Habitat quality matters for the distribution of an endangered leaf beetle and its egg parasitoid in a fragmented landscape

Fragmentation, deterioration, and loss of habitat patches threaten the survival of many insect species. Depending on their trophic level, species may be differently affected by these factors. However, studies investigating more than one trophic level on a landscape scale are still rare. In the present study we analyzed the effects of habitat size, isolation, and quality for the occurrence and population density of the endangered leaf beetle Cassida canaliculata Laich. (Coleoptera: Chrysomelidae) and its egg parasitoid, the hymenopteran wasp Foersterella reptans Nees (Hymenoptera: Tetracampidae). C. canaliculata is strictly monophagous on meadow sage (Salvia pratensis), while F. reptans can also parasitize other hosts. Both size and isolation of habitat patches strongly determined the occurrence of the beetle. However, population density increased to a much greater extent with increasing host plant density ( = habitat quality) than with habitat size. The occurrence probability of the egg parasitoid increased with increasing population density of C. canaliculata. In conclusion, although maintaining large, well-connected patches with high host plant density is surely the major conservation goal for the specialized herbivore C. canaliculata, also small patches with high host plant densities can support viable populations and should thus be conserved. The less specialized parasitoid F. reptans is more likely to be found on patches with high beetle density, while patch size and isolation seem to be less important.     

A comparison of the responses of two tropical specialist herbivores to host plant patch size

Summary The effects of host plant patch size on the abundances of two specialist herbivores (the chrysomelid beetle, Acalymma innubum and the pentatomid bug, Piezosternum subulatum) were investigated in a natural forest community in the Virgin Islands. Abundances were compared early and late in the season in different sized patches of the cucurbit host plant (Cayaponia americana) growing in open habitat (with no surrounding plant community) and forest habitat (with diverse surrounding plant community). For both herbivore species, adult abundances per patch were positively correlated with patch leaf area, but there was a significant patch size effect (i.e., correlation between herbivore density per unit plant and patch leaf area) only for beetles in the forest habitat. Both herbivore species were significantly affected by surrounding plant diversity, but in opposite ways: beetles were more abundant in open patches whereas bugs were more abundant in forest patches. Relationships between abundance and patch size in open and forest patches changed through the season for both herbivore species. These changing abundance patterns are discussed with respect to (1) increases in the diversity of the plant community surrounding host plant patches, and (2) differences in herbivore movement patterns.     

Butterfly larvae reduce host plant survival in vicinity of alternative host species

Summary Mortality of an annual plant. Collinsia torreyi, was increased in the vicinity of Pedicularis semibarbata plants. This was a consequence of feeding by Euphydryas editha, a herbivore which feeds on both plant species. Mortality of C. torreyi was better predicted by distance from P. semibarbata than by density of C. torreyi. Since not all herbivores are monophagous, interactions involving distance and density effects on plant survival and on vegetational diversity need to be considered with respect to plant and herbivore communities. Some of the complexities of one predator, two prey systems are discussed.     

Does fragmentation of Urtica habitats affect phytophagous and predatory insects differentially?

Effects of habitat fragmentation on the insect community of stinging nettle (Urtica dioica L.) were studied, using 32 natural nettle patches of different area and degree of isolation in an agricultural landscape. Habitat fragmentation reduced the species richness of Heteroptera, Auchenorrhyncha, and Coleoptera, and the abundance of populations. Habitat isolation and area reduction did not affect all insect species equally. Monophagous herbivores had a higher probability of absence from small patches than all (monophagous and polyphagous) herbivore species, and the percentage of monophagous herbivores increased with habitat area. Abundance and population variability of species were negatively correlated and could both be used as a predictor of the percentage of occupied habitats. Species richness of herbivores correlated (positively) with habitat area, while species richness of predators correlated (negatively) with habitat isolation. In logistic regressions, the probability of absence of monophagous herbivores from habitat patches could only be explained by habitat area (in 4 out of 10 species) and predator absence probability only by habitat isolation (in 3 out of 14 species). Presumably because of the instability of higher-trophic-level populations and dispersal limitation, predators were more affected by habitat isolation than herbivores, while they did not differ from herbivore populations with respect to abundance or variability. Thus increasing habitat connectivity in the agricultural landscape should primarily promote predator populations. Received: 4 February 1998 / Accepted: 4 May 1998     

