis) reduced growth of monarch larvae (Danaus plexippus) and leaf beetle larvae (Labidomera clivicollis), suggesting that plant quality is reduced by weevil damage. To better understand the potential for initial herbivore damage to affect subsequent colonization by herbivores in the field, we compared undamaged controls to plants experimentally damaged with one of three herbivores: weevils, monarchs, or leaf beetles. We counted seven species of naturally colonizing herbivores on all plants for the next two months to assess colonization, damage, and insect community richness. Our results showed that initial herbivory by different species altered host plant use by herbivores in two years of experiments. Similarly, induced resistance and suscep- tibility occurred in both years, but due to different initial damaging species on individual plants. Treatment effects also scaled up to alter herbivore community richness. Initial treatments varied in their persistence through the season. For example, in 2001, the influence of initial monarch damage dissipated due to subsequent damage by colonizing herbivores, but the impacts of initial weevil treatment were unaffected. This result suggests that, al- though induced responses to weevil feeding persisted through the season, monarch herbivory was more likely to affect the herbivore community via a cascade of indirect effects. In 2002, plant and insect responses were more specific, depending on the identity of both initial and colonizing herbivore species. Despite year-to-year variation, considerable con- sistency in many responses to our treatments indicates that the identity of the initially colonizing herbivore can affect subsequent plant use and community structure. Given the preponderance of influential early-season herbivores, the effects of induced plant responses similar to those presented here may be widespread and may strongly contribute to the structure of phytophagous insect communities.

Induced plant responses to herbivory appear to be universal, yet the degree to which they are specific to sets of herbivores is poorly understood. The generalist/specialist hypothesis predicts that generalist herbivores are more often negatively affected by host plant defenses, wheras specialists may be either unaffected by or attracted to these same “plant defenses”. Therefore, specialists should be less predictable than generalists in their responses to induced plant resistance traits. To better understand the variation in plant responses to herbivore attack, and the impacts these responses have on specialist herbivores, we conducted a series of experiments examining pairwise interactinos between two specialaist herbivores of the common milkweed (Asclepias syriaca). We damaged plants mechnically, with swamp milkweed beetles (Labidomera clivicollis), or with monarchs (Danaus plexippus), and then asessed specificity of elicitation, both by measuring a putative defensive trait (latex volume) and by challenging plants with insects of both species in bioasays. Latex production increased by 34% and 13% following beetle and monarch herbivory, respectively, but only beetles significantly elevated latex production compared to undamaged controls. While beetle growth was negatively affected by latex across all experiments, beetles were not affected by previous damage caused by conspecifies or by monarchs. In contrast, monarchs feeding on previously damaged plants were 20% smaller, and their response was the same on plants damaged mechnically or by either herbivore. Therefore, these specialist herbivores exhibit both specificity of elicitation in plant responses and specificity of effects in response to prior damage.

Summary 1 We studied the genetic and environmental determinants of attack by the specialist stem-attacking weevil, Rhyssomatus lineaticollis on Asclepias syriaca. 2 In natural populations, the extent of stem damage and oviposition were positively correlated with stem width, but not stem height. We hypothesized that both genotypic and environmental factors influencing stem morphology would affect attack by weevils. 3 In a common garden study with 21 full-sib families of milkweed, both phenotypic and genetic correlations indicated that weevils impose more damage and lay more eggs on thicker stemmed plants. 4 Of three other putative resistance traits, only latex production showed a negative genetic correlation with weevil attack. 5 When neighbouring grasses were clipped to reduce light competition, focal milkweed plants received up to 2.6 times the photosynthetically active radiation and 1.6 times the red to far red ratio of light compared with plants with intact grass neighbours. Focal milkweed plants were therefore released from the classic neighbour avoidance response and had 20% shorter internode lengths, were 30% shorter, and had 90% thicker stems compared with controls. 6 Clipping of grass neighbours resulted in nearly 2.7 times the damage and oviposition by stem weevils, thus supporting the hypothesis of an environmental or trait-mediated indirect influence on resistance. 7 Although attack of plants by weevils strongly increases the probability of stem mortality, thicker stems experience lower mortality, thus counteracting the selective impact of weevil-induced plant mortality. 8 The determinants of attack on milkweeds include both genetic variation for stem thickness and an indirect environmental influence of plant neighbours. If milkweeds and weevils are coevolving, the interaction is diffuse because the ecological neighbourhood is likely to modify the patterns of reciprocal natural selection.

