The Influence of Dispersal and Diet Breadth on Patterns of Genetic Isolation by Distance in Phytophagous Insects

To determine the effects of dispersal ability and diet breadth on population-genetic structure, we reviewed the allozyme literature and estimated genetic isolation by distance (IBD) for 43 species/host races of phytophagous insects. Subsequently, we tested two opposing hypotheses regarding the influence of dispersal ability on IBD: that IBD slopes do not vary with mobility, but that intercepts increase with mobility, and, alternatively, that IBD slopes vary with dispersal ability. We found that from tens of kilometers to more than 1,000 km, IBD is weak in sedentary and highly mobile species but pronounced in moderately mobile species. We attribute the weak IBD in strong dispersers to the homogenizing effects of gene flow, whereas in sedentary species, limited gene flow allows nearly all populations to diverge. In intermediate dispersers, genetic homogeneity is achieved at small spatial scales, but limited dispersal promotes genetic divergence over long distances. We also tested the hypothesis that IBD increases with decreasing diet breadth. We discovered no such pattern, casting doubt on the supposition that specialization promotes speciation by influencing population-genetic subdivision. Finally, we found that the number of populations is a more important consideration than the number of polymorphic loci in studies of IBD

Direct filtration using coarse media and dual media filters in series

A filtration system consisting of coarse media filters in series with standard dual media filters was proposed and evaluated as an alternative for direct filtration of high turbidity waters. The coarse media filters utilized a 3.08 mm uniform sand with a media depth of 1.32 m;The study was divided into three phases. The first phase involved the use of jar tests to determine optimum coagulants and approximate dosages for direct filtration. The jar test results were used in the second phase of study which involved laboratory pilot filter studies. The optimum alum dosage in the pilot plant studies was less than that determined in the jar test. Successful filter runs were possible with influent solids concentrations of 100 mg/l of kaolin or less. Headloss development (3 m maximum) limited the filter run lengths at kaolin concentrations of 25 and 50 mg/l. With 100 mg/l of kaolin, filter runs were terminated due to turbidity breakthrough. The third phase of the study involved a pilot plant field investigation using Des Moines river water as the raw water source. The chemical coagulants used in the laboratory did not produce acceptable results during the field study. The difference in chemical requirements was attributed to the nature of the raw water turbidity. The turbidity in the Des Moines river water was attributed mainly to the presence of diatoms;Particle deposition in the coarse media filters was recorded photographically during the laboratory filter runs. The particles were deposited in the filters by first coating the media. Next, the particles filled the media void spaces. The filling of the void spaces resulted in the development of narrow channels through which the suspension was transferred deeper into the filter;Two mathematical models were applied to the laboratory pilot filter data. Both models agreed reasonably well with the experimental data and were subsequently used to develop design curves. Design considerations for both small and large system applications were evaluated;The results of the study indicated that coarse media filters in series with standard dual media filters can be used successfully in extending the process of direct filtration to waters of higher turbidity than presently acceptable

Feeding-Induced Changes in Plant Quality Mediate Interspecific Competition Between Sap-Feeding Herbivores

Feeding-induced plant resistance is a well-documented phenomenon for leaf-chewing insects. Furthermore, feeding-induced resistance provides the mechanistic basis for many cases of delayed interspecific competition, whereby previous feeding by one species diminishes the performance of other herbivores which attack the same plant later in the season. This phenomenon, however, has been very poorly investigated for sap-feeding insects. The results we present here for salt marsh-inhabiting planthoppers (Prokelisia dolus and P. marginata) provide one of the few known examples of delayed, plant-mediated interspecific competition between two sap-feeding insects. Three lines of experimental evidence from the laboratory, field cages, and open field plots provide support for the detrimental effects of previous feeding by one planthopper species on the subsequent survival and performance of the other. Laboratory experiments showed that prior feeding on cordgrass by one congener resulted in reduced performance of the other in the following generation. However, the effect was asymmetric. Prior feeding by P. dolus resulted in prolonged development and reduced body size (a correlate of fecundity) in P. marginata, whereas only development was protracted in P. dolus when plants were previously exposed to P. marginata. Consequently, P. dolus appears to be the superior competitor in the context of delayed, plant-mediated interactions. The negative effects of previous feeding by P. dolus on the development time, body size, and survival of P. marginata obtained in the laboratory were confirmed both in cages and on cage-free islets of cordgrass in the field. Feeding-induced reductions in host-plant quality by P. dolus may provide additional impetus for P. marginata to migrate from shared habitats on the high marsh to nutritionally superior plants in the low marsh rarely occupied by P. dolus. The mechanism underlying the delayed competitive effects between Prokelisia planthoppers is most likely diminished plant nutrition, because feeding by P. dolus significantly reduces the concentration of essential amino acids in cordgrass. The asymmetry of plant-mediated competition between the Prokelisia species may be due to the ability of P. dolus to better tolerate feeding-depleted levels of plant nitrogen via compensatory feeding. Even though these two planthoppers do not suffer significant fitness reductions during contemporaneous interactions, they compete severely in the context of feeding-induced plant resistance which is expressed later in the season. This result, coupled with the fact that most studies of interspecific interaction between herbivorous insects are contemporaneous, indicates that interspecific competition may be profoundly underestimated as a structuring force in phytophagous insect communities