Visual cues override olfactory cues in the host-finding process of the monophagous leaf beetle Altica engstroemi

It is generally assumed that specialist insect herbivores utilize plant odours to find their particular host plants and that visual cues are of minor importance in the host‐finding process. We performed Y‐tube olfactometer bioassays and small‐scale field experiments to determine whether, under laboratory and field conditions, the monophagous herbivore Altica engstroemi J. Sahlberg (Coleoptera: Chrysomelidae: Alticinae) is guided to its host plant Filipendula ulmaria (L.) Maxim. (Rosaceae) by visual or olfactory cues. The olfactometer tests showed that A. engstroemi was never attracted to odours, either from undamaged or from damaged plants. Even starvation for 24 h did not change this behaviour. However, the field experiment showed that visual cues alone were sufficient to attract a significant number of starved beetles when offered a choice between bagged host plants and bagged green plastic control ‘plants’. Our findings contrast with the general view that plant odours constitute the major cue in the host‐finding process among specialized phytophagous insects. A review of the literature for the period 1986–2006 inclusive, relating to host‐plant finding in Chrysomelidae, identified studies of 19 chrysomelid species, all of which were guided by olfactory cues. No species were guided to their host by visual cues. Although some studies demonstrated that chrysomelids may exhibit orientation responses to colour or contrast, our study on A. engstroemi is the only one demonstrating that visual cues affect host‐plant selection in a chrysomelid species. We suggest that the use of visual cues in host‐finding may evolve among chrysomelids with limited dispersal ability in persistent habitats and may be found among species monophagous on abundant host plants that dominate the structure of the plant community, that is, where the host plant's presence is predictable in time and space.     

Genetic and soil-nutrient effects on the abundance of herbivores on willow

The effects of soil-nutrient environment, plant genotype, and the interaction between the two on the resistance of the willow, Salix sericea, to insect species in a diverse herbivore community was measured. We found that soil-nutrient environment influenced plant growth and the abundance of most herbivores of S. sericea. However, environmental effects on herbivore abundance were often modified by plant genetics; the abundance of four of seven herbivores exhibited significant genotypeby-environment interaction effects. Pure genotype effects were mostly small and non-significant. The effects of fertilization differed among herbivores. Several herbivores were more abundant on fertilized plants, one was less abundant, and the abundance of others did not change. We found that feeding guild was a poor predictor of herbivore response. Finally we found significant phenotypic and genetic correlations among growth rate, internode length, and the abundances of several herbivores.     

Individually mark–mass release–resight study elucidates effects of patch characteristics and distance on host patch location by an insect herbivore

1. How organisms locate their hosts is of fundamental importance in a variety of basic and applied ecological fields, including population dynamics, invasive species management and biological control. However, tracking movement of small organisms, such as insects, poses significant logistical challenges. 2. Mass‐release and individual–mark–recapture techniques were combined in an individually mark–mass release–resight (IMMRR) approach to track the movement of over 2000 adult insects in an economically important plant–herbivore system. Despite its widespread use for the biological control of the invasive thistle Carduus nutans, the host‐finding behaviour of the thistle head weevil Rhinocyllus conicus has not previously been studied. Insects were released at different distances from a mosaic of artificially created host patches with different areas and number of plants to assess the ecological determinants of patch finding. 3. The study was able to characterize the within‐season dispersal abilities and between‐patch movement patterns of R. conicus. Weevils found host plant patches over 900 m away. Large patches, with tall plants, situated close to the nearest release point had the highest first R. conicus resights. Patch area and plant density had no effect on the number of weevils resighted per plant; however, R. conicus individuals were more likely to disperse out of small patches and into large patches. 4. By understanding how R. conicus locates host patches of C. nutans, management activities for the control of this invasive thistle can be better informed. A deeper mechanistic understanding of host location will also improve prediction of coupled plant–herbivore spatial dynamics in general.     

