Michael D. Ulyshen

A comparison of the Beetle (Coleoptera) Fauna Captured at two heights above the ground in a North American temperate deciduous forest

ABSTRACT We compared the beetle fauna captured in 12 pairs of flight intercept traps suspended at two different heights above the ground (≥15 m and 0.5 m) in a temperate deciduous forest in the southeastern United States to better understand how the abundance, species richness, diversity and composition of insect communities differ among forest strata. A total of 15,012 beetle specimens were collected representing 73 families and 558 morphospecies. Shannons diversity and evenness were both higher near the ground than in the canopy, but no differences in total abundance or species richness between the two layers were observed. There were many differences at the family level, however, and species composition differed considerably between the two layers. About 29% and 31% of species were captured exclusively in the canopy or near the ground, respectively. The canopy traps were more similar to one another than they were to those near the ground and vice-versa based on both Sorensens quotient of similarity and Raabes percentage of similarity. The canopy and ground trap locations were quite distinct based on nonmetric multidimensional scaling. The degree to which species composition was similar (i.e., Sorensens quotient of similarity) between pairs of traps decreased significantly with inter-trap distance for the traps in the canopy, but not for those near the ground, suggesting a more uniform community near the ground. Of the 41 families or subfamilies represented by more than 40 individuals, 12 were more abundant in the canopy and 14 were more abundant near the ground. Similarly, of the 16 families or subfamilies represented by more than 10 species, five were more species rich in the canopy and four were more species rich near the ground. Three families (Cerambycidae, Cleridae and Coccinellidae) were both more abundant and species rich in the canopy, whereas four other groups (Carabidae, Pselaphinae (Staphylinidae), Scolytinae (Curculionidae) and other Staphylinidae) were both more abundant and species rich near the ground. In addition to differing considerably among families, the vertical distribution patterns varied within many families as well. The distribution patterns for several groups are discussed in some detail with respect to known life history information.

Wood decomposition as influenced by invertebrates

The diversity and habitat requirements of invertebrates associated with dead wood have been the subjects of hundreds of studies in recent years but we still know very little about the ecological or economic importance of these organisms. The purpose of this review is to examine whether, how and to what extent invertebrates affect wood decomposition in terrestrial ecosystems. Three broad conclusions can be reached from the available literature. First, wood decomposition is largely driven by microbial activity but invertebrates also play a significant role in both temperate and tropical environments. Primary mechanisms include enzymatic digestion (involving both endogenous enzymes and those produced by endo‐ and ectosymbionts), substrate alteration (tunnelling and fragmentation), biotic interactions and nitrogen fertilization (i.e. promoting nitrogen fixation by endosymbiotic and free‐living bacteria). Second, the effects of individual invertebrate taxa or functional groups can be accelerative or inhibitory but the cumulative effect of the entire community is generally to accelerate wood decomposition, at least during the early stages of the process (most studies are limited to the first 2–3 years). Although methodological differences and design limitations preclude meta‐analysis, studies aimed at quantifying the contributions of invertebrates to wood decomposition commonly attribute 10–20% of wood loss to these organisms. Finally, some taxa appear to be particularly influential with respect to promoting wood decomposition. These include large wood‐boring beetles (Coleoptera) and termites (Termitoidae), especially fungus‐farming macrotermitines. The presence or absence of these species may be more consequential than species richness and the influence of invertebrates is likely to vary biogeographically.

Microclimate and habitat heterogeneity as the major drivers of beetle diversity in dead wood

Summary Resource availability and habitat heterogeneity are principle drivers of biodiversity, but their individual roles often remain unclear since both factors are usually correlated. The biodiversity of species dependent on dead wood could be driven by either resource availability represented by dead-wood amount or habitat heterogeneity characterized by dead-wood diversity or both. Understanding their roles is crucial for improving evidence-based conservation strategies for saproxylic species in managed forests. To disentangle the effects of dead-wood amount and dead-wood diversity on biodiversity relative to canopy openness (microclimate), we experimentally exposed different amounts of logs and branches of two different tree species representing a gradient of dead-wood diversity in 190 sunny and shady forest plots. During the 3 years after exposing dead wood, we sampled saproxylic beetles, which are together with fungi the most diverse and important taxonomic group involved in decomposition of wood. The composition of saproxylic beetle assemblages differed clearly between shady and sunny forest plots, with higher richness in sunny plots. Both dead-wood amount and dead-wood diversity positively and independently affected species richness of saproxylic beetles, but these effects were mediated by canopy openness. In sunny forest, species richness increased with increasing amount of dead wood, whereas in shady forest, dead-wood diversity was the prevailing factor. The stepwise analysis of abundance and species richness, however, indicated that effects of both factors supported only the habitat-heterogeneity hypothesis, as the positive effect of high amounts of dead wood could be explained by cryptic variability of dead-wood quality within single objects. Synthesis and applications. As canopy openness and habitat heterogeneity seem to be the major drivers of saproxylic beetle diversity in temperate forests, we recommend that managers aim to increase the heterogeneity of dead-wood substrates under both sunny and shady forest conditions. Intentional opening of the canopy should be considered in anthropogenically homogenized, dense forests. Specifically in temperate mixed montane forests, dead wood should be provided in the form of large logs in sunny habitats and a high diversity of different dead-wood substrates should be retained or created in shady forests.

