Predator interference limits fly egg biological control by a guild of ground-active beetles

We examined whether predator interference could prevent effective conservation biological control of Delia spp. flies, important pests of cole crops, by an assemblage of carabid and staphylinid beetles. In laboratory feeding trials we found that the smaller (<1 cm) beetle species common at our site readily ate dipteran eggs, while the most common large carabid species, Pterostichus melanarius, rarely did. However, P. melanarius did eat several of the smaller beetle species. We conducted two field experiments where we manipulated immigration rates of the ground predator guild and then measured predation on fly eggs. Predation rates were consistently higher in cages where predators were added at ambient densities, compared to cages where ground predators were removed. However, in the second field experiment, when we quadrupled predator immigration rates neither beetle activity-density nor predation rate increased. High immigration rate plots had a higher proportion of P. melanarius in the predator community, compared to plots with beetles added at ambient densities, suggesting that P. melanarius was reducing activity-densities of the smaller beetles, perhaps through intraguild predation. Thus, tactics to improve the biological control of Delia spp. by conserving generalist predators, such as providing in- or extra-field refuges, could be thwarted if the primary predators of fly eggs, small carabids and staphylinids, are the targets of intraguild predation by also-conserved larger predators.     

Phoretic Poecilochirus mites specialize on their burying beetle hosts

Recurring species interactions can cause species to adapt to each other. Specialization will increase the fitness of symbionts in the coevolved association but may reduce the flexibility of symbiont choice as it will often decrease fitness in interactions with other than the main symbiont species. We analyzed the fitness interactions between a complex of two cryptic mite species and their sympatric burying beetle hosts in a European population. Poecilochirus mites (Mesostigmata, Parasitidae) are phoretic on burying beetles and reproduce alongside beetles, while these care for their offspring at vertebrate carcasses. While Poecilochirus carabi is typically found on Nicrophorus vespilloides beetles, P. necrophori is associated with N. vespillo. It has long been known that the mites discriminate between the two beetle species, but the fitness consequences of this choice remained unknown. We experimentally associated both mite species with both beetle species and found that mite fitness suffered when mites reproduced alongside a nonpreferred host. In turn, there is evidence that one of the beetle species is better able to cope with the mite species they are typically associated with. The overall fitness effect of mites on beetles was negative in our laboratory experiments. The Poecilochirus mites studied here are thus specialized competitors or parasites of burying beetles.     

The rove beetle Drusilla sparsa (Coleoptera: Staphylinidae) is a myrmecophilous species associated with a myrmicine ant,Crematogaster osakensis (Hymenoptera: Formicidae)

The rove beetle genus Drusilla includes some myrmecophilous species. The Japanese species Drusilla sparsa (Sharp, 1874) has been regarded as a non‐myrmecophilous beetle. In Kagawa Prefecture, Shikoku Island, western Japan, however, we often observed that D. sparsa adults were walking in the vicinity of foraging workers of the myrmicine ant Crematogaster osakensis Forel, 1990. The body color of the beetle is similar to C. osakensis as in other myrmecophilous beetles found near the trails of the host ants. To examine whether D. sparsa is myrmecophilous, we investigated the distribution of D. sparsa and C. osakensis in the field, as well as their behavior including prey preference of the beetle in the laboratory. Drusilla sparsa beetles were collected only in sites where C. osakensis ants occurred. When the beetles encountered the ant workers, they bent the abdominal tip toward the ants. The ants licked the abdominal tip, and then the beetles usually walked away. Such behavioral reaction of the ants was not observed when the beetles encountered workers of the formicine ant Nylanderia flavipes (Smith, 1874) that continuously attacked the beetles. Drusilla sparsa preferred to feed on dead workers of C. osakensis even when other ants were available as food, indicating that D. sparsa is a myrmecophilous species associated with C. osakensis. Crematogaster osakensis was frequently found in the stomach in the ant predator, the Japanese treefrog Hyla japonica Günther, 1859. Thus, the significance of body color similarity between the host ants and beetles is not a case of Batesian mimicry.     

Emergence patterns of univoltine and bivoltine Ips typographus (L.) populations and associated natural enemies

Control measures aiming at reducing bark beetle populations and preserving their natural enemies require a sound knowledge on their overwintering and emergence behaviour. These behavioural traits were investigated in univoltine and bivoltine populations of the European spruce bark beetle (Ips typographus [L.], Coleoptera: Scolytinae) and its predators and parasitoids over several consecutive years. In univoltine populations, roughly 50% of the bark beetles left their brood trees in fall together with most parasitoids and some significant predatory flies and beetles. In bivoltine populations, <10% of the second bark beetle generation emerged in fall and the remainder overwintered under the bark of their brood trees. Likewise, most predatory beetles and flies spent wintertime with their prey under the bark, while most parasitic wasps emerged in fall. The spring emergence of bivoltine predatory beetles was found to occur up to 3 weeks earlier than that of I. typographus, while that of the predatory flies and the parasitoids was delayed by up to 1 month. In univoltine populations, the bark beetles emerged several weeks prior to most antagonistic taxa. In the heat year 2003, three I. typographus generations were produced at the lower location, 36% of the third generation emerged in fall, while the proportions of overwintering predators remained largely the same as in previous years. Similar to their host, more parasitoids left their brood trees in fall after warm years. The results show that sanitation felling during winter probably kills most bark beetles in bivoltine populations, but also eliminates many natural enemies. In univoltine populations, sanitation felling might be less detrimental to both I. typographus and natural enemies because a fair fraction of their populations will already have left the trees before cutting. Warmer climates may affect the interactions of bark beetles and natural enemies and thus the impact of control measures.     

Attraction of a kleptoparasitic sphaerocerid fly (Norrbomia frigipennis) to dung beetles (Phanaeus spp. andCanthon sp.)

Norrbomia frigipennis (Diptera: Sphaeroceridae) is phoretic on dung-feeding scarab beetles (Coleoptera: Scarabaeidae). In this study we investigate the attractiveness of three beetle species,Phanaeus ignius, P. vindex, andCanthon pilularis, to the fly. Stationary, moving-dead, and live beetles were used. More flies were attracted toPhanaeus. However, this attractiveness may be due to the larger average size ofPhanaeus. A preference for larger individuals was found withinPhanaeus, though not withinC. pilularis. Flies mounted beetles on the thorax and the elytra at similar rates.Phanaeus males that possesed horns did not attract more flies than did hornless ones, and there was no effect of host sex on attractiveness. In hornedPhanaeus, about 11–16% of the flies mounting those beetles landed on the horn.     

Social encapsulation of the small hive beetle (<Emphasis Type="Italic">Aethina tumida</Emphasis> Murray) by European honeybees (<Emphasis Type="Italic">Apis mellifera</Emphasis> L.)

Summary European and African subspecies of honeybees (Apis mellifera L.) utilize social encapsulation to contain the small hive beetle (Aethina tumida Murray), a honeybee colony scavenger. Using social encapsulation, African honeybees successfully limit beetle reproduction that can devastate host colonies. In sharp contrast, European honeybees often fail to contain beetles, possibly because their social encapsulation skills may be less developed than those of African honeybees. In this study, we quantify beetle and European honeybee behaviours associated with social encapsulation, describe colony and time (morning and evening) differences in these behaviours (to identify possible circadian rhythms), and detail intra-colonial, encapsulated beetle distributions. The data help explain the susceptibility of European honeybees to depredation by small hive beetles. There were significant colony differences in a number of social encapsulation behaviours (the number of beetle prisons and beetles per prison, and the proportion of prison guard bees biting at encapsulated beetles) suggesting that successful encapsulation of beetles by European bees varies between colonies. We also found evidence for the existence of circadian rhythms in small hive beetles, as they were more active in the evening rather than morning. In response to increased beetle activity during the evening, there was an increase in the number of prison guard bees during evening. Additionally, the bees successfully kept most (~93%) beetles out of the combs at all times, suggesting that social encapsulation by European honeybees is sufficient to control small populations of beetles (as seen in this study) but may ultimately fail if beetle populations are high.Received 20 January 2003; revised 21 April 2003; accepted 29 April 2003.     

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.     

Geographic variation for colour in the sandburrowing beetle Chaerodes trachyscelides White (Goleoptera: Tenebrionidae) on New Zealand beaches analysed using CIELAB L * values

Chaerodes trachyscelides White is a highly specialized, flightless burrowing beetle confined to the narrow strip of sand at and just above high water level on sandy marine beaches in New Zealand. Although the ventral surface of the beetle is always pale, the dorsal surface varies from pale to almost black. Large samples of this beetle were taken, together with the sand, from 11 beaches on New Zealand's three main islands. The colour of the dorsal surface of each individual beetle and that of the sand samples was measured using reflectance spectroscopy and expressed as CIE L*,a*,b* (CIELAB) values. The L* values, which are objective, quantitative measures of the degree of lightness of the beetles, were subjected to statistical and frequency analysis. Although the species was very variable in colour and the variation appeared to be continuous, a highly significant correlation was obtained between the mean of the L* values for the samples on each beach and that of the sand, the correlation coefficient being 0.961. This close association between the lightness of the beetles and that of the sand suggests the variable melanism functions as cryptic colouration. On most beaches, the distribution oflightness among the beetles sampled conformed to a normal curve. For beetles from sites where the sand was relatively uniform, such as the black Taranaki beaches, the L* frequency distribution curves were narrow and the coefficient of variation of mean beetle colour was relatively small indicating low colour variability. In contrast, the greatest within-site variability occurred on the two Stewart Island beaches sampled, where in each case there was less uniformity in the colour of the sand. At one of these sites, Maori Beach, darker sand present below the high water level is often deposited on the zone occupied by the beetles after storms. On Lonneker's Beach, the distribution of L* values among the beetles sampled was actually bimodal. On this small beach, there was an area of intensely black sand in the zone occupied by the beetles, but most of the rest was covered with light golden sand. These results are interpreted as evidence that the variability of colour of Chaerodes beetles has the effect of populations being able to match the colour of the sand of their home beaches, presumably as a consequence of the differential survival of individuals.     

Overwintering biology ofMyiopharus aberrans andMyiopharus doryphorae (Dipt.: Tachinidae) larval parasitoids of the colorado potato beetle (Col.: Chrysomelidae)

The overwintering biology ofMyiopharus aberrans (Townsend) andMyiopharus doryphorae (Riley), tachinid parasitoids of the Colorado potato beetle, was investigated. Sixty seven overwintered Colorado potato beetle adults were dug from the soil before beetle emergence in the spring of 1990 from a field that had been planted to potatoes the previous year. Five adultM. aberrans and two adultM. doryphorae flies were reared from seven beetles. This is the first record of howM. aberrans andM. doryphorae flies overwinter.      

Sampling indigenous ground‐living beetles in a stand of non‐native tree privet (Ligustrum lucidum) in New Zealand raises new management questions

Small urban forest reserves in New Zealand have been shown to have value in conserving indigenous beetle diversity. However there is little information available on the ability of non‐native vegetation areas such as tree privet to support indigenous beetle assemblages. To investigate this for one site, ground‐living beetles were collected using pitfall traps over a year at a small urban forest of the invasive tree Ligustrum lucidum (tree privet) in Auckland, New Zealand. A total of 815 beetles were found, from 20 families and 42 relative taxonomic units. Using monthly data, there was no correlation between soil moisture and diversity index (P = 0.805) or species richness (P = 0.375). These results raise the question of whether urban patches of non‐native tree privet may have potential as reservoirs of beetle diversity, if only until they are replaced with native vegetation.     

The dispersal behaviour of the phoretic mite Poecilochirus carabi (Mesostigmata,Parasitidae): adaptation to the breeding biology of its carrier Necrophorus vespilloides (Coleoptera,Silphidae)

Summary When the phoretic mite Poecilochirus carabi reproduces in the brood chamber of its carrier Necrophorus vespilloides, a beetle with biparental brood care, the first deuteronymphs of the new mite generation aggregate on the male beetle. They do not use sex-specific traits to discriminate between male and female beetles in the brood chamber, but traits that are related to the beetles' behaviour and may be displayed by both parent beetles. When the male beetle departs, it carries virtually all deuteronymphs then present in the brood chamber. Deuteronymphs that develop later congregate on the female, which leaves the crypt some days after the male. Only those deuteronymphs that miss the female's departure disperse on the beetle larvae, meaning they have to wait in their pupal chambers until the beetles have completed their development. On average, 86% of the deuteronymphs leave the brood chamber on the parent beetles, thereby gaining the advantage of an early departure. As soon as their carrier arrives at one of the beetles' meeting places, the deuteronymphs can transfer between the beetles present. Choice experiments revealed that the deuteronymphs tend to even out density differences between congregating carriers, and prefer sexually mature to immature beetles. Therefore, transferring between beetles results in a dispersion of deuteronymphs on the sexually mature beetles of the population.     

Host-associated differences in fitness within and between populations of a seed beetle (Bruchidae): effects of plant variability

Summary Experiments were conducted with the sexually reproducing seed beetle Stator limbatus and its hosts in north-central Arizona to determine if it was substructured into units, each specialized for higher fitness on a specific host species. Unlike many studies, we incorporated scale, i.e., conducting experiments between and within beetle populations on seeds from within and between plant species. Of particular interest was whether intraspecific plant variability prevented beetle specialization within beetle populations. Results suggest that S. limbatus is specialized to certain hosts. On the palo verde Cercidium floridum, beetles originally reared from this host had significantly higher emergence compared to beetles transferred from other hosts. We did not test directly for a genetic basis for this. Alternative hypotheses of variation in symbiotic microorganisms in larval guts and maternal effects were assessed. Essentially no bacteria, yeast or protozoa were found, and maternal effects as expressed by varying egg weights were not detected; however, other microorganisms might have been present and maternal effects through inducible enzymes was possible. Caution, then, is needed in any genetic interpretations of our results. The differences on C. floridum were detected from tests between and within beetle populations. Evidence for specialization was not detected on the other hosts, Cercidium microphyllum and Acacia greggii. On the other hosts, beetles performed well regardless of their source. Significant differences were detected among individual plants of C. floridum as to the suitability of their seeds for deveoopment of S. limbatus. No such differences were detected among the other host plants. These patterns of conspecific plant variability are opposite of what is expected if plant variability prevents specialization of beetles to particular species of hosts. Thus, the data suggest seed variability among plants does not prevent specialization to host species in this system. We discuss how the patterns of host use in this study relate to the hypothesis of sympatric host race formation.     

Flowers and Beetles in the South American Tropics

Examples of beetle pollination in the South American tropics are given. The case histories refer to representatives of the families Cyclanthaceae, Annonaceae, and Araceae. In most cases the cantharophilous syndrome does not seem to be the result of coevolution but of a one-sided adaptation of flowers to the behavior, necessities, and sense-capacities of beetle species. It is emphasized that cantharophily may not necessarily be an indication of ancientness of a flower/insect relationship. Besides primitive angiosperms, there are derived and modern groups which, in their pollination, are specialized for beetles. Carludovica palmata (Cyclanthaceae) is pollinated by small curculionids which are attracted by odors in the early morning hours. Annona coriacea (Annonaceae) and Philodendron selloum (Araceae) are pollinated by large dynastid scarab beetles in the evening hours. In all three species thermogenesis occurs. Although belonging to widely separate systematic groups, Annona flowers and Philodendron inflorescences have a similar pollination biology, which indicates that their adaptations to beetles are the result of convergent evolutionary developments.     

Confinement Behavior of Cape Honey Bees (Apis mellifera capensis Esch.) in Relation to Population Densities of Small Hive Beetles (Aethina tumida Murray)

We quantified the effects of increasing small hive beetle (Aethina tumida Murray) populations on guarding behavior of Cape honey bees (Apis mellifera capensis, an African subspecies). We found more confinement sites (prisons) at the higher (50 beetles per colony) rather than lower (25 beetles per colony) beetle density. The number of beetles per prison did not change with beetle density. There were more guard bees per beetle during evening than morning. Neither guard bee nor beetle behavior varied with beetle density or over time. Forty-six percent of all beetles were found among the combs at the low beetle density and this increased to 58% at the higher one. In neither instance were beetles causing depredation to host colonies. Within the limits of the experiment, guarding behavior of Cape honey bees is relatively unaffected by increasing beetle density (even if significant proportions of beetles reach the combs).     

Delaying insect access alters community composition on small carrion: a quantitative approach

We present a study of interactions in the highly competitive insect communities inhabiting the carrion of small mammals. Via manipulation in a fully quantitative design, we delayed community development by excluding insect colonization in mouse and rat carcasses for 3 days, to study the role of early competitively dominant colonizers [burying beetles (Nicrophorus spp.; Coleoptera: Silphidae) and blowfly larvae (Diptera: Calliphoridae)] in the course of heterotrophic succession on small cadavers. Earlier studies demonstrated that in the case of large mammalian carrion, exclusion of insects’ access to the carcass in the early stages of decomposition altered the successional trajectory and species assemblages. However, the effect of such manipulation in easy monopolizable small vertebrate carrion remained unknown. Our results demonstrate that delaying insect access to carrion significantly lowered blowfly larvae abundances, while it simultaneously had no effect on colonization and carrion burial by burying beetles. Higher abundances of blowfly larvae seem to deter necrophagous beetles, whereas they are not harmful to the larvae of flesh flies, at least in larger rat carcasses. Predatory beetle species preferred the lower abundances of blowfly larvae, presumably due to better accessibility of their prey. Our results therefore suggest that in the course of the entire season, larvae of blowflies are the dominant competitors in small carcasses, and significantly affect the assembly of other insect groups, whereas burying beetles may exhibit a more temporal pattern of dominance.     

Indirect effects of a non-target species, Pyrrhalta luteola (Chrysomelidae) on the biodeterioration of Brussels tapestries

Treatments to preserve artwork are usually designed to control for known target species. Nevertheless, indirect effects of non-target species may cause important biodeterioration. During the restoration of a series of Brussels tapestries preserved in the Abbey of Sacromonte (Granada, Spain), we studied one of the tapestries which presented a high degree of deterioration, principally large losses of material and color alterations. We found important biodeterioration effects produced by a non-target insect species, the beetle Pyrrhalta luteola (Chrysomelidae). We raised P. luteola beetles in the laboratory on different constituent materials to identify the effects of this insect on the tapestry. Results showed that beetle remains were mainly responsible for the chromatic changes. Although beetles did not consume textile materials, and had not direct effects in tapestry destruction, its presence indirectly affected the tapestry due to the higher mold-growth obtained in samples of materials with beetle remains. Based on these results, we propose some recommendations to consider control for these potential biodeterioration agents in artwork conservation programs.     

Effects of different tree harvesting intensities on saproxylic beetle diversity in coniferous and broad-leaved mixed forest in the Changbai Mountain,Northeast, China

1. Saproxylic beetles have gained increasing attention due to their role in the decomposition of rotting wood in forests. Studying the response of saproxylic beetles to tree harvesting is important for developing harvesting strategies that consider conservation of saproxylic beetle diversity.
2. We report results from a case study in which we designed four treatment stands to test the effects of forest harvesting intensity on saproxylic beetle diversity, harvest intensities of 0% untreated control (CK), 17.2% light harvest (LT), 34.7% moderate harvest (MT) and 51.9% high harvest (HT). Flight intercept traps were used to collect specimens of saproxylic beetles in each stand at 2, 3 and 4 years post-harvest.
3. The richness and abundance of saproxylic beetles were higher in MT and HT than in CK. Twelve of the 15 indicator species were significantly associated with MT or HT, whereas only three species were significantly associated with CK.
4. We found that moderate and high intensity harvesting affected the composition and increased beetle abundance and richness of saproxylic beetles, and light intensity harvesting had no effect on the beetle community in years 2, 3 and 4 post-harvest. There was no difference in the beetle community composition between moderate and high harvest stands.

     

Passive aerodynamic stabilization by beetle elytra (wing covers)

Summary. Most beetles extend their elytra laterally during flight, where they can act as fixed airfoils; typically, such elytra are held at a pronounced dihedral angle. We fixed extended elytra of June beetles (Scarabaeidae: Melolonthinae) and tiger beetles (Cicindelidae) over a range of dihedral angles (-10° to 45°) and towed them in a water tunnel. We also towed them with flexed elytra (i.e. in the resting position). (Reynolds number was used to maintain dynamic similarity with air.) We measured the maximum rolling angle and the lateral oscillation rate as two indices of stability. There was no relationship between oscillation rate and dihedral angle, or between maximum roll angle and dihedral angle. At all dihedral angles tested, however, beetles were much more stable than when they were towed with the elytra flexed. We also included a third beetle type, flower beetle (Scarabaeidae: Cetoniine), which normally flies with flexed elytra. We measured stability indices for these beetles with elytra only in the flexed position. When all beetles were tested with flexed elytra, flower beetles showed no difference in maximum roll angle, but more stability in oscillation rate than the other two beetle types.     

Surviving submerged—Setal tracheal gills for gas exchange in adult rheophilic diving beetles

The gas exchange in adult diving beetles (Coleoptera: Dytiscidae) relies on a subelytral air store, which has to be renewed in regular intervals at the water surface. The dive duration varies from a few minutes to 24 h depending on the species, activity, and temperature. However, some species remain submerged for several weeks. Stygobiont species do not ascend to the surface and gas exchange of these species remains unclear, but it is assumed that they require air filled voids for respiration or they use cutaneous respiration. In this study, we investigate the gas exchange in the running water diving beetle Deronectes aubei, which survive submerged for over 6 weeks. The diffusion distance through the cuticle is too great for cutaneous respiration. Therefore, the dissolved oxygen uptake of submerged beetles was determined and an oxygen uptake via the rich tracheated elytra was observed. Fine structure analyses (SEM and TEM) of the beetles showed tracheated setae mainly on the elytral surface, which acts as tracheal gills. Prevention of the air bubble formation at the tip of the abdomen, which normally act as physical gill in Dytiscidae, resulted in no effect in oxygen uptake in D. aubei, but this was the sole way for a submerged Hydroporus palustris to get oxygen. The setal gas exchange technique explains the restriction of D. aubei to rivers and brooks with high oxygen concentration and it may also be used by subterran living diving beetles, which lack access to atmospheric oxygen. The existence of setal tracheal gills in species in running water which are often found in the hyporheic zone and in stygobiont species supports the known evolution of stygobiont Dytiscidae from species of the hyporheic zone. For species in running water, setal tracheal gills could be seen as an adaptation to avoid drifting downstream by the current. J. Morphol., 2009. © 2009 Wiley‐Liss, Inc.     

Influence ofOnthophagus gazella on hornfly,Haematobia irritans density in irrigated pastures

Haematobia irritans (L.) breeding in flood irrigated pastures of the lower Colorado Desert of southeastern California continues to remain unacceptably high during warm seasons (>1,000 adult flies per bovine head) despite the presence of moderately abundant populations ofOnthophagus gazella F. This study suggests that densities of > 40–70 adult beetles per dung pad and giving pronounced dung shredding activity, caused fly mortality of 38–56 %. The continued high abundance of adult horn flies on cattle suggests that at > 50% mortality, the pasture environment still produces sufficient flies to saturate cattle, although emigration might be reduced. Additional species of scarabs may be necessary to increase fly mortality. However, the dung drying activity of existingO. gazella significantly could interfere with resident staphylinid beetle breeding, which was significantly lower in pastures whereO. gazella reached densities of 40 per dung pad. Scarab beetle activity might also impede the introduction of superior predatory species for biological control.