Movement patterns of an Australian chrysomelid beetle in a stand of two Eucalyptus host species

Summary We examined the movements of Chrysophtharta hectica, a eucalypt-feeding chrysomelid beetle in New South Wales, Australia, in relation to the beetle's sex, age and life-history, and to attributes of its Eucalyptus host plants. Beetle movements within the site were not influenced by beetle age or sex, but may be related to generation. Beetle distributions on the two host plant species, Eucalyptus stellulata and E. pauciflora, were generally clumped. Some of this clumping resulted from preference for E. stellulata over E. pauciflora. Clumping of beetles also occurred within host plant species; some plant individuals were consistently heavily used by beetles over the course of three years. We examined nutritional, spatial and biomass attributes of plants and found plant height and foliage production to be the best predictors of beetle numbers.     

Effects of environmental context on the susceptibility of Atriplex patula to attack by herbivorous beetles

The susceptibility of plants to attack by insect herbivores often depends on local environmental conditions. This study documents variation in herbivore damage by the chrysomelid beetle Erynephala maritima to the annual forb Atriplex patula in two microhabitats within New England salt marshes: bare patches and dense matrix vegetation. Environmental conditions within bare patches differ from those within matrix vegetation in a number of ways. Bare patches are characterized by the absence of perennial grasses and rushes (matrix vegetation) and greater levels of physical stress, and are rapidly colonized by the fugitive annual, Salicornia europaea, a second host plant of these beetles. Surveys of herbivore damage across three marshes revealed that A. patula in bare patches had a greater proportion of leaves damaged by beetles than those within matrix vegetation. Presence or absence of matrix vegetation and presence or absence of S. europaea were experimentally manipulated to determine the proximate cause of this pattern. The presence of S. europaea significantly increased the susceptibility of A. patula to herbivory in experimental plots. Both the extent of herbivore damage to plants and the proportion of plants damaged through time were greater in treatments with S. europaea than in controls, regardless of the presence or absence of matrix vegetation. Plants in S. europaea addition treatments were also less likely to survive to reproduction. Decreased survival appears to result from increased herbivory, suggesting that the negative effect of S. europaea on A. patula is mediated indirectly through shared insect herbivores. These results support the hypothesis that indirect interactions between alternative host plants, mediated by insect herbivores, can be important in natural communities. Received: 9 January 1999 / Accepted: 29 April 1999     

Habitat selection of three chrysomelid species associated with Rumex spp.

Summary Habitat selection of three chrysomelid beetles feeding on the Rumex plants was studied in relation to the ecological requirements of the three species and the habitat characteristics arising in their unstable and often unpredictable environment. The different ecological requirements and bionomic characteristics of the three chrysomelids were associated not only with their differing degrees of feeding specialization; they also influenced habitat selection. For example, the small body size and lower food demands of Mantura clavareaui, which is a feeding specialist, allowed it to exploit any habitat, irrespective of habitat size and duration. The other specialist, Gastrophysa atrocyanea, selected larger habitats because of the relatively brief lifespan of its adults after overwintering, its reduced mortality, and greater food demand. On the other hand, Galerucella vittaticollis, which is a multivoltine, general feeder, selected the habitat that lasted longer, rather than one that was larger. This species' life cycle is not synchronized with Rumex phenology, but its adults are long-lived and it has a flexible life cycle. The different patterns in habitat selection among these three chrysomelids prevented serious overlapping of their habitats; thus, the habitat overlapping of 2 or 3 species was 34.62%, that of all 3 species 3.85%. Most of the habitats utilized by Gastrophysa, however, overlapped those of the other two species, because Gastrophysa's habitat utilization range was the narrowest of the three. In Mantura, the degree of overlap with the other two species was lowest because this beetle had the broadest range of habitat utilization. On the other hand, Galerucella adjusted the temporal, rather than the spatial coordinates of its habitat usage when it occupied habitats where the other two specialists were also present. Galerucella, unlike the other two species, is multivoltine. Consequently, it was able to coexist in habitats occupied by the others by exploiting the host plants at times when the other species were numerically few, or even absent. No direct interspecific competition among adults of the three species was observed. Their different patterns of habitat selection created by their different responses to available food allowed them to coexist on Rumex without direct competition.     