Host Plant Phenology and Dispersal by a Montane Butterfly: Causes and Consequences of Uphill Movement

In the Wenatchee Mountains of central Washington State, populations of the lycaenid butterfly Euphilotes enoptes occur patchily with their sole host plant, Eriogonum compositum (Polygonaceae). Nearly all courtship and adult feeding occur on the inflorescences of this long-lived perennial. Furthermore, because females oviposit on inflorescences and larvae feed only on flowers and developing seeds, the window of opportunity for exploiting this resource is narrow. I demonstrated that inflorescence phenology varied according to the aspect and elevation of plant patches, and butterflies were most likely to occur in patches nearing full bloom. A mark–release–recapture study revealed that individual butterflies can disperse far enough to sample from a wide range of patch phenologies, and the movements of females are apparently influenced by those phenologies. In particular, an uphill bias in female movement appeared to be the result of greater emigration from senescing low-elevation patches than from phenologically delayed, high-elevation patches. Males showed no such uphill movement. The apparent result of the uphill movement by females was that larval densities increased fourfold over a 300-m elevational gradient in each of two years. Differential mortality is an unlikely explanation for this variation in larval density, as both parasitism rates and frequency of visitation by ants, which tend larvae, did not vary with elevation. To demonstrate the influence of host plant phenology on adult behavior, I manipulated flowering phenology in an artificially established grid of small host plant patches and observed the encounters of adult butterflies with those patches. Inflorescences in patches that were nearing full bloom received disproportionately many visits by females, indicating that patches in this phenological state were more attractive to females. Small sample sizes prevented me from determining if this result was due to increased entry into those patches by females, an increased number of inflorescence visitations per female, or both. The frequency with which inflorescences in the different patches were visited by males did not vary with patch phenology. These results suggest that the uphill bias in dispersal by females is indeed a response to host plant phenology. Thus, in this system, among-patch variation in host plant phenology apparently influences the relative densities of larvae in different host plant patches by altering patterns of dispersal among those patches

Behavioral, Molecular, and Morphological Evidence for a Hybrid Zone between Chrysochus auratus and C. cobaltinus (Coleoptera: Chrysomelidae)

In this article, we describe a hybrid zone between the chrysomelid beetles, Chrysochus auratus (F.), andC. cobaltinus LeConte, which have historically been considered as having allopatric distributions. By combining field studies with surveys of museum specimens, we documented that in western North America there are two regions in which these beetles are sympatric, and four additional regions in which populations of the two species are Washington, we found an ≈25 km wide area of sympatry in which the two species freely interbreed. Morphological and allozyme differences between the species allowed us to demonstrate that individuals with intermediate coloration in this area are indeed hybrids; all 22 putative hybrids we assayed for allozyme variation were heterozygous at each of three species-specific loci. Museum specimens revealed that the two species have been hybridizing in this region at least since 1952. Within the hybrid zone, ≈10-15% of the beetles is apparently F1 hybrids. At one focal site, 22.9% of all matings involved heterospecific pairs and 20.8% of all matings involved at least one hybrid individual. Although we found no molecular evidence of introgression between the two species, morphometric results and preliminary ecological data suggest possible past introgression or weak ongoing introgression. We discuss the implications of our findings for the specific status of these two species. This system appears well suited to provide answers to long-standing questions concerning the evolution of premating barriers between hybridizing species. In addition, hybridization between these two beetle species with differing host ranges will allow us to test the hypothesis that ecologically significant traits such as diet breadth can be gained via introgression

First report of Oecophora bractella (L.) (Lepidoptera: Oecophoridae) in North America

The first report of Oecophora bractella (L.) from North America is given, based on collection records from Washington and British Columbia. This species is found throughout Europe, but is generally rare to uncommon through most of its range. Larvae occur on rotting wood in association with certain fungi and are not expected to reach pest status. Adults were captured at nine locations, with the first record from Seattle, WA in 1998. This first record and some of the subsequent records are from sites immediately adjacent to or associated with international shipping routes. Other sites with O. bractella are from residential areas, two of which have established breeding populations. The introduction of this moth is surprising, particularly given the low interception rate at United States ports and its specialized feeding niche. A diagnosis and photographs of adults are provided to enable North American researchers to identify this species and to monitor its spread

Sawflies (Hymenoptera, Symphyta) Newly Recorded from Washington State

Examination of museum specimens, unpublished collection data, and field surveys conducted between 2010 and 2014 resulted in records for 22 species of sawflies new to Washington State, seven of which are likely to be pest problems in ornamental landscapes. These data highlight the continued range expansion of exotic species across North America. These new records also indicate that our collective knowledge of Pacific Northwest arthropod biodiversity and biogeography is underdeveloped, even for a relatively well known and species-poor group of insects. Notable gaps in the knowledge of Washington State’s Symphyta remain for the Olympic Peninsula, the Cascade Mountain Range, and the arid interior of the state. Washington’s shrub-steppe appears to be particularly poorly surveyed for sawflies