Responses of Arthropods to large scale manipulations of dead wood in loblolly pine stands of the southeastern United States

ABSTRACT Large-scale experimental manipulations of dead wood are needed to better understand its importance to animal communities in managed forests. In this experiment, we compared the abundance, species richness, diversity, and composition of arthropods in 9.3-ha plots in which either (1) all coarse woody debris was removed, (2) a large number of logs were added, (3) a large number of snags were added, or (4) no coarse woody debris was added or removed. The target taxa were ground-dwelling arthropods, sampled by pitfall traps, and saproxylic beetles (i.e., dependent on dead wood), sampled by flight intercept traps and emergence traps. There were no differences in total ground-dwelling arthropod abundance, richness, diversity, or composition among treatments. Only the results for ground beetles (Carabidae), which were more species rich and diverse in log input plots, supported our prediction that ground-dwelling arthropods would benefit from additions of dead wood. There were also no differences in saproxylic beetle abundance, richness, diversity, or composition among treatments. The findings from this study are encouraging in that arthropods seem less sensitive than expected to manipulations of dead wood in managed pine forests of the southeastern United States. Based on our results, we cannot recommend inputting large amounts of dead wood for conservation purposes, given the expense of such measures. However, the persistence of saproxylic beetles requires that an adequate amount of dead wood is available in the landscape, and we recommend that dead wood be retained whenever possible in managed pine forests.

Population dynamics of an invasive forest insect and associated natural enemies in the aftermath of invasion: implications for biological control

Summary 1. Understanding the population dynamics of exotic pests and associated natural enemies is important in developing sound management strategies in invaded forest ecosystems. The emerald ash borer (EAB) Agrilus planipennis Fairmaire is an invasive phloem-feeding beetle that has killed tens of millions of ash Fraxinus trees in North America since first detected in 2002. 2. We evaluated populations of immature EAB life stages and associated natural enemies over a 7-year period (2008–2014) in six stands of eastern deciduous forest in southern Michigan, where Tetrastichus planipennisi Yang and two other Asian-origin EAB parasitoids were released for biological control between 2007 and 2010. 3. We observed 90% decline in densities of live EAB larvae in infested ash trees at both parasitoid-release and control plots from 2009 to 2014 and found no significant differences in EAB density or mortality rates by parasitoids, avian predators or other undetermined factors between parasitoid-release and control plots. The decline in EAB larval density in our study sites was correlated with significant increases in EAB larval parasitism, first by native parasitoids, then by T. planipennisi. 4. Life table analyses further indicated that parasitism by the introduced biocontrol agent and the North American native parasitoids contributed significantly to the reduction of net EAB population growth rates in our study sites from 2010 to 2014. 5. Synthesis and applications. Our findings indicate that successful biocontrol of emerald ash borer (EAB) may involve suppression of EAB abundance both by local, generalist natural enemies (such as Atanycolus spp.) and by introduced specialist parasitoids (such as T. planipennisi). Biological control programmes against EAB in the aftermath of invasion should focus on establishing stable populations of T. planipennisi and other introduced specialist parasitoids for sustained suppression of low-density EAB populations. Moreover, we recommend releasing the introduced specialist biocontrol agents as soon as possible to prevent the outbreak of EAB populations in both newly infested and aftermath forests when EAB densities are still low.

On the vertical distribution of bees in a temperate deciduous forest

Abstract.  1. Despite a growing interest in forest canopy biology, very few studies have examined the vertical distribution of forest bees. In this study, bees were sampled using 12 pairs of flight‐intercept traps suspended in the canopy (≥15 m) and near the ground (0.5 m) in a bottomland hardwood forest in the southeastern United States.

Suitability and Accessibility of Immature Agrilus planipennis (Coleoptera: Buprestidae) Stages to Tetrastichus planipennisi (Hymenoptera: Eulophidae)

ABSTRACT Tetrastichus planipennisi Yang (Hymenoptera: Eulophidae), a gregarious larval endoparasitoid, is one of three biocontrol agents from Asia currently being released in the United States to combat the invasive emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae). The current protocol for rearing T. planipennisi involves presenting the wasps with artificially infested ash sticks made by placing field-collected larvae into shallow grooves beneath flaps of bark. Although third and fourth instars are readily accepted by T. planipennisi in these exposures, the suitability of younger or older developmental stages, which are often more readily available in the field, has not been tested. In this study, we used both artificially infested ash sticks and naturally infested ash logs to test which emerald ash borer developmental stages (second to fourth instars, J larvae [preprepupae], prepupae, and pupae) are most suitable for rearing T. planipennisi. T. planipennisi parasitized all stages except for pupae, but parasitized fewer J larvae and prepupae in naturally infested logs than in artificially infested ash sticks. This is probably because, in naturally infested ash logs, these stages were confined to pupal chambers excavated in the sapwood and may have been largely beyond the reach of ovipositing T. planipennisi. The number of T. planipennisi progeny produced was positively correlated (logarithmic) with host weