Spillover of Agriculturally Subsidized Predators as a Potential Threat to Native Insect Herbivores in Fragmented Landscapes

Abstract:  Habitat loss and fragmentation can have strong negative impacts on populations of some native species. Spillover of generalist natural enemies from the surrounding landscape matrix is one mechanism potentially generating such effects, yet this has been rarely studied in insects. We examined the influence of habitat conversion to agriculture on the abundance and potential effects of predatory coccinellid beetles on native insect herbivores within 12 grassland remnants in central Nebraska (U.S.A.). Results of sweep sampling revealed that coccinellids were three to six times more abundant at native grassland sites embedded within cropland-dominated landscapes compared with control sites in grassland-dominated landscapes over the 3 years of the study. Exclusion experiments further demonstrated that predation intensity was strongly related to coccinellid abundances across sites and that coccinellids can dramatically reduce densities of a native aphid herbivore. In contrast to studies of specialized insect parasitoids, which have generally found reduced enemy pressure in fragmented landscapes, our results suggest that native herbivores may in some cases experience increased consumer pressure in landscapes with increasing habitat loss because of spillover of generalist predators from surrounding cropland habitats.     

Impact of minimum winter temperatures on the population dynamics of Dendroctonus frontalis.

Predicting population dynamics is a fundamental problem in applied ecology. Temperature is a potential driver of short-term population dynamics, and temperature data are widely available, but we generally lack validated models to predict dynamics based upon temperatures. A generalized approach involves estimating the temperatures experienced by a population, characterizing the demographic consequences of physiological responses to temperature, and testing for predicted effects on abundance. We employed this approach to test whether minimum winter temperatures are a meaningful driver of pestilence from Dendroctonus frontalis (the southern pine beetle) across the southeastern United States. A distance-weighted interpolation model provided good, spatially explicit, predictions of minimum winter air temperatures (a putative driver of beetle survival). A Newtonian heat transfer model with empirical cooling constants indicated that beetles within host trees are buffered from the lowest air temperatures by approximately 1-4 degrees C (depending on tree diameter and duration of cold bout). The life stage structure of beetles in the most northerly outbreak in recent times (New Jersey) were dominated by prepupae, which were more cold tolerant (by >3 degrees C) than other life stages. Analyses of beetle abundance data from 1987 to 2005 showed that minimum winter air temperature only explained 1.5% of the variance in interannual growth rates of beetle populations, indicating that it is but a weak driver of population dynamics in the southeastern United States as a whole. However, average population growth rate matched theoretical predictions of a process-based model of winter mortality from low temperatures; apparently our knowledge of population effects from winter temperatures is satisfactory, and may help to predict dynamics of northern populations, even while adding little to population predictions in southern forests. Recent episodes of D. frontalis outbreaks in northern forests may have been allowed by a warming trend from 1960 to 2004 of 3.3 degrees C in minimum winter air temperatures in the southeastern United States. Studies that combine climatic analyses, physiological experiments, and spatially replicated time series of population abundance can improve population predictions, contribute to a synthesis of population and physiological ecology, and aid in assessing the ecological consequences of climatic trends.     

Antennal responses of four species of tree-killing bark beetles (Coleoptera: Scolytidae) to volatiles collected from beetles,and their host and nonhost conifers

Summary. Host selection in tree-killing bark beetles (Coleoptera: Scolytidae) is mediated by a complex of semiochemical cues. Using gas chromatographic-electroantennographic detection (GC-EAD) and GC-mass spectrometric analyses, we conducted a comparative study of the electrophysiological responses of four species of tree-killing bark beetles, the Douglas-fir beetle, Dendroctonus pseudotsugae, Hopkins, the mountain pine beetle, D. ponderosae Hopkins, the spruce beetle, D. rufipennis Kirby, and the western balsam bark beetle, Dryocoetes confusus Swaine, to volatiles captured by aeration of 1) bole and foliage of four sympatric species of conifers, Douglas-fir, Pseudotsuga menziesii (Mirb.) Franco, lodgepole pine, Pinus contorta var. latifolia Engelm., interior spruce, Picea engelmannii x glauca, and interior fir, Abies lasiocarpa x bifolia, and 2) con- and heterospecific beetles at three stages of attack. We identified 13 monoterpenes in the conifers and nine compounds in the volatiles of beetles that elicited antennal responses. There was no qualitative difference in the terpene constitution of the four species of conifers and very little difference across beetle species in their antennal response to compounds from conifers or beetles. The lack of species-specific major or minor components in conifers suggests that beetles would need to detect differences in the ratios of different compounds in conifers to discriminate among them. Attraction to hosts and avoidance of nonhost conifers may be accentuated by perception of compounds emitted by con- and heterospecific beetles, respectively. The 22 compounds identified are candidate semiochemicals with potential behavioural roles in host location and discrimination.     

Do birds and beetles show similar responses to urbanization?

To date, the vast majority of studies in urban areas have been carried out on birds, yet it is not known whether the responses of birds to urbanization are congruent with those of other taxa. In this paper, we compared the responses of breeding birds and carabid beetles to urbanization, specifically asking whether the emerging generalizations of the effects of extreme levels of urbanization on birds (declines in total species richness and the richness of specialist species, increases in total abundance and the abundances of native generalist and introduced species, and community simplification, including increasing similarity) could also be applied to ground beetles. We also directly tested for congruence between birds and ground beetles using correlations between variables describing bird and beetle community structure and correlations between bird and beetle distance matrices describing community dissimilarity between pairs of sampling locations. Breeding bird and carabid beetle community data were collected in Ottawa, Ontario, and Gatineau, Quebec, Canada, in two groups of sites: developed sites representing the predictor variable within-site housing density, and forested sites adjacent to development representing the predictor variable neighboring housing density (each site was 0.25 km2). Breeding birds and carabid beetles do not respond similarly to increasing within-site housing density but do exhibit some similar responses to increasing neighboring housing density. Birds displayed strong declines in diversity, compositional changes, and community simplification in response to increasing within-site housing density. Forest and introduced species of birds and beetles responded similarly to increasing housing density within a site, but responses of overall diversity and open-habitat species richness and patterns of community simplification differed between birds and beetles. Increasing neighboring housing density resulted in increases in the abundances of introduced birds and introduced beetles and similar patterns of community simplification in both taxa. To better understand and mitigate the effects of urbanization on biodiversity, we suggest that, in addition to the responses of birds, future research should focus on the responses of other taxa in the urban matrix.     

Effects of Forest Fragmentation on a Dung Beetle Community in French Guiana

Abstract:  Fragmentation is the most common disturbance induced by humans in tropical forests. Some insect groups are particularly suitable for studying the effects of fragmentation on animal communities because they are taxonomically and ecologically homogenous. We investigated the effects of forest fragmentation on a dung beetle species community in the forest archipelago created in 1994–1995 by the dam of Petit Saut, French Guiana. We set and baited an equal number of pitfall traps for dung beetles on three mainland sites and seven island sites. The sites ranged from 1.1 to 38 ha. In 250 trap days, we captured 50 species in 19 genera. Diversity indices were high (2.18–4.06). The lowest diversity was on the small islands and one mainland site. Species richness and abundance were positively related to fragment area but not to distance from mainland or distance to the larger island. The islands had lower species richness and population than mainland forest, but rarefied species richness was relatively invariant across sites. There was a marked change in species composition with decreasing fragment that was not caused by the presence of a common fauna of disturbed-area species on islands. Small islands differed from larger islands, which did not differ significantly from mainland sites. Partial correlation analyses suggested that species richness and abundance of dung beetle species were positively related to the number of species of nonflying mammals and the density index of howler monkeys ( Alouatta seniculus ), two parameters positively related to fragment area.     

Host-plant-mediated competition via induced resistance: interactions between pest herbivores on potatoes.

Plant-mediated competition among insect herbivores occurs when one species induces changes in plant chemistry, nutrition, or morphology that render plants resistant to attack by others. We explored plant-mediated interspecific interactions between the potato leafhopper (Empoasca fabae) and the Colorado potato beetle (Leptinotarsa decemlineata), two important pests on potatoes. Leafhoppers colonize fields in advance of beetles, and thus the possibility exists that previous feeding by leafhoppers induces changes in potato plants that have adverse consequences for beetles. The consequences of leafhopper-induced resistance for beetle performance were studied in the greenhouse, field cages, and in large open-field plots. Potato plants were exposed to four densities of leafhoppers (none, low, moderate, and high), and visible feeding symptoms were measured as percentage leaf curling, chlorosis, and necrosis. The oviposition preference, performance, and survivorship of Colorado potato beetles were then measured on the four categories of induced plants in field-cage and greenhouse settings. In open field plots, survival on the four categories of induced plants was determined by placing cohorts of beetle adults onto plants and measuring the densities of resulting eggs, larvae, and emerging Fl adults. Leafhopper-induced symptoms on potato plants were density dependent, with the percentage of curled, chlorotic, and necrotic leaves increasing with leafhopper density. Previous feeding by leafhoppers adversely affected oviposition and larval performance of beetles. Fewer egg masses were deposited on plants that incurred high levels of leafhopper feeding. Similarly, larval development was delayed and emerging adult beetles weighed less when fed induced foliage from the high leafhopper-density treatment. Beetles survived less well in the field on plants experiencing moderate and high levels of leafhopper feeding as evidenced by lower densities of eggs, larvae, and emerging F1 adults. Overall, leafhoppers and beetles competed through feeding-induced changes in plant quality. Notably, the asymmetric interaction took place at a large spatial scale in open field plots and had negative consequences that persisted to the next beetle generation. Ultimately, to establish an effective management strategy for crop pests such as leafhoppers, it is essential to consider the positive indirect effects of induced resistance along with the negative direct effects on crop yield.     

Plague bacterium as a transformer species in prairie dogs and the grasslands of western North America

Invasive transformer species change the character, condition, form, or nature of ecosystems and deserve considerable attention from conservation scientists. We applied the transformer species concept to the plague bacterium Yersinia pestis in western North America, where the pathogen was introduced around 1900. Y. pestis transforms grassland ecosystems by severely depleting the abundance of prairie dogs (Cynomys spp.) and thereby causing declines in native species abundance and diversity, including threatened and endangered species; altering food web connections; altering the import and export of nutrients; causing a loss of ecosystem resilience to encroaching invasive plants; and modifying prairie dog burrows. Y. pestis poses an important challenge to conservation biologists because it causes trophic‐level perturbations that affect the stability of ecosystems. Unfortunately, understanding of the effects of Y. pestis on ecosystems is rudimentary, highlighting an acute need for continued research.     

Predicting Plant Extinction Based on Species-Area Curves in Prairie Fragments with High Beta Richness

Abstract:  Species-area relationships and island biogeography theory are commonly used to predict how species richness will decline with fragmentation. There are a variety of largely untested assumptions in these approaches, including the assumptions that populations are distributed uniformly before fragmentation, and that local extinctions are due to effects of small population sizes. If populations are not distributed uniformly, then populations can be abundant locally but rare globally. This would cause extinction rates to be smaller than predicted. We tested extinction theory by developing estimates of the number of plant species that should be present in small tallgrass prairie fragments and then testing the uniformity assumption by partitioning species richness into α (within site) and β (among site) components in Iowa prairies. Many more native prairie plant species were present in surveys of prairie fragments (491) than was predicted based on theory (27–207). A large proportion (75%) of the total species richness was β richness. We suggest that the high proportion of β richness was responsible for the shallow species-area slopes and the lower than expected number of species losses and that a better understanding of what determines β diversity will improve predictions of fragmentation effects on richness of plants. We also suggest that plants in prairie remnants may be best conserved by protecting different prairie types rather than by protecting a few large areas containing a single prairie type.     

New repellent semiochemicals for three species of <Emphasis Type="Italic">Dendroctonus</Emphasis> (Coleoptera: Scolytidae)

Summary. Nine compounds identified from captured volatiles of the Douglas-fir beetle, Dendroctonus pseudotsugae, the mountain pine beetle, D. ponderosae, and the spruce beetle, D. rufipennis, that elicited antennal responses in males and females of one or more of these species were tested in the field to determine behavioural activity. 1-Octen-3-ol, found in the volatiles of females of all three species decreased the response of male and female coastal and male interior D. pseudotsugae and both sexes of D. ponderosae to their aggregation pheromones. Acetophenone, identified in the volatiles of females of all three species, significantly decreased the response of interior female D. pseudotsugae. trans-Verbenol, a potent aggregation pheromone of D. ponderosae, decreased the response of both sexes of D. pseudotsugae, while 3-methyl-2-cyclohexen-1- one (MCH), the antiaggregation pheromone of D. pseudotsugae and D. rufipennis decreased the response of both sexes of D. ponderosae. While it has been demonstrated that semiochemical mediated interspecific communication occurs among bark beetles infesting the same host, this study demonstrates that beetles can perceive signals emitted by heterospecifics attacking nonhosts and can potentially use them to avoid attacking the wrong species of conifer.     

Effects of Knowledge of an Endangered Species on Recreationists’ Attitudes and Stated Behaviors and the Significance of Management Compliance for Ohlone Tiger Beetle Conservation

Recreation is a leading cause of species decline on public lands, yet sometimes it can be used as a tool for conservation. Engagement in recreational activities, such as hiking and biking, in endangered species habitats may even enhance public support for conservation efforts. We used the case of the endangered Ohlone tiger beetle (Cicindela ohlone) to investigate the effect of biking and hiking on the beetle's behavior and the role of recreationists’ knowledge of and attitudes toward Ohlone tiger beetle in conservation of the species. In Inclusion Area A on the University of California Santa Cruz (U.S.A.) campus, adult Ohlone tiger beetles mate and forage in areas with bare ground, particularly on recreational trails; however, recreation disrupts these activities. We tested the effect of recreation on Ohlone tiger beetles by observing beetle behavior on trails as people walked and road bikes at slow and fast speed and on trails with no recreation. We also surveyed recreationists to investigate how their knowledge of the beetle affected their attitudes toward conservation of the beetle and stated compliance with regulations aimed at beetle conservation. Fast cycling caused the beetles to fly off the trail more often and to fly farther than slow cycling or hiking. Slow cycling and hiking did not differ in their effect on the number of times and distance the beetles flew off the trail. Recreationists’ knowledge of the beetle led to increased stated compliance with regulations, and this stated compliance is likely to have tangible conservation outcomes for the beetle. Our results suggest management and education can mitigate the negative effect of recreation and promote conservation of endangered species. Efectos del Conocimiento de una Especie en Peligro sobre las Actitudes y Comportamientos Declarados de los Recreacionistas y el Significado del Manejo de la Conformidad para la Conservación del Escarabajo Tigre de Ohlone     

Evaluating the Predicted Local Extinction of a Once-Common Mouse

Abstract:  In an earlier paper ( Pergams & Nyberg 2001 ) we found that the proportion of the prairie deer mouse ( Peromyscus maniculatus bairdii ), among all local Peromyscus museum specimens collected in the Chicago region, had significantly declined over time. This proportion changed from about 50% before 1900 to <10% in the last 25 years. Based on this proportion a regression model predicted the local extinction of the prairie deer mouse in 2009. To evaluate that prediction, we estimated current deer mouse abundance by live trapping small mammals at 15 preserves in Cook and Lake counties, Illinois (USA) at which prairie deer mice had previously been caught or that still contained their preferred open habitat. In 1900 trap nights, 477 mammals were caught, including 251 white-footed mice ( P. leucopus ), but only one prairie deer mouse. The observed proportion of Peromyscus that were prairie deer mice, 0.4%, was even lower than the 4.5% predicted for 2000. Here we also introduce a simple, new community proportions model, which for any given geographic region compares the proportions of species recently caught with the proportions of species in museums. We compared proportions of seven species collected in Cook and Lake counties and examined by Hoffmeister (1989) with proportions of these species that we caught. Ten percent of the museum community was prairie deer mice, but only 0.2% of our catch was. The current local scarcity of the prairie deer mouse is consistent with the regression-based prediction of its eminent local extinction. More conservation attention should be paid to changes in relative abundance of once-common species.     

Pesticide treatments affect mountain pine beetle abundance and woodpecker foraging behavior.

In British Columbia, Canada, management efforts used to control mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreaks have included treatment of infested trees with an organic arsenic pesticide, monosodium methanearsonate (MSMA). Cumulative pesticide applications over a large geographic area have generated concerns about arsenic loading in the environment and potential toxicity to nontarget wildlife. We investigated woodpecker foraging patterns in infested stands with and without MSMA treatment using a combination of tree debarking indices, point count surveys, and radiotelemetry methods in addition to insect flight traps to measure mountain pine beetle emergence. Debarking indices indicated woodpecker foraging of MSMA-treated trees was significantly lower than nontreated trees in all sampling years. However, approximately 40% of MSMA trees had some evidence of foraging. Focal observations of foraging woodpeckers and point count surveys in MSMA treatment areas further confirmed that several species of woodpeckers regularly used MSMA stands during the breeding season. Radio-tagged Hairy (Picoides villosus) and Three-toed (Picoides dorsalis) Woodpeckers spent on average 13% and 23% (range 0-66%) of their time, respectively, in treated stands, despite the fact that these areas only comprised on average 1-2% of their core home range (1 km2). MSMA strongly reduced the emergence of several bark beetle (Coleoptera, Scolytidae) species including the mountain pine beetle, and there was a highly significant positive relationship between Dendroctonus beetle abundance and Three-toed Woodpecker abundance. This study identifies the potential negative impact that forest management practices using pesticides can have on woodpecker populations that depend on bark beetles and their host trees.     

Associations of Grassland Bird Communities with Black‐Tailed Prairie Dogs in the North American Great Plains

Colonial burrowing herbivores can modify vegetation structure, create belowground refugia, and generate landscape heterogeneity, thereby affecting the distribution and abundance of associated species. Black‐tailed prairie dogs (Cynomys ludovicianus) are such a species, and they may strongly affect the abundance and composition of grassland bird communities. We examined how prairie dog colonies in the North American Great Plains affect bird species and community composition. Areas occupied by prairie dogs, characterized by low percent cover of grass, high percent cover of bare soil, and low vegetation height and density, supported a breeding bird community that differed substantially from surrounding areas that lacked prairie dogs. Bird communities on colony sites had significantly greater densities of large‐bodied carnivores (Burrowing Owls [Athene cunicularia], Mountain Plovers, [Charadrius montanus], and Killdeer [Charadrius vociferus]) and omnivores consisting of Horned Larks (Eremophila alpestris) and McCown's Longspurs (Rhynchophanes mccownii) than bird communities off colony sites. Bird communities off colony sites were dominated by small‐bodied insectivorous sparrows (Ammodramus spp.) and omnivorous Lark Buntings (Calamospiza melanocorys), Vesper Sparrows (Pooecetes gramineus), and Lark Sparrows (Chondestes grammacus). Densities of 3 species of conservation concern and 1 game species were significantly higher on colony sites than off colony sites, and the strength of prairie dog effects was consistent across the northern Great Plains. Vegetation modification by prairie dogs sustains a diverse suite of bird species in these grasslands. Collectively, our findings and those from previous studies show that areas in the North American Great Plains with prairie dog colonies support higher densities of at least 9 vertebrate species than sites without colonies. Prairie dogs affect habitat for these species through multiple pathways, including creation of belowground refugia, supply of prey for specialized predators, modification of vegetation structure within colonies, and increased landscape heterogeneity. Asociaciones de Comunidades de Aves de Pastizales con Perros de la Pradera en la Gran Llanura de Norte América     