Summary 1 Salinization is a growing environmental stress in wetland ecosystems world-wide. Several models have been proposed that predict clonal plant responses to stress, including that environmental stress stimulates sexual reproduction. 2 We conducted a common-garden experiment to investigate the effects of salinity on 10 natural populations of Iris hexagona, a clonal perennial endemic to freshwater and brackish wetlands of the North American Gulf Coast. 3 Salinity reduced vegetative growth but either increased or had neutral effects on sexual reproduction, consistent with the clonal stress hypothesis. Salinity of 4 µg g−1 more than doubled the number of seeds produced compared with freshwater controls, but flower number and seed mass were unaffected. 4 Salinity reduced total below-ground mass by nearly 50% compared with controls, with no significant change in rhizome numbers. 5 Plants from 10 randomly selected I. hexagona populations differed dramatically in growth and reproduction, independent of salinity. Total biomass that accumulated over the 20-month experiment ranged across all treatments from 52 to 892 g, and flower numbers varied from 2.3 to 11.3 per replicate. 6 Populations did not respond differently to salinity, except with respect to above- : below-ground ratios, thus providing no conclusive evidence for local adaptation to salinity stress. 7 Our results concur with published models of plant reproductive strategies in variable environments, in that environmental stress stimulated sexual reproduction at the expense of growth. However, these models do not predict the observed sharp decline in seed production at near lethal salinity levels.

Adults and larvae of the loberine erotylid beetle Loberus impressus (LeConte) were found associated with fungi growing on corolla and seed capsule tissue of the blue flag iris, Iris hexagona. We examined adult beetle specimens using light and scanning electron microscopy to determine if specialized structures (mycangia) may function in transporting fungi. Two pairs of deep pits on the ventral aspect of the gena between the eyes and the maxillae of both sexes contained fungal spores and hyphae, suggesting a possible role as mycangia in addition to their role as glandular outlets. Inoculation from the surface of cleaned beetle specimens produced colonies of Cladosporium and Fusarium. These genera are widespread, usually air-dispersed conidial fungi that sometimes are associated with insects.

Stilbosis quadricustatella leafminers are microlepidopteran specialists of sand-live oak (Quercus geminata). These tiny moths produce one generation per year and have a parasitic life-cycle and long larval stage that develops entirely within a single oak leaf. Differences in host-plant age, phenotype, and phenology generate a coarse-grained, spatially heterogeneous environment for the leafminer population. Previous reciprocal trans- fers of leafminer eggs among mature oaks revealed that S. quadricustatella are locally adapted to individual oak trees. In this paper we use genetic markers and an extinction- recolonization experiment to explore further variation in leafminer population structure. Allozyme loci indicate significant interdemic genetic structure among recent colonists of new host trees, which weakens in the 10th generation and disappears by the 40th generation. In contrast, adaptive demic structure is evident by the 10th generation and is strong in the 40th generation, despite the potential for substantial intertree dispersal. We propose that host heterogeneity combined with leafminer fidelity to natal trees promotes divergent se- lection and rapid demic evolution on individual oaks, despite potentially high gene flow between the leafminers inhabiting them.

The larva of Loberus impressus LeConte is described based on adult-associated and reared specimens collected from corollas and seed pods of Iris hexagona on the gulf coast of Louisiana. The larva of L. impressus is similar to larvae of Zavaljus brunneus (Gyllenhal) Hapalips prolixus Sharp, Pharaxonotha spp., and Bolerus angulosus (Arrow), the other described larvae within the Xenoscelinae. Larval characters typical for the Loberini that are found in L. impressus include arrangement of integumental granules into rows, frayed and aciculate setae borne on spiny tubercles, and urogomphi spiny or tuberculate and recurved. The tarsungular setae of L. impressus and B. angulosus are single, in contrast to the dual setae present in other described larvae of Languriidae. Larva and adults of L. impressus were common in dried corollas and seed pods of I. hexagona that had begun to split open or had been invaded by Crematogaster ants. Fusarium and Cladosporium fungi were potential sources of food for L. impressus and other fungivorous beetles in the iris seed pods and on corollas.

