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.     

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     

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.     

Preferences of six leaf beetle species among qualitatively different leaf age classes of three Salicaceous host species

Summary. As Salicaceous plants produce new leaves for a prolonged period of time, they expose a wide range of differentially aged leaves to herbivores during the growing season. In this work, I show that young leaves of three Salicaceous species, Populus tremula L., Salix phylicifolia L. and S. pentandra L., contain more nitrogen than conspecific old leaves. In P. tremula and S. pentandra young leaves also contained more low-molecular weight secondary compounds, phenolic glucosides. Leaves of S. phylicifolia did not contain phenolic glucosides in detectable amounts. Furthermore, in P. tremula and S. pentandra young leaves contained less polymeric digestability-reducing phenolics, condensed tannins, than old leaves. In S. phylicifolia, higher concentrations of condensed tannins were found in young leaves. In laboratory feeding trials with six leaf beetle species, young leaves of the studied plants were invariably preferred in all tested herbivore × host species combinations. In particular, it is remarkable that three leaf beetle species with known different overall relationships to phenolic glucosides equally preferred more glucoside-containing young S. pentandra leaves over conspecific old ones. Four beetle species were found to prefer young leaves of S. phylicifolia despite the higher content of condensed tannins in young leaves. These results indicate that the general preference of leaf beetles for young leaves of Salicaceous plants probably does not primarily result from variable distribution of secondary compounds. Apparently, the preference for young leaves is fundamentally due to variation in leaf nutritive traits, such as nitrogen content. Received 9 February 2001.     

Semiochemicals mediating host-finding behaviour in the phoretic association between Macrocheles saceri (Acari: Mesostigmata) and Scarabaeus species (Coleoptera: Scarabaeidae)

Summary. Phoresy is an interspecific association where mobile organisms ensure small and/or wingless organisms to colonize patchy distributed environment. Macrocheles saceri (Acari: Mesostigmata) is a phoretic mite which specialized on Scarabaeus dung beetles (Coleoptera: Scarabaeidae). Variation for phoretic load and sites for mite fixation on beetle body has been tested according to Scarabaeus species. The number of setae on beetle body cannot explain the observed patterns in mite fixation. Evidence was provided that the semiochemicals of the host cuticle play a kairomonal role in the host finding behaviour of phoretic mites. In addition semiochemicals ensure host discrimination.     

Effects of carbaryl on the aminotransferases in the mid gut of blister beetle,Mylabris pustulata (Thunb)

The activity levels of aminotransferases such as alanine aminotransferases (AlAT), aspartate aminotransferases (AAT), isoleucine aminotranferases (IlAT), leucine aminotransferases (LAT), valine aminotrasferases (VAT) were measured in the mid gut of the blister beetle; Mylabris pustulata (Thunb) exposed to sublethal doses of carbaryl. The insecticide markedly inhibited the activity levels of all aminotransferases exposed for either short or prolonged treatments. The beetles regained normal enzymic activity after withdrawal from the treatment. It is possible that chronic exposures of beetles to carbaryl may induce impairment in protein synthesis in the mid gut of the beetle.     

Emission of oilseed rape volatiles after pollen beetle infestation; behavioural and electrophysiological responses in the parasitoid <Emphasis Type="Italic">Phradis morionellus</Emphasis>

Summary. The pollen beetle, Meligethes aeneus, is an important pest of oilseed rape, Brassica napus. Larvae of this species feed only in the buds and flowers of Brassicaceae. One important natural enemy of this beetle is the parasitoid Phradis morionellus that attacks larvae in buds and flowers and also feeds on the flowers. The preferences for odours of non-infested and infested rape were tested for both starved and fed parasitoids in Y-tube olfactometer experiments. The volatile blend released from pollen beetle-infested and non-infested flowering rape and from pollen beetle larvae was identified and quantified. Gas chromatography-electroantennodetection analyses were performed with female P. morionellus. Parasitoids in both treatment groups preferred infested rape, but the proportion of responding female P. morionellus was significantly lower for the group that was starved. Six of the 20 volatiles identified were released at higher rates from infested rape than from non-infested. None of these compounds was found in pollen beetle larvae headspace. P. morionellus antennae detected both major and minor components in the volatile blend. The volatiles released at a significantly higher rate from infested rape and detected by P. morionellus antennae were (Z)-3-hexenylacetate, (Z)-3-hexenol, 3-butenyl isothiocyanate and (E,E)-α-farnesene.     

Behavioral evidence for eavesdropping on prey song in two Palearctic sibling bat species

Eavesdropping on prey communication signals has never before been reported for a Palearctic bat species. In this study, we investigated whether lesser and greater mouse-eared bats, Myotis blythii oxygnathus and Myotis myotis, find tettigoniid bushcrickets (Tettigoniidae) by eavesdropping on their mate-attraction song. Tettigoniids are known to be the most important prey item for M. blythii oxygnathus, while carabid beetles and other epigaeic arthropods are the most important prey for its sibling species, M. myotis, in many places in Europe. M. myotis locates walking beetles by listening for their rustling sounds. We compared these two species’ response to four acoustic prey cues: calling song of two tettigoniid species, the rustling sound made by walking carabid beetles, and a control tone. Individuals of both bat species attacked the speaker playing tettigoniid song, which clearly indicates that both species eavesdrop on prey-generated advertisement signals. There were, however, species differences in response. M. blythii oxygnathus exhibited stronger predatory responses to the calling song of two species of tettigoniid than to the beetle rustling sound or the control. M. myotis, in contrast, exhibited stronger predatory responses to the beetle rustling and to one tettigoniid species but not the other tettigoniid or the control. Our study (1) for the first time demonstrates eavesdropping on prey communication signals for Palearctic bats and (2) gives preliminary evidence for sensory niche partitioning between these two sympatric sibling bat species.     

Effects of Postfire Salvage Logging on Deadwood‐Associated Beetles

Abstract: In Canada and the United States pressure to recoup financial costs of wildfire by harvesting burned timber is increasing, despite insufficient understanding of the ecological consequences of postfire salvage logging. We compared the species richness and composition of deadwood‐associated beetle assemblages among undisturbed, recently burned, logged, and salvage‐logged, boreal, mixed‐wood stands. Species richness was lowest in salvage‐logged stands, largely due to a negative effect of harvesting on the occurrence of wood‐ and bark‐boring species. In comparison with undisturbed stands, the combination of wildfire and logging in salvage‐logged stands had a greater effect on species composition than either disturbance alone. Strong differences in species composition among stand treatments were linked to differences in quantity and quality (e.g., decay stage) of coarse woody debris. We found that the effects of wildfire and logging on deadwood‐associated beetles were synergistic, such that the effects of postfire salvage logging could not be predicted reliably on the basis of data on either disturbance alone. Thus, increases in salvage logging of burned forests may have serious negative consequences for deadwood‐associated beetles and their ecological functions in early postfire successional forests.     

Experimental Test of Postfire Management in Pine Forests: Impact of Salvage Logging versus Partial Cutting and Nonintervention on Bird–Species Assemblages

Abstract: There is an intense debate about the effects of postfire salvage logging versus nonintervention policies on regeneration of forest communities, but scant information from experimental studies is available. We manipulated a burned forest area on a Mediterranean mountain to experimentally analyze the effect of salvage logging on bird–species abundance, diversity, and assemblage composition. We used a randomized block design with three plots of approximately 25 ha each, established along an elevational gradient in a recently burned area in Sierra Nevada Natural and National Park (southeastern Spain). Three replicates of three treatments differing in postfire burned wood management were established per plot: salvage logging, nonintervention, and an intermediate degree of intervention (felling and lopping most of the trees but leaving all the biomass). Starting 1 year after the fire, we used point sampling to monitor bird abundance in each treatment for 2 consecutive years during the breeding and winter seasons (720 censuses total). Postfire burned‐wood management altered species assemblages. Salvage logged areas had species typical of open‐ and early‐successional habitats. Bird species that inhabit forests were still present in the unsalvaged treatments even though trees were burned, but were almost absent in salvage‐logged areas. Indeed, the main dispersers of mid‐ and late‐successional shrubs and trees, such as thrushes (Turdus spp.) and the European Jay (Garrulus glandarius) were almost restricted to unsalvaged treatments. Salvage logging might thus hamper the natural regeneration of the forest through its impact on assemblages of bird species. Moreover, salvage logging reduced species abundance by 50% and richness by 40%, approximately. The highest diversity at the landscape level (gamma diversity) resulted from a combination of all treatments. Salvage logging may be positive for bird conservation if combined in a mosaic with other, less‐aggressive postfire management, but stand‐wide management with harvest operations has undesirable conservation effects.     

An Endangered Longhorn Beetle Associated with Old Oaks and Its Possible Role as an Ecosystem Engineer

Abstract:  For more than 10 years, ecologists have been discussing the concept of ecosystem engineering (i.e., nontrophic interactions of an organism that alters the physical state of its environment and affects other species). In conservation biology, the functional role of species is of interest because persistence of some species may be necessary for maintaining an entire assemblage with many threatened species. The great capricorn ( Cerambyx cerdo ), an endangered beetle listed in the European Union's Habitats Directive, has suffered a dramatic decline in the number of populations and in population sizes in Central Europe over the last century. The damage caused by C. cerdo larvae on sound oak trees has considerable effects on the physiological characteristics of these trees. We investigated the impacts of these effects on the species richness and heterogeneity of the saproxylic beetle assemblage on oaks. We compared the catches made with flight interception traps on 10 oaks colonized and 10 oaks uncolonized by C. cerdo in a study area in Lower Saxony (Germany). Our results revealed a significantly more species-rich assemblage on the trees colonized by C. cerdo . Colonized trees also harbored more red-listed beetle species. Our results suggest that an endangered beetle species can alter its own habitat to create favorable habitat conditions for other threatened beetle species. Efforts to preserve C. cerdo therefore have a positive effect on an entire assemblage of insects, including other highly endangered species. On the basis of the impact C. cerdo seems to have on the saproxylic beetle assemblage, reintroductions might be considered in regions where the species has become extinct.     

Pyrrolizidine alkaloids on three trophic levels – evidence for toxic and deterrent effects on phytophages and predators

Summary. Pyrrolizidine alkaloids (PAs) present a model system in the investigation of tritrophic interactions mediated by plant secondary compounds. However, their toxicity for insect herbivores has never been experimentally proven. Here, we demonstrate the toxic effects of a PA on growth and survival of the eri silk moth Philosamia ricini. In a feeding experiment, larvae of this generalist herbivore fed with an artificial PA diet gained weight significantly slower than control animals, and died as pupae. We suggest that derivatives of the ingested PA N-oxide damage developmental functions during metamorphosis. A tracer test with [¹⁴C]senecionine N-oxide revealed that the caterpillars lack adaptations that would prevent conversion of the chemical into the pro-toxic free base. In contrast, the PA adapted leaf beetle Longitarsus anchusae accumulates PAs as N-oxides. We tested the purpose of sequestration in this species as defence against predators. Through a series of prey choice experiments with three carabid predator species, chemically non-protected bark beetle pupae were chosen almost uniformly over L. anchusae pupae. In a following choice test with one of these predators, artificially PA-treated mealworm segments deterred the predator from feeding. Overall the study corroborates the immediate toxic effect of PAs on non-adapted herbivores and the protective effect that adapted insects may gain by sequestering them. It thereby underlines the potential for PAs to play a central role in multitrophic interactions between plants, phytophages and their predators.     

Effect of bombardier beetle spray on a wolf spider: repellency and leg autotomy

Summary. Data are presented on the repellency of the spray of a bombardier beetle (Pheropsophus aequinoctialis) to a lycosid spider (Lycosa ceratiola). The secretion is shown to cause the spider to desist from its assault on the beetle within, on average, 58 ms of onset of the beetle’s secretory emission, a reaction time that is at a par with latencies previously reported for startle, escape, and avoidance reactions of cockroaches, flies, and moths. Spray ejections by the beetle, are shorter in duration (43 ms, on average) than the response time of the spider, an indication that the beetle does indeed pack a formidable “punch” into its ejection. After being hit by a beetle’s spray, L. ceratiola were found occasionally to autotomize one or two of their legs. It is argued, but not proven, that this unusually severe effect from exposure to an arthropodan defensive secretion may be caused by the high temperature of the bombardier beetle spray.     