Simulation and analysis of outbreaks of bark beetle infestations and their management at the stand level

Outbreaks of bark beetles in forests can result in substantial economic losses. Understanding the factors that influence the development and spread of bark beetle outbreaks is crucial for forest management and for predicting outbreak risks, especially with the expected global warming. Although much research has been done on the ecology and phenology of bark beetles, the complex interplay between beetles, host trees, beetle antagonists and forest management makes predicting beetle population development especially difficult. Using the recent infestations of the European Spruce Bark Beetle (Ips typographus L. Col. Scol.) in the Bavarian Forest National Park (Germany) as a case study, we developed a spatially explicit agent-based simulation model (SAMBIA) that takes into account individual trees and beetles. This model primarily provides a tool for analysing and understanding the spatial and temporal aspects of bark beetles outbreaks at the stand scale. Furthermore, the model should allow an estimation of the effectiveness of concurrent impacts of both antagonists and management to confine outbreak dynamics in practice. We also used the model to predict outbreak probabilities in various settings. The simulation results indicated a distinct threshold behaviour of the system in response to pressure by antagonists or management of the bark beetle population. Despite the different scenarios considered, we were able to extract from the simulations a simple rule of thumb for the successful control of an outbreak: if roughly 80% of individual beetles are killed by antagonists or foresters, outbreaks will rarely take place. Our model allows the core dynamics of this complex system to be reduced to this inherent common denominator.     

Long-term reactions of plants and macroinvertebrates to extreme floods in floodplain grasslands

Extreme summertime flood events are expected to become more frequent in European rivers due to climate change. In temperate areas, where winter floods are common, extreme floods occurring in summer, a period of high physiological activity, may seriously impact floodplain ecosystems. Here we report on the effects of the 2002 extreme summer flood on flora and fauna of the riverine grasslands of the Middle Elbe (Germany), comparing pre- and post-flooding data collected by identical methods. Plants, mollusks, and carabid beetles differed considerably in their response in terms of abundance and diversity. Plants and mollusks, displaying morphological and behavioral adaptations to flooding, showed higher survival rates than the carabid beetles, the adaptation strategies of which were mainly linked to life history. Our results illustrate the complexity of responses of floodplain organisms to extreme flood events. They demonstrate that the efficiency of resistance and resilience strategies is widely dependent on the mode of adaptation.     

Quantitative analysis of pyrrolizidine alkaloids sequestered from diverse host plants in Longitarsus flea beetles (Coleoptera, Chrysomelidae)

Summary. Several species of the flea beetles genus Longitarsus are able to sequester pyrrolizidine alkaloids (PAs) from their host plants. In five Longitarsus species we compare the concentration of PAs present in their host plants belonging to the Asteraceae or Boraginaceae with those found in the beetles. To get an estimate of the intrapopulation variability, three samples of five beetles each and five individual plants were analyzed for each comparison. A strong intrapopulation variability could be detected both among plant and beetle samples. The total concentration found in the beetles varied strongly between species. The local host plant and its phenology influence the concentrations present in the beetles as evidenced in comparisons of a single beetle species from two different hosts and of one beetle species collected at the same site at different times of the year. In addition, different beetle species apparently vary in their capacity to sequester the alkaloids, at the lowest extreme the mean PA concentration in the beetles (0.034 μg PA/mg dry weight) was 1/30 of the mean concentration found in the plant leaves (L. aeruginosus from Eupatorium cannabinum), at the highest extreme (2.098 μg PA/mg dw) the concentration in the beetles was a 1000 fold higher than in the plant leaves (L. nasturtii from Symphytum officinale). The highest mean concentration found in the beetles was 3.446 μg/mg dw (L. exoletus from Cynoglossum officinale). The absolute concentrations found in the beetles are comparable to other insects which have been shown to be effectively defended against their potential predators. Received 22 June 1999; accepted 25 August 1999     