The adaptive deme formation (ADF) hypothesis pre- likely to become locally adapted than are dispersive in- dicts that herbivorous insects become locally adapted to their host sects; and haplodiploid insects are more likely to become plants over time. Since its inception, approximately 17 indepen- dent studies have tested ADF, and they are divided in support and locally adapted than are diplodiploid species. rejection of the hypothesis. This field of insect evolutionary ecol- Since its inception, the predictions of the ADF hypoth- ogy has a contentious history, and the contradictory studies ob- esis have been examined in approximately 17 indepen- scure our understanding of the general evolutionary importance of dent studies. Recent reviews of these studies (Mopper adaptive deme formation in phytophagous insects. We conducted 1996; Boecklen and Mopper 1998; Strauss and Karban a meta-analysis in an attempt to clarify this issue. Meta-analysis is 1998) conclude that they are about evenly divided in a statistical method for quantitatively comparing and synthesizing support and rejection of the ADF hypothesis. Addition- the results of different studies in a way that is more objective than a traditional literature review. Our analysis indicates that local ad- ally, Alstad (1998) recently retracted the experimental re- aptation is an important phenomenon in diverse insect systems. sults and deme formation hypothesis proposed in the Contrary to predictions of the original hypothesis, there was no original study by Edmunds and Alstad (1978). This field evidence that insect dispersal ability, and ostensibly gene flow, was of insect evolutionary ecology has been somewhat con- associated with local adaptive differentiation. There was some in- tentious, and combined with the inconsistent results of dication that breeding (parthenogenetic, haplodiploid, diplodi- the experimental field studies, a clear picture of the bio- ploid) and feeding (exophagous, endophagous) modes may influ- logical role of adaptive deme formation in the evolution ence the evolution of locally adapted demes. Our analysis supports the theory of adaptive deme formation and provides guidance for of phytophagous insect populations has been impossible future research directions. to obtain using traditional literature reviews. Meta-analysis offers a solution to this dilemma be

Saltwater intrusion into wetland ecosystems has destroyed or damaged many native plant populations. Iris hexagona is a salt-sensitive species that exhibits intraspecific variation in salinity tolerance. To investigate the effect of salinity on flowering, we exposed I. hexagona collected from natural populations to salt treatments in a common garden. Experimental salinity additions strongly delayed flowering phenology, but the effect was not apparent until the second year, when less than 4 g/L NaCl delayed flowering up to 3 d. In the field, soil salinity and flowering phenology varied substantially within I. hexagona populations. Iris flowers are receptive to pollinators for 2 d or less, therefore a 3-d delay could affect outcrossing dynamics, and ultimately, the evolutionary ecology of iris populations. Salinity also had a carryover effect; prior salinity exposure delayed flowering in irises that had been replanted in freshwater conditions for 6 mo. This is an important result because it suggests that episodic stress (such as tropical storms) can influence performance well after the stress has disappeared. Our research further underscores the importance of long-term studies because a 1-yr experiment would have failed to reveal the strong effects of salinity that emerged in the second year.

The growth rate (GR) hypothesis relates the evolution of plant defense to resource availability and predicts that plants that have evolved in abiotically stressful environments grow inherently more slowly and are more constitutively resistant to herbivory than plants from more productive habitats. Stress-adapted plants are also predicted to have reduced inducibility, but this prediction has not been previously tested. To evaluate this hypothesis, I compared the growth of nine species of herbaceous plants from Missouri glade habitats to congeners from more productive non-glade habitats. I also conducted bioassays using larvae of the generalist herbivore Spodoptera exigua to estimate constitutive and inducible resistance in these congeners. Glade congeners tended to grow more slowly and have higher constitutive resistance and lower inducibility than non-glade species. However, none of these comparisons was statistically significant due to the conflicting response of one congeneric pair (Salvia azurea and S. lyrata). Analyses without this genus were consistent with the GR hypothesis, as were analyses that categorized congeners by relative growth rate. These results highlight the complexity in searching for factors that determine plant growth rates and resistance traits across multiple genera and support the hypothesis that both constitutive and induced resistance may be influenced by selection on traits that alter plant growth rates. Future studies should attempt to determine whether variation in inducibility is better explained by habitat or relative plant growth rates.

Extinction of populations occurs naturally, but global extinction rates are accelerating, making understanding extinction a high priority for conservation. Extinction in experimental populations of brine shrimp ( Artemia franciscana ) was measured to assess hypothesized extinction processes. Greater initial population size, greater maximum population size supported by the environment, and lower variation in environmental conditions reduced the likelihood of extinction, as hypothesized. However, initial population size was less important, and maximum population size and environmental variation were more important than often hypothesized. Unexpectedly, deterministic oscillations in population size due to inherent nonlinear dynamics and overcrowding were as important or more important than hypothesized processes.