Trade-offs in plant responses to herbivory influence trophic routes of production in a freshwater wetland

Responses of aquatic macrophytes to leaf herbivory may differ from those documented for terrestrial plants, in part, because the potential to maximize growth following herbivory may be limited by the stress of being rooted in flooded, anaerobic sediments. Herbivory on aquatic macrophytes may have ecosystem consequences by altering the allocation of nutrients and production of biomass within individual plants and changing the quality and quantity of aboveground biomass available to consumers or decomposers. To test the effects of leaf herbivory on plant growth and production, herbivory of a dominant macrophyte, Nymphaea odorata, by chrysomelid beetles and crambid moths was controlled during a 2-year field experiment. Plants exposed to herbivory maintained, or tended to increase, biomass and aboveground net primary production relative to controls, which resulted in 1.5 times more aboveground primary production entering the detrital pathway of the wetland. In a complementary greenhouse experiment, the effects of simulated leaf herbivory on total plant responses, including biomass and nutrient allocation, were investigated. Plants in the greenhouse responded to moderate herbivory by maintaining aboveground biomass relative to controls, but this response occurred at the expense of belowground growth. Results of these studies suggest that N. odorata may tolerate moderate levels of herbivory by reallocating biomass and resources aboveground, which in turn influences the quantity, quality and fate of organic matter available to herbivores and decomposers.     

Increased Reproductive Allocation of Rumex japonicus Caused by Leaf Clipping

Abstract To elucidate the effects of herbivory by chrysomelid beetles on Rumex japonicus, rosette leaves were clipped and the subsequent fruit production and root growth were observed. The increase of leaf biomass of some clipped plants was greater than that of control plants, although this varied among individual plants. The root growth of clipped plants was less than that of control plants. Fruit production increased with plant size, and there was no difference in fruit production between clipped and control plants. Reproductive allocation (fruit biomass, relative to fruit biomass plus root growth) increased with plant size; it was greater in clipped plants than in control ones. Based on these results, reproductive allocation strategy against herbivory was discussed.     

Heterogeneity of plant phenotypes caused by herbivore-specific induced responses influences the spatial distribution of herbivores

Abstract.  1. Herbivory can induce resistance in a plant and the induced phenotype may be disfavoured by subsequent herbivores. Yet, as the distance between plants in a population increases, limited mobility may make a herbivore more likely to feed and oviposit on host plants in its immediate surroundings.
2. The present study tested whether a herbivore's preference and distribution across plants with different induced phenotypes was influenced by the spatial distribution of plants. A fragmented population of Solanum dulcamara plants was created. This consisted of discrete, spatially separated patches with different histories of damage, either herbivory from adult flea beetles ( Psylliodes affinis ), tortoise beetles ( Plagiometriona clavata ), or mechanical damage. Each patch was separated by 7 m and consisted of 12 plants that were spaced 30 cm apart. Then a fixed number of adult tortoise beetles were introduced to each patch, and movement and oviposition within and between spatially separate homogeneous patches (receiving one type of damage) were compared with movement and oviposition within heterogeneous patches (containing all three types of damage) over the growing season.
3. Flea beetle and tortoise beetle herbivory consistently induced different phytochemical responses in S. dulcamara (polyphenol oxidase and peroxidase), and adult tortoise beetles avoided oviposition on the flea beetle induced plants within heterogeneous patches. However, between homogeneous patches, plant phenotype did not influence oviposition. Colonisation by naturally occurring flea beetle adults followed a similar pattern.
4. These results suggest that the heterogeneity of plant phenotypes can influence herbivore choice and distribution at small but not large spatial scales.     

The role of aggregation in the response of Mexican bean beetles to host-plant density

Summary According to the resource concentration hypothesis, specialized insect herbivores predominantly attack host plants growing in pure, large, and/or dense stands because they are more likely to find and less likely to leave such stands. This study examines movement of a herbivorous beetle, Epilachna varivestis, in an attempt to understand why the beetle's numbers per plant increase with plant density. I studied immigration into, emigration from, and movement within host patches by following movements of individually marked beetles, while experimentally varying host-plant density. In order to study the effect of conspecifics on movement, I varied the number of beetles released on the same plant.The probability of intrapatch movement decreased, and the probability of emigration increased when plant density was reduced. Both immigration rate and intrapatch movement were strongly affected by presence or absence of conspecifics. Beetles were much more likely to stop at plants with a high number of beetles, compared to plants without beetles.On the basis of these results I advance a model that provides a mechanistic explanation of why dense patches should acquire high herbivore loads in the E. varivestis-garden bean system. Movement in Mexican bean beetles is strongly aggregative, and in dense patches frequent intrapatch movement provides ample opportunity for beetle aggregations to build up. In sparse patches, however, intrapatch movement is virtually absent and such aggregations are less likely to arise.     