Movement patterns of Rhyssomatus lineaticollis Say (Coleoptera: Curculionidae) within and among Asclepias syriaca (Asclepiadaceae) patches in a fragmented landscape

Abstract.  1. Dispersal capabilities of organisms are critical in determining the landscape population structure of species as well as their likelihood of survival in fragmented landscapes. Using mark–recapture techniques on the monophagous weevil Rhyssomatus lineaticollis Say (Curculionidae), within- and between-patch dispersal capabilities, landscape level population structure, and the role of beetle density and host patch characteristics in setting distances, amounts, and timing of dispersal were studied.
2. The data indicate that R. lineaticollis is sedentary, with 50% of recaptured beetles moving < 1 m and the maximum distance moved < 1 km. Within- and between-patch movement of beetles was unrelated to host plant patch characteristics and beetle densities.
3. Despite limited dispersal, R. lineaticollis probably functions as a patchy population in east-central Iowa, U.S.A. because dispersals between patches are common and because all host patches surveyed contained this herbivore, indicating a lack of suitable vacant patches, a prerequisite for metapopulation structure.
4. Between-patch distances are well within the dispersal capabilities of R. lineaticollis , although this may be the result of an increase in the density of patches of its host, Asclepias syriaca , in the landscape over the last 150 years as a result of human disturbance and this species' weedy habit.
5. Metapopulation structure in monophagous prairie herbivores may be most likely in species whose non-weedy host plants form highly predictable resources in space and time, but which are now widely scattered in habitat fragments.     

Effect of plant patch shape on the distribution and abundance of three lepidopteran species associated with Brassica oleracea

• 1 Plant patch shape may affect the abundance of herbivorous insects. Patches of the same size but longer or irregular have a higher perimeter/area relationship (P/A) than square or regular ones, which may determine the immigration, emigration and abundance of individuals in the patch.
• 2 Only specialist species should be affected by plant patch shape. Those species that are more abundant in smaller patches should be more abundant in patches with higher P/A, whereas those that are more abundant in larger patches should be more abundant in patches with lower P/A.
• 3 We studied the density of eggs, larvae and pupae of Pieris brassicae, Plutella xylostella and Trichoplusia ni in square (low P/A) and I‐shaped (high P/A) patches of 144 plants of Brassica oleracea. We also estimated their immigration to these patches, and the final plant weight.
• 4 Plant patch shape affected the abundance, but not the distribution, of the two specialist species. Whereas P. brassicae was denser in I‐shaped patches, P. xylostella was more abundant in square patches. The generalist T. ni was not affected by patch shape. Immigration of P. brassicae was higher in I‐shaped patches, but immigration of P. xylostella and T. ni was not affected by patch shape. Plants were heavier in the centre of square patches.
• 5 Our results suggest that plant patch shape affects the density of herbivorous insects and should be considered independently from other plant patch variables when studying the population dynamics of these organisms.

     

Dispersal ability and host-plant characteristics influence spatial population structure of monophagous beetles

Abstract.  1. Dispersal plays an integral role in determining spatial population structure and, consequently, the long-term survival of many species. Theoretical studies indicate that dispersal increases with population density and decreasing habitat stability. In the case of monophagous insect herbivores, the stability of host-plant populations may influence their spatial population structure.
2. The tallgrass prairie in Iowa, U.S.A. is highly fragmented and most prairie insects face a landscape with fewer habitat patches and smaller host-plant populations than 150 years ago, potentially making dispersal between patches difficult. Some herbivores, however, use native plant species with weedy characteristics that have increased in abundance because of disturbances.
3. Mark–recapture data and presence–absence surveys were used to examine dispersal and spatial population structure of two monophagous beetles with host plants that exhibit different population stability and have responded differently to fragmentation of tallgrass prairie.
4. Chrysochus auratus Fabricius exhibits a patchy population structure and has relatively large dispersal distances and frequencies. Its host plant is variable locally in time and space, but is more abundant than 150 years ago. The other species, Anomoea laticlavia Forster, exhibits a metapopulation or non-equilibrium population structure and has relatively small dispersal distances and frequencies. Its host-plant populations are stable in time and space.
5. The results indicate that dispersal ability of monophagous beetles reflects the life-history dynamics of their host plants, but the spatial population structure exhibited today is strongly influenced by how the host plants have responded to the fragmentation process over both time and space.     

Distribution and colonisation ability of three parasitoids and their herbivorous host in a fragmented landscape

Habitat fragmentation can disrupt communities of interacting species even if only some of the species are directly affected by fragmentation. For instance, if parasitoids disperse less well than their herbivorous hosts, habitat fragmentation may lead to higher herbivory in isolated plant patches due to the absence of the third trophic level. Community-level studies suggest that parasitoids tend to have limited dispersal abilities, on the order of tens of metres, much smaller than that of their hosts, while species-oriented studies document dispersal by parasitoids on the scale of kilometres. In this study the distribution patterns of three parasitoid species with different life histories and their moth host, Hadena bicruris, a specialist herbivore of Silene latifolia, were compared in a large-scale network of natural fragmented plant patches along the rivers Rhine and Waal in the Netherlands. We examined how patch size and isolation affect the presence of each species. Additionally, experimental plots were used to study the colonisation abilities of the species at different distances from source populations.In the natural plant patches the presence of the herbivore and two of the parasitoids, the gregarious specialist Microplitis tristis and the gregarious generalist Bracon variator were not affected by patch isolation at the scale of the study, while the solitary specialist Eurylabus tristis was. In contrast to the herbivore, the presence of all parasitoid species declined with plant patch size. The colonisation experiment confirmed that the herbivore and M. tristis are good dispersers, able to travel at least 2 km within a season. B. variator showed intermediate colonisation ability and E. tristis showed very limited colonisation ability at this spatial scale. Characteristics of parasitoid species that may contribute to differences in their dispersal abilities are discussed.     

Do distances among host patches and host density affect the distribution of a specialist parasitoid?

The effect of spatial habitat structure and patchiness may differ among species within a multi-trophic system. Theoretical models predict that species at higher trophic levels are more negatively affected by fragmentation than are their hosts or preys. The absence or presence of the higher trophic level, in turn, can affect the population dynamics of lower levels and even the stability of the trophic system as a whole. The present study examines different effects of spatial habitat structure with two field experiments, using as model system the parasitoid Cotesia popularis which is a specialist larval parasitoid of the herbivore Tyria jacobaeae. One experiment examines the colonisation rate of the parasitoid and the percentage parasitism at distances occurring on a natural scale; the other experiment examines the dispersal rate and the percentage parasitism in relation to the density of the herbivore and its host plant. C. popularis was able to reach artificial host populations at distances up to the largest distance created (at least 80 m from the nearest source population). Also, the percentage parasitism did not differ among the distances. The density experiment showed that the total number of herbivores parasitised was higher in patches with a high density of hosts, regardless of the density of the host plant. The percentage parasitism, however, was not related to the density of the host. The density of the host plant did have a (marginally) significant effect on the percentage parasitism, probably indicating that the parasitoid uses the host plant of the herbivore as a cue to find the herbivore itself. In conclusion, the parasitoid was not affected by the spatial habitat structure on spatial scales that are typical of local patches.     

Exotic Slugs Pose a Previously Unrecognized Threat to the Herbaceous Layer in a Midwestern Woodland

Developing effective restoration strategies requires first identifying the underlying factors limiting native plant recovery. The slug Deroceras reticulatum is an important herbivore in Europe, a global agricultural pest, and is introduced and abundant throughout eastern North America, but little information is available on the effect of this exotic herbivore on the forest herbaceous layer. Here, we test the palatability of 12 forest herbs to the introduced slug D. reticulatum and use field surveys to determine the degree to which slugs are damaging plants in the field. In laboratory feeding trials, slugs readily consumed most plants, but avoided the grass Elymus virginicus, the invasive forb Alliaria petiolata (garlic mustard), and thicker leaved plants. In the field, we documented significant slug damage, with close to 50% or more of plant leaves damaged by slugs on five of the six native species tested. Slug damage in the field was predicted by laboratory‐determined acceptability, but was significantly greater on short‐statured rosette species than on erect plants for a given acceptability value. Our results identify introduced slugs as an important, but overlooked obstacle to forest herb restoration and potential drivers of larger scale understory compositional change. The relaxed herbivore pressure on A. petiolata, relative to native competitors, suggests that invasive plant removal alone may not result in the recovery of native flora. Rather, restoration of unpalatable native species should accompany invasive plant control in slug invaded areas. Erect forbs, thick‐leaved plants, and graminoids should have the greatest success where introduced slugs are abundant.     

Quality or quantity: the direct and indirect effects of host plants on herbivores and their natural enemies

Resource quality (plant nitrogen) and resource quantity (plant density) have often been argued to be among the most important factors influencing herbivore densities. A difficulty inherent in the studies that manipulate resource quality, by changing nutrient levels, is that resource quantity can be influenced simultaneously, i.e. fertilized plants grow more. In this study we disentangled the potentially confounding effects of plant quality and quantity on herbivore trophic dynamics by separately manipulating nutrients and plant density, while simultaneously reducing pressure from natural enemies (parasitoids) in a fully factorial design. Plant quality of the sea oxeye daisy, Borrichia frutescens, a common coastal species in Florida, was manipulated by adding nitrogen fertilizer to increase and sugar to decrease available nitrogen. Plant density was manipulated by pulling by hand 25 or 50% of Borrichia stems on each plot. Because our main focal herbivore was a gall making fly, Asphondylia borrichiae, which attacks only the apical meristems of plants, manipulating plant nitrogen levels was a convenient and reliable way to change plant quality without impacting quantity because fertilizer and sugar altered plant nitrogen content but not plant density. Our other focal herbivore was a sap-sucker, Pissonotus quadripustulatus, which taps the main veins of leaves. Parasitism of both herbivores was reduced via yellow sticky traps that caught hymenopteran parasitoids. Plant quality significantly affected the per stem density of both herbivores, with fertilization increasing, and sugar decreasing the densities of the two species but stem density manipulations had no significant effects. Parasitoid removal significantly increased the densities of both herbivores. Top-down manipulations resulted in a trophic cascade, as the density of Borrichia stems decreased significantly on parasitoid removal plots. This is because reduced parasitism increases gall density and galls can kill plant stems. In this system, plant quality and natural enemies impact per stem herbivore population densities but plant density does not.     

Ploidy level interacts with population size and habitat conditions to determine the degree of herbivory damage in plant populations

Recently it has been suggested that ploidy level of a plant population may have important effects on plant‐animal interactions. Plant‐animal interactions can also be strongly altered by factors such as plant population size and habitat conditions. It is, however, not known how these factors interact to shape the overall pattern of plant‐animal interactions. I studied the interaction between a perennial plant, Aster amellus, and a monophagous herbivorous moth, Coleophora obscenella, and investigated the effect of ploidy level of the plant population, plant population size, isolation and habitat conditions on density of the insect, damage by the insect, and plant performance. Ploidy level, plant population size and habitat conditions, but not isolation, strongly influence plant‐herbivore interactions. Furthermore, there are significant interactions between effects of ploidy level and plant population size and between ploidy level and isolation. Hexaploid plants suffer higher seed damage by the herbivore, but their seed production is still higher than that of diploids. Herbivores thus partly limit the evolutionary success of the hexaploid plants. Plant‐animal interactions are also strongly determined by plant population size. Small populations of A. amellus (below forty flowering ramets) host no C. obscenella larvae, indicating a minimum A. amellus population size that can sustain a viable C. obscenella population. Negative and positive effects of plant population size balance and result in no relationship between plant population size and number of developed seeds per flower head. The results also show a significant interaction between ploidy level and plant population size, indicating that the increase in density of C. obscenella larvae with plant population size is greater in hexaploid than in diploid populations. The results also indicate that the effect of ploidy level on plant‐herbivore interactions can be altered by plant population size, which suggests that plant‐herbivore interactions are driven by a complex of interactions among different factors. Studying each factor separately could thus lead to biased conclusions about patterns of interactions in such systems.     