Sawflies (Hymenoptera, Symphyta) Newly Recorded from Washington State

Examination of museum specimens, unpublished collection data, and field surveys conducted between 2010 and 2014 resulted in records for 22 species of sawflies new to Washington State, seven of which are likely to be pest problems in ornamental landscapes. These data highlight the continued range expansion of exotic species across North America. These new records also indicate that our collective knowledge of Pacific Northwest arthropod biodiversity and biogeography is underdeveloped, even for a relatively well known and species-poor group of insects. Notable gaps in the knowledge of Washington State’s Symphyta remain for the Olympic Peninsula, the Cascade Mountain Range, and the arid interior of the state. Washington’s shrub-steppe appears to be particularly poorly surveyed for sawflies

Habitat Persistence Underlies Intraspecific Variation in the Dispersal Strategies of Planthoppers

Dispersal is considered a vital life history characteristic for insects exploiting temporary habitats, and life history theorists have often hypothesized an inverse relationship between dispersal capability and habitat persistence. Most often, this hypothesis has been tested using interspecific comparisons of dispersal capability and qualitative estimates of habitat persistence. Consequently, most assessments have failed to control for possible phylogenetic nonindependence and they also lack quantitative rigor. We capitalized on existing intraspecific variation on the dispersal capability of Prokelisia planthoppers to examine the relationship between habitat persistence and dispersal, thereby minimizing possible phylogenetic effects. Two congeneric species (Prokelisia marginata and P. dolus) occur in the intertidal marshes of North America, where they feed exclusively on cordgrasses (Spartina). Because these planthoppers exhibit wing dimorphism, flight-capable adults (macropters with fully developed wings). Thus, dispersal capability can be readily estimated by the percentage of macropters in a population. At a regional spatial scale, we found a highly significant negative relationship between dispersal capability (present macroptery) and habitat persistence. In this system, habitat persistence is influenced by a combination of marsh elevation, winter severity, and tidal range, which interact to determine the ability of planthoppers to endure through winter in their primary habitat for development. P. marginata develops primarily in low-marsh habitats during summer, habitats that can be subjected to pronounced winter disturbance due to ice scouring and/or extensive tidal inundation. Levels of winter disturbance of the low marsh are extreme along the Atlantic coast, intermediate along the Pacific, and low along the Gulf. Both the failure of P. marginata populations to remain through winter in the habitat, and the dispersal ability of these populations (92%, 29%, and 17% macroptery, respectively), are correlated with levels of disturbance. Thus, in regions where winter disturbance is high, levels of dispersal are correspondingly high to allow for recolonization of extirpated habitats from overwintering sites on the high marsh. Unlike P. marginata, P. dolus develops primarily in high-marsh habitats, which are much less disturbed on all coasts during winter. Consequently, this species remains year-round in its primary habitat for development, and most populations exhibit relatively low levels of macroptery (\u3c10%). When raised under common garden conditions, many more macropters of both species were produced from Atlantic compared to Gulf populations. Thus the proportion of macropters produced from the populations used in this experiment paralleled the incidence of macroptery measured in the field, providing evidence that the geographic variation in dispersal capability in both species has in part a genetic basis. The results of this study provide strong intraspecific evidence for an inverse relationship between the dispersal capability of insects and the persistence of their habitats

The Effects of Heterospecific Mating Frequency on the Strength of Cryptic Reproductive Barriers

Heterospecific mating frequency is critical to hybrid zone dynamics and can directly impact the strength of reproductive barriers and patterns of introgression. The effectiveness of post-mating prezygotic (PMPZ) reproductive barriers, which include reduced fecundity via heterospecific matings and conspecific sperm precedence, may depend on the number, identity and order of mates. Studies of PMPZ barriers suggest that they may be important in many systems, but whether these barriers are effective at realistic heterospecific mating frequencies has not been tested. Here, we evaluate the strength of cryptic reproductive isolation in two leaf beetles (Chrysochus auratus and C. cobaltinus) in the context of a range of heterospecific mating frequencies observed in natural populations. We found both species benefited from multiple matings, but the benefits were greater in C. cobaltinus and extended to heterospecific matings. We found that PMPZ barriers greatly limited hybrid production by C. auratus females with moderate heterospecific mating frequencies, but that their effectiveness diminished at higher heterospecific mating frequencies. In contrast, there was no evidence for PMPZ barriers in C. cobaltinus females at any heterospecific mating frequency. We show that integrating realistic estimates of cryptic isolation with information on relative abundance and heterospecific mating frequency in the field substantially improves our understanding of the strong directional bias in F1 production previously documented in the Chrysochus hybrid zone. Our results demonstrate that heterospecific mating frequency is critical to understanding the impact of cryptic post-copulatory barriers on hybrid zone structure and dynamics, and that future studies of such barriers should incorporate field-relevant heterospecific mating frequencies