A survey of antennal responses by five species of coniferophagous bark beetles (Coleoptera: Scolytidae) to bark volatiles of six species of angiosperm trees

Summary. Using Porapak Q traps, we collected the bark volatiles of six angiosperm trees native to British Columbia: black cottonwood, Populus trichocarpa Torr. & A. Gray (Salicaceae), trembling aspen, P. tremuloides Michx. (Salicaceae), paper birch, Betula papyrifera Marsh. (Betulaceae), bigleaf maple, Acer macrophyllum Pursh (Aceraceae), red alder, Alnus rubra Bong. (Betulaceae), and Sitka alder, A. viridis ssp. sinuata (Regel) á. L?ve & D. L?ve (Betulaceae). Utilising coupled gas chromatographic-electroantennographic detection analysis, the captured volatiles were assayed for antennal responses in five species of coniferophagous bark beetles (Coleoptera: Scolytidae), sympatric with most or all of the angiosperm trees: the Douglas-fir beetle, Dendroctonus pseudotsugae Hopkins, the mountain pine beetle, D. ponderosae Hopkins, the spruce beetle, D. rufipennis (Kirby), the western balsam bark beetle, Dryocoetes confusus Swaine, and the pine engraver, Ips pini (Say). The identities of 25 antennally-active compounds were determined by coupled gas chromatographic-mass spectroscopic analysis, and co-chromatographic comparisons with authentic chemicals. The compounds identified were: hexanal, (E)-2-hexenal, (Z)-3-hexen-1-ol, 1-hexanol, heptanal, α-pinene, frontalin, benzaldehyde, β-pinene, 2-hydroxycyclohexanone, 3-carene, limonene, β-phellandrene, benzyl alcohol, (E)-ocimene, salicylaldehyde, conophthorin, guaiacol, nonanal, methyl salicylate, 4-allylanisole, decanal, thymol methyl ether, (E)-nerolidol, and dendrolasin. A number of these compounds are known semiochemicals that are active in the behaviour of other organisms, including bark beetles, suggesting a high degree of semiochemical parsimony. Antennally-active compounds ranged from seven in A. viridis to 17 in P. trichocarpa. The fewest number of compounds (9) were detected by I. pini and the largest number (24) were detected by D. pseudotsugae. Six compounds excited the antennae of all five species of bark beetles. The large number of antennally-active compounds detected in common by numerous bark beetles and present in common in numerous nonhost trees supports the hypothesis of olfaction-based recognition and avoidance of nonhost angiosperm trees during the process of host selection by coniferophagous bark beetles. Received 13 December 1999; accepted 14 March 2000     

Species traits as predictors of lepidopteran composition in restored and remnant tallgrass prairies.