Impact of ecological factors on the initial invasion of Bt transgenes into wild populations of birdseed rape (Brassica rapa)

The inevitable escape of transgenic pollen from cultivated fields will lead to the emergence of transgenic crop-wild plant hybrids in natural patches of wild plants. The fate of these hybrids and that of the transgene depend on their ability to compete with their wild relatives. Here we study ecological factors that may enhance the fitness of genetically modified hybrids relative to wild plants for a Bacillus thuringiensis (Bt) transgene conferring resistance to insects. Mixed stands of wild plants and first-generation hybrids were grown under different conditions of herbivore pressure and density, with Bt oilseed rape (Brassica napus) as the crop and B. rapa as the wild recipient. Biomass and fitness components were measured from plant germination to the germination of their offspring. The frequency of transgenic seedlings in the offspring generation was estimated using the green fluorescent protein marker. The biomass of F₁ Bt-transgenic hybrids relative to that of wild-type plants was found to be sensitive to both plant density and herbivore pressure, but herbivore pressure appeared as the major factor enhancing their relative fitnesses. In the absence of herbivore pressure, Bt hybrids produced 6.2-fold fewer seeds than their wild neighbors, and Bt plant frequency fell from 50% to 16% within a single generation. Under high herbivore pressure, Bt hybrids produced 1.4-fold more seeds, and Bt plant frequency was 42% in the offspring generation. We conclude that high-density patches of highly damaged wild plants are the most vulnerable to Bt-transgene invasion. They should be monitored early to detect potential transgene spread.     

Impact of habitat modification on the distribution and abundance of fruit flies (Diptera: Tephritidae) in Southeast Queensland

Loss of rainforest because of agricultural and urban development may impact the abundance and diversity of species that are rainforest natives. Tropical fruit flies are one group of such organisms indigenous to rainforests. In southeast Queensland, a region subject to rapid urbanization, we assessed the impact of habitat disturbance on the distribution and abundance of native fruit flies. Data on four species (Bactrocera tryoni, Bactrocera neohumeralis, Bactrocera chorista, and Dacus aequalis) were gathered and analyzed over 6 months in three habitat types: suburbia, open sclerophyll forest, and rainforest. We also analyzed the data at a combined "dacine fruit fly" level incorporating all fruit fly species trapped over the period of study (as might occur in a biodiversity assessment): these included the four species already named and Bactrocera melas, Bactrocera bryoniae, Bactrocera newmani, and Dacus absonifacies. Analysis at the species level showed that the polyphagous pest species responded differently to the monophagous species. Bactrocera tryoni, which has more exotic than native hosts, was positively affected by transformation of natural habitat into suburbia whereas B. neohumeralis, which has nearly identical numbers of native and exotic hosts, was found equally across habitat types. Bactrocera chorista and Dacus aequalis, each monophagous on a species-specific rainforest host plant, were most abundant in rainforest. The analysis based on the combined data suggests that replacing rainforest with suburbia has a neutral, or even positive, effect on the abundance of fruit flies as a whole. At the species level, however, it can be seen that this is an erroneous conclusion biased by the abundance of a single pest species. Our discussion raises the issue of analyses at supraspecific levels in biodiversity and impact assessment studies. Received: March 6, 2000 / Accepted: June 19, 2000     

Asymmetric priority effects influence the success of invasive forest insects

1. Settlement timing is often an important factor in interspecific herbivore interactions, as early‐arriving species may encounter higher resource availability and/or avoid induced defences. Despite the general importance of priority effects to the outcome of herbivore interactions, there has been little exploration of such interactions on woody host plants where their impact can only be measured over multiple years. 2. In the eastern U.S.A., two invasive species, the hemlock woolly adelgid Adelges tsugae and the elongate hemlock scale Fiorinia externa, share a native host, eastern hemlock Tsuga canadensis. Their interaction and its consequences were investigated for plant growth – hemlock saplings that had been inoculated with either A. tsugae or F. externa, starting in spring 2007, were cross‐infested with the other insect in spring 2009. A set of uninfested trees was simultaneously infested with A. tsugae, F. externa, both, or neither insect (= control), and insect density and plant growth was assessed in all treatments. 3. Adelges tsugae settlement rates did not differ if it settled alone or simultaneously with F. externa, but were ～45% lower on trees previously infested with F. externa. There was no difference in F. externa settlement rates, and plant growth did not differ substantively between any of the herbivore treatments. 4. At a temporal scale (i.e. multiple growing seasons) appropriate to interactions between woody plants and their herbivores, this work demonstrates that plant‐mediated priority effects can substantially affect herbivore settlement and thus the outcome of interspecific competition.     

Combined effects of patch size and plant nutritional quality on local densities of insect herbivores

Plant–insect interactions occur in spatially heterogeneous habitats. Understanding how such interactions shape density distributions of herbivores requires knowledge on how variation in plant traits (e.g. nutritional quality) affects herbivore abundance through, for example, affecting movement rates and aggregation behaviour. We studied the effects of plant patch size and herbivore-induced differences in plant nutritional quality on local densities of insect herbivores for two Brassica oleracea cultivars, i.e. white cabbage and Brussels sprouts. Early season herbivory as a treatment resulted in measurable differences in glucosinolate concentrations in both cultivars throughout the season. Herbivore induction and patch size both influenced community composition of herbivores in both cultivars, but the effects differed between species. Flea beetles (Phyllotreta spp.) were more abundant in large than in small patches, and this patch response was more pronounced on white cabbage than on Brussels sprouts. Herbivore-induction increased densities in all patches. Thrips tabaci was also more abundant in large patches and densities of this species were higher on Brussels sprouts than on white cabbage. Thrips densities were lower on induced than on control plants of both cultivars and this negative effect of induction tended to be more pronounced in large than in small patches. Densities of the cabbage moth (Mamestra brassicae) were lower on Brussels sprouts than on white cabbage and lower on herbivore-induced than on uninduced plants, with no effect of patch size. No clear effects of patch size and induction were found for aphids. This study shows that constitutive and herbivore-induced differences in plant traits interact with patch responses of insect herbivores.     

Insect herbivory and herbivores of Ficus species along a rain forest elevational gradient in Papua New Guinea

Classic research on elevational gradients in plant–herbivore interactions holds that insect herbivore pressure is stronger under warmer climates of low elevations. However, recent work has questioned this paradigm, arguing that it oversimplifies the ecological complexity in which plant–insect herbivore interactions are embedded. Knowledge of antagonistic networks of plants and herbivores is however crucial for understanding the mechanisms that govern ecosystem functioning. We examined herbivore damage and insect herbivores of eight species of genus Ficus (105 saplings) and plant constitutive defensive traits of two of these species, along a rain forest elevational gradient of Mt. Wilhelm (200–2,700 m a.s.l.), in tropical Papua New Guinea. We report overall herbivore damage 2.4% of leaf area, ranging from 0.03% in Ficus endochaete at 1,700 m a.s.l. to 6.1% in F. hombroniana at 700 m a.s.l. Herbivore damage and herbivore abundances varied significantly with elevation, as well as among the tree species, and between the wet and dry season. Community-wide herbivore damage followed a hump-shaped pattern with the peak between 700 and 1,200 m a.s.l. and this pattern corresponded with abundance of herbivores. For two tree species surveyed in detail, we observed decreasing and hump-shaped patterns in herbivory, in general matching the trends found in the set of plant defenses measured here. Our results imply that vegetation growing at mid-elevations of the elevational gradient, that is at the climatically most favorable elevations where water is abundant, and temperatures still relatively warm, suffers the maximum amount of herbivorous damage which changes seasonally, reflecting the water availability.