Interpatch movement of the red milkweed beetle,Tetraopes tetraophthalmus: individual responses to patch size and isolation

Individual movement patterns and the effects of host plant patch size and isolation on patch occupancy were examined for red milkweed beetles, Tetraopes tetraophthalmus, residing in a heterogeneous landscape. Male beetles were found to move both more often and farther between host plant patches than female beetles, and this difference affected the patterns of patch occupancy observed. Overall, unoccupied milkweed patches were smaller and more isolated than patches occupied by beetles. Patches uninhabited by females tended to be more isolated, but not necessarily smaller, than patches with female beetles, indicating that females may be affected more by patch isolation than patch size. Presence of male beetles on patches showed a stronger response to patch size than to patch isolation. Differences in movement between males and females illustrate the need for demographically based dispersal data. Comparisons of Tetraopes interpatch movement patterns between landscapes composed of patches of different size revealed that landscapes with overall smaller patches may have greater rates of interpatch movement.     

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.     

Host‐searching responses to herbivory‐associated chemical information and patch use depend on mating status of female solitary parasitoid wasps

1. In a tritrophic interaction system consisting of plants, herbivores, and their parasitoids, chemicals released from plants after herbivory are known to play important roles for many female parasitoids to find their hosts efficiently. On the plant side, chemical information associated with herbivory can act as an indirect defence by attracting the natural enemies of the host herbivores. 2. However, mated and virgin females of haplodiploid parasitoids might not necessarily respond to such chemical cues in the same way. Since virgin females can produce only sons, they might refrain from searching for hosts to invest eggs until copulation, in order to produce both sexes. 3. Here, we investigated differential host‐searching behaviours shown by mated and virgin females in the solitary parasitoid wasp, Cotesia vestalis, in response to herbivory‐associated chemical information from cruciferous plants infested by their host larvae, Plutella xylostella. 4. Mated females showed a significantly higher flight preference for host‐infested plants over intact plants, while no preference was observed with virgin females. Mated females also showed more intensive antennal searching and ovipositor probing behaviours to leaf squares with wounds caused by hosts than did virgin females. Furthermore, mated females stayed longer in host patches with higher parasitism rates than virgin females. 5. These results indicate that mating status of C. vestalis females clearly influences their host‐searching behaviour in response to herbivory‐associated chemical information and patch exploitation. Female parasitoids seem to forage for hosts depending on their own physiological condition in a tritrophic system.     

Host-specificity of a root borer,Bembecia chrysidiformis [Lep.: Sesiidae], a potential control agent forRumex spp. [Polygonaceae] in Australia

Bembecia chrysidiformis (Esper) [Lep.: Sesiidae] was examined for its natural history and specificity toRumex spp. (Polygonaceae) which are weeds in Australia. Adults of this southern European insect appear in late spring to summer. Eggs are laid on the dried, seed bearing stems of perennialRumex plants. The larvae tunnel inside the root during summer through to the next spring. In nature, the larvae are round inRumex species of the subgeneraRumex andAcetosa. In host-specificity tests with 1st instar larvae, the roots of a number of genera within thePolygonaceae were attacked. Larvae died on a range of plants from other families except inPersea americana Miller(Lauraceae), Helianthemum nummularium (L.) Miller (Cistaceae) andQuercus ilex L. (Fagaceae) where larvae fed on the stems. The insect was judged safe for release in Australia by assessing aspects of its biology, its known host plants, and the lack of reported attack on other plants.      

Influence of Forest Types and Effects of Forestry Activities on Species Richness and Composition of Chrysomelidae in the Central Mountainous Region of Japan

Species richness and composition of the Chrysomelidae (Coleoptera) were studied in larch (Larix kaempheri [Lamb.] Carrière) plantations, secondary forests, and primary forests. In addition, the effects of forest management practices, such as thinning and long rotation, were examined in the larch plantation. The species richness of Chrysomelidae was higher in the larch plantation than in the secondary forest or in the primary forest. Among the larch plantations, the species richness in old-aged plantations was higher than that in middle-aged plantations. The composition of the beetle assemblages in the larch plantation differed from that in the secondary forest or in the primary forest. Exosoma akkoae (Chujo), Batophila acutangula Heikertinger, and Calomicrus nobyi Chujo were caught with a bias toward the larch plantation. Longitarsus succineus (Foudras) and Sphaeroderma tarsatum Baly were caught more in the secondary forest and the primary forest, respectively. More B. acutangula and S. tarsatum were caught in stands where their host plants occurred at higher rates. Species richness of understory plants was an important factor for chrysomlid species richness, and frequency of host occurrence affected the number of individuals of leaf beetles examined. It seems that forest types and forest management practices affect host plants as well as Chrysomelidae, and that these effects on the host plants also influence chrysomelid assemblages.     

Colonization of host patches following long-distance dispersal by a goldenrod beetle, Trirhabda virgata

1. The arrival of the chrysomelid beetle Trirhabda virgata on isolated patches of its host Solidago altissima was closely monitored to determine how conspecific density and host condition influence colonization.
2. Experimental host patches, which were set on the roof of a four-storey building located 0.7 km from the nearest naturally occurring hosts, were frequently colonized by beetles over a 2-week dispersal period.
3. Females preferred lush host patches that were free from simulated Trirhabda chewing damage. Females colonized lush patches more often than defoliated patches at two spatial scales, when patches were 2 m and 25 m apart. Males did not show a strong preference for lush plants.
4. Males aggregated on plants that already contained adult conspecifics, apparently increasing their reproductive success. Females did not respond to the presence of adults on the patch.
5. Ninety-five per cent of the females arriving on the isolated plants had mated before flying, indicating that lone females are able to colonize empty stands.
6. By avoiding heavily defoliated plants, females should dissipate local outbreaks and spread their offspring away from over-exploited areas.     

Coreus marginatus (Heteroptera: Coreidae) as a natural enemy of Rumex obtusifolius (Polygonaceae)

Rumex obtusifolius is the preferred host plant of Coreus marginatus. The survival of nymphal C. marginatus, its development on a diet of R. obtusifolius and the effects of herbivory on Rumex seed were investigated. Although the diet consisting solely of seed did not allow complete development, growth rates and survival of nymphs were proportionate to the quantity of provided seed. Nymphal feeding did not noticeably decrease seed mass but significantly decreased the proportion of germinating seed. In this way C. marginatus affects Rumex reproduction. Germination ability decreased with decreasing seed availability and increasing duration of seed predation. Nymphs confined to Rumex inflorescences in the field-completed development and significantly reduced both seed mass and germination ability. The mortality of seed overwintering on standing dry shoots of dock plants was moderately affected by feeding of C. marginatus.     

Leafhopper-induced plant resistance enhances predation risk in a phytophagous beetle

Many herbivores elicit biochemical, physiological, or morphological changes in their host plants that render them more resistant to co-occurring herbivores. Yet, despite the large number of studies that investigate how induced resistance affects herbivore preference and performance, very few have simultaneously explored the cascading effects of induction on higher trophic levels and consequences for prey suppression. In our study system, early-season herbivory by leafhoppers elevated plant resistance to subsequent attack by chrysomelid beetles sharing the same host plant. Notably, beetles feeding on leafhopper-damaged plants incurred developmental penalties (e.g., prolonged time in early larval instars) that rendered them more susceptible to predation by natural enemies. As a result, the combined bottom-up effect of leafhopper-induced resistance and the top-down effect of enhanced predation resulted in the synergistic suppression of beetle populations. These results emphasize that higher trophic level dynamics should be considered in conjunction with induced resistance to better understand how plants mediate interspecific interactions in phytophagous insect communities.     

Locating host plants via orientation to standing visual targets has dispersal benefits for the monophagous leaf beetle Ambrostoma quadriimpressum

The highly specialized chrysomelid Ambrostoma quadriimpressum Motschulsky (Coleoptera: Chrysomelidae) is strictly monophagous on elm trees, Ulmus pumila L. (Ulmaceae) and a few of its close relatives. In order to elucidate how the adult beetles find their host plants, we performed both choice and no‐choice arena field experiments to examine their spatial maneuvering under seminatural conditions. In the no‐choice experiments, three treatments were tested in which individual beetles in thanatosis were released in the vicinity of a host plant, in which they were released facing (1) toward the host, (2) away from the host but toward a black paper column, or (3) away from the host with no standing target in view. Videorecordings of the beetles’ walking tracks showed that a standing visual target in field of vision strongly influenced their behavior. The host plant and black paper column equally attracted beetles released facing toward them. In choice experiments, the beetles randomly moved to both host and non‐host plants, as well as to the host plant and the black paper column, indicating that they do not discriminate host plants from a distance. The results suggest that this monophagous beetle locates host plants via orientation to standing visual targets. The results also indicate that the beetle encounters hosts by chance. The benefit of such a host‐finding mechanism in light of dispersion and emigration of the beetle is discussed.     