How maize root volatiles affect the efficacy of entomopathogenic nematodes in controlling the western corn rootworm?

Because the ferocious maize pest Diabrotica virgifera virgifera LeConte can adapt to all currently used control strategies, focus has turned to the development of novel, more sustainable control methods, such as biological control using entomopathogenic nematodes (EPN). A good understanding of the biology and behaviour of these potential control agents is essential for their successful deployment. Root systems of many maize varieties emit (E)-β-caryophyllene (EβC) in response to feeding by larvae of the beetle D. v. virgifera. This sesquiterpene has been shown to attract certain species of EPN, thereby enhancing their control potential. In this study, we tested the effect of this root-produced volatile on the field efficacy of the three EPN Heterorhabditis bacteriophora, Heterorhabditis megidis and Steinernema feltiae against D. v. virgifera larvae in southern Hungary. By comparing beetle emergence and root damage for two maize varieties, one that emits EβC and one that does not, it was found that root protection by H. megidis and S. feltiae was higher on the emitting variety, but this was not the case for H. bacteriophora. Overall, all three nematode species showed good control potential. We conclude that, if properly applied and in combination with the right maize variety, the release of these nematodes can be as effective as other control methods.     

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     

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     

Mother’s choice of the oviposition site: balancing risk of egg parasitism and need of food supply for the progeny with an infochemical shelter?

Summary. Oviposition site selection of herbivorous insects depends primarily on host plant presence which is essential for offspring survival. However, parasitoids can exploit host plant cues for host location. In this study, we hypothesised that herbivores can solve this dilemma by ovipositing within high plant diversity. A diverse plant species composition might represent an ‘infochemical shelter’, as a potentially complex volatile blend can negatively affect the host location ability of parasitoids. We examined this exemplarily for the egg-laying response of the generalist leaf beetle, Galeruca tanaceti, in relation to (1) host plant availability and (2) plant species diversity in the field. Further, we investigated the effect of odours from mixed plant species compositions on (3) leaf beetle oviposition site selection and on (4) the orientation of its specialised egg parasitoid, Oomyzus galerucivorus. In the field, egg clutch occurrence was positively related to the presence and quantity of two major host plants, Achillea millefolium (yarrow) and Centaurea jacea, and to the number of herbaceous plant species. In two-choice bioassays, female beetles oviposited more frequently on sites surrounded by an odour blend from a diverse plant species composition (including yarrow) than on sites with a pure grass odour blend. In the presence of yarrow odour and an odour blend from a diverse plant mixture (including yarrow) no difference in the oviposition response was recorded. Experienced parasitoid females were attracted to yarrow odours, but showed no response when yarrow odours were offered simultaneously with odours of a non-host plant. In conclusion, it could be shown in laboratory bioassays that the parasitoid responds only to pure host plant odours but not to complex odour blends. In contrast, the herbivore prefers to oviposit within diverse vegetation in the field and in the laboratory. However, the laboratory results also point to a priority of host plant availability over the selection of a potential ‘infochemical shelter’ for oviposition due to high plant diversity.     

Dynamics of pheromone production and communication in the mountain pine beetle, Dendroctonus ponderosae Hopkins, and the pine engraver, Ips pini (Say) (Coleoptera: Scolytidae)

Summary. The mountain pine beetle, Dendroctonus ponderosae Hopkins, and the pine engraver, Ips pini (Say), often co-exist in lodgepole pine, Pinus contorta var. latifolia Engelmann. Intra- and interspecific semiochemical communication occurs in both species and their complete semiochemical repertoire and precise dynamics of pheromone production have not been elucidated. Porapak-Q extracts of captured volatiles from beetles of each species aerated at different attack phases (freshly emerged, pioneer sex alone in the log and both sexes paired in new galleries), followed by gas chromatographic-electroantennographic detection (GC-EAD) and GC-mass spectroscopic analyses identified 17 compounds (seven compounds common to both species, six present in D. ponderosae and four present in I. pini) that excited the antennae of either or both species. Seven compounds for D. ponderosae and nine for I. pini had not been assessed for behavioural activity. In field trapping experiments, 2-phenylethanol produced by both species inhibited the response of D. ponderosae to its aggregation pheromones. exo- and endo-Brevicomin produced by D. ponderosae significantly decreased the response of I. pini to its aggregation pheromone ipsdienol. Nonanal, a ubiquitous compound found in the volatiles of lodgepole pine, various nonhosts and in both beetle species deterred the response of I. pini to ipsdienol. The occurrence of cis-verbenol, trans-verbenol and verbenone in emergent I. pini, and verbenone and 2-phenylethanol in emergent D. ponderosae suggests that these compounds may inhibit aggregation and induce dispersal following emergence. Termination of aggregation in D. ponderosae appears to depend on the production of frontalin in combination with changes in the relative ratios of verbenone, exo-brevicomin, trans-verbenol and 2-phenylethanol. In I. pini, the cessation of ipsdienol production by males is probably the main factor in terminating aggregation. Received 16 November 1999; accepted 7 August 2000     

Three beetles—three concepts. Different defensive strategies of congeneric myrmecophilous beetles

Myrmecophiles, i.e., organisms associated with ants live in a variety of ecological niches in the vicinity or inside ant colonies and employ different strategies to survive ant encounters. Because different niches are characterized by different encounter rates with host ants, strategies used to avoid ant aggressions should depend on these niches. This hypothesis was studied with three rove beetle species of the genus Pella, which are myrmecophiles of the ant Lasius fuliginosus and the non-myrmecophilous relative Drusilla canaliculata. Behavioral tests in the field revealed that Pella species are better adapted to interactions with ants than D. canaliculata, but that they use species-specific strategies in ant interactions. Pella cognata and Pella funesta avoid encounters with ants by swift movements. Chemical analyses of the defensive tergal gland secretions showed that P. cognata has replaced the aggression inducing undecane by the behaviorally neutral tridecane. P. funesta repels the ants by releasing the panic alarm pheromone sulcatone from its tergal gland resulting in an “ant free space” around the beetles. Finally, Pella laticollis uses a specific and unique appeasing behavior. Behavioral and chemical data did not reveal any indication for the mimicry of the ants' cuticular hydrocarbon profiles by any of the beetle species. It is discussed that the employed strategies correlate with the ecological niches of the beetles. P. cognata and P. funesta are living along ant trails with ample space to escape and the employed strategies are probably sufficient to escape from dangerous conflicts. In contrast, P. laticollis lives in refuse areas of ant nests with frequent ant encounters, and its appeasement strategy allows it to stay at the encounter site.     

Understanding Trait‐Dependent Community Disassembly: Dung Beetles,Density Functions,and Forest Fragmentation

Abstract: Anthropogenic disturbances such as fragmentation are rapidly altering biodiversity, yet a lack of attention to species traits and abundance patterns has made the results of most studies difficult to generalize. We determined traits of extinction‐prone species and present a novel strategy for classifying species according to their population‐level response to a gradient of disturbance intensity. We examined the effects of forest fragmentation on dung beetle communities in an archipelago of 33 islands recently created by flooding in Venezuela. Species richness, density, and biomass all declined sharply with decreasing island area and increasing island isolation. Species richness was highly nested, indicating that local extinctions occurred nonrandomly. The most sensitive dung beetle species appeared to require at least 85 ha of forest, more than many large vertebrates. Extinction‐prone species were either large‐bodied, forest specialists, or uncommon. These explanatory variables were unrelated, suggesting at least 3 underlying causes of extirpation. Large species showed high wing loading (body mass/wing area) and a distinct flight strategy that may increase their area requirements. Although forest specificity made most species sensitive to fragmentation, a few persistent habitat generalists dispersed across the matrix. Density functions classified species into 4 response groups on the basis of their change in density with decreasing species richness. Sensitive and persistent species both declined with increasing fragmentation intensity, but persistent species occurred on more islands, which may be due to their higher baseline densities. Compensatory species increased in abundance following the initial loss of sensitive species, but rapidly declined with increasing fragmentation. Supertramp species (widespread habitat generalists) may be poor competitors but strong dispersers; their abundance peaked following the decline of the other 3 groups. Nevertheless, even the least sensitive species were extirpated or rare on the smallest and most isolated islands.