Restoration ecologists are increasingly turning to the development of trait-filter models, which predict how evolved traits limit species membership within assemblages depending on existing abiotic or biotic constraints, as a tool to explain how species move from a regional species pool into a restored community. Two often untested assumptions of these models, however, are that species traits can reliably predict species' broadscale distribution and that the effects of traits on community membership do not vary between restored and remnant habitats. The goals of this study were to determine whether combinations of ecological traits predispose moth species toward recolonization of restored prairies and to assess the degree to which restored prairies contain moth assemblages comparable with prairie remnants. In 2004, we collected 259 moth species from 13 tallgrass prairie remnants and restorations in central Iowa. Principal components analysis (PCA) was used to identify significant combinations of ecological traits that were shared by groups of moth species. Logistic regression was then employed to test for significant effects of the trait combinations on the frequency of prairie sites occupied by moth species. PCA partitioned moth traits into four axes that explained a total of 81.6% of the variance. Logistic regression detected significant effects for all four PCA axes on the fraction of sites occupied by moths. Species frequently filtered from the regional species pool into prairies were those that had long flight periods and were multivoltine, displayed a feeding preference for legumes but not other forb families, and were regionally abundant but relatively small in body size. Ordination revealed significant differences in moth communities among prairies, suggesting that species traits and habitat characteristics likely interact to create observed patterns of species recolonization of restorations. Thus, the optimal approach to restoring the lepidopteran fauna of tallgrass prairies may involve locating prairie plantings adjacent to habitat remnants.     

Application of the New Keystone-Species Concept to Prairie Dogs: How Well Does It Work?

Abstract: It has been suggested that the keystone-species concept should be dropped from ecology and conservation, primarily because the concept is poorly defined. This prompted Power et al. (1996) to refine the definition: keystone species have large effects on community structure or ecosystem function (i.e., high overall importance), and this effect should be large relative to abundance (i.e., high community importance). Using prairie dogs ( Cynomys spp.) as an example, I review operational and conceptual difficulties encountered in applying this definition. As applied to prairie dogs, the implicit assumption that overall importance is a linear function of abundance is invalid. In addition, community importance is sensitive to abundance levels, the definition of community, and sampling scale. These problems arise largely from the equation for community importance, as used in conjunction with removal experiments at single abundance levels. I suggest that we shift from the current emphasis on the dualism between keystone and nonkeystone species and instead examine how overall and community importance vary (1) with abundance, (2) across spatial and temporal scales, and (3) under diverse ecological conditions. In addition, I propose that a third criterion be incorporated into the definition: keystone species perform roles not performed by other species or processes. Examination of how these factors vary among populations of keystone species should help identify the factors contributing to, or limiting, keystone-level functions, thereby increasing the usefulness of the keystone-species concept in ecology and conservation. Although the quantitative framework of Power et al. falls short of being fully operational, my conceptual guidelines may improve the usefulness of the keystone-species concept. Careful attention to the factors that limit keystone function will help avoid misplaced emphasis on keystone species at the expense of other species.     

Factors Influencing Movement Patterns of Keel-Billed Toucans in a Fragmented Tropical Landscape in Southern Mexico

Abstract: Various factors influence animal movements in fragmented landscapes, and determining these factors is key to understanding ecological processes at a landscape scale. My goals were (1) to determine what factors influence movements of Keel-billed Toucans (    Ramphastos sulfuratus ) in a fragmented landscape in southern Mexico and (2) to use this information to predict how movement patterns might change if the landscape was altered. I developed a cost-distance geographic information system model that adjusts Euclidean distances by a cost of moving through a certain habitat type. Cost was based on habitat preferences exhibited by toucans. I then used this model to predict how movements might be affected by removal of isolated trees and living fences from the pasture matrix and by removal of forest remnants. Toucans moved more frequently between remnants separated by a low cost-distance value. There was a cost-distance threshold beyond which movements between remnants were rare. Below this threshold, fruit abundance influenced toucan movements but remnant area was not influential in that toucans did not preferentially move to large patches. Remnants close to various other remnants were more frequently visited by toucans, indicating that landscape connectivity influences toucan movements. Toucans incurred a 10–30% cost increase when moving in computer-simulated landscapes, indicating that changes in forest cover or configuration of habitats may negatively affect toucan populations, assuming that increased cost has a fitness consequence. Cost-distance modeling has been relatively unexplored and may be a valuable tool for determining how the configuration of a landscape impedes or facilitates animal movements.