Differences in morphology and reproductive traits of Galerucella nymphaeae from four host plant species

The water lily beetle Galerucella nymphaeae L. (Coleoptera: Chrysomelidae) exploits different hosts, including Nuphar lutea Sm. and Nymphaea alba L. (both Nymphaeaceae), as well as Polygonum amphibium L. and Rumex hydrolapathum Hudson (both Polygonaceae). The present study investigates whether within-species differences in morphological and reproductive traits are associated with differences in host species exploitation. A total of 1103 adult beetles were collected from 11 localities in The Netherlands, one of which contained all four hosts and three other localities contained hosts from both families (sympatric localities). Adults originating from Nuphar and Nymphaea were on average darker in colour and larger in size and had disproportionally bigger mandibles than beetles originating from Polygonum and Rumex across the 11 localities. Head capsules of first instar larvae from Nymphaeaceae hosts were between 17% and 28% larger than those of larvae from Polygonaceae hosts. Furthermore, beetles from Nuphar and Nymphaea laid larger sized eggs, but fewer eggs per clutch than beetles originating from Polygonum and Rumex. Although host related variation was less pronounced at the sympatric localities than in the allopatric localities, differences in larval and adult size were still highly significant at the sympatric localities. It is not clear whether the observed differences are genetically based, as opposed to host induced. However, leaf toughness varied among species in a way suggesting that leaf toughness may be partly responsible for host related differences in G. nymphaeae.     

Impact of Host Plant Connectivity,Crop Border and Patch Size on Adult Colorado Potato Beetle Retention

Tagged Colorado potato beetles (CPB), Leptinotarsa decemlineata (Say), were released on potato plants, Solanum tuberosum L., and tracked using a portable harmonic radar system to determine the impact of host plant spatial distribution on the tendency of the pest to remain on the colonized host plant or patch. Results confirmed the long residency time on the host plant and showed that close connection of the plant to neighboring plants hastened dispersal between plants. Tracking walking CPB for over 6 h in small potato plots revealed that all types of mixed borders tested (potato/bare ground, potato/timothy and potato/woodland) acted as a strong barrier and retained beetles within the patch. In another experiment in potato patches surrounded by bare ground borders, tracked walking CPB displayed similar behaviour for up to four days. The distribution of turning angles in the CPB walking paths was not uniform and corresponded to beetles following the edge rows of potato patches in response to the crop border barrier or reversing their direction as they reached the end of a row and therefore a border. Patch size had no or little effect on beetle retention in the patch. The relative distribution of counts of tagged beetles detected among small (16 m²), medium (64 m²) and large size (256 m²) patches of potato four days after initial release remained similar to that of numbers released. Even though mixed crop borders were a strong barrier to walking CPB emigrating from potato patches, the departure rate of beetles over time was high. Results suggest that the effect of mixed borders is largely limited to dispersal by walking and does not apply to beetles leaving host patches by flight. The manipulation of crop borders and patch size seem to have limited potential for the management of CPB emigrating from potato fields.     

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.     

Insect- and herbicide-resistant transgenic eucalypts*

Transgenic Eucalyptus camaldulensis containing both the insecticidal cry3A gene and the bar gene (conferring tolerance to the herbicide glufosinate ammonium) have been produced by Agrobacterium tumefaciens-mediated transformation of seedling explants. Transgenic plants from two lines tested were resistant to first instars of chrysomelid beetles that are important pests of commercial Australian eucalypt plantations. Both lines also exhibit tolerance to the broad-spectrum herbicide Liberty® at 6 l/ha (1.2 kg active ingredient per hectare), twice the field application rate. Transgenic insect- and herbicide-resistant eucalypts like these are likely to provide better insect and weed control options in plantations, particularly during the vulnerable establishment phase, provided that any adverse ecological impacts of releasing transgenic trees into the environment can be assessed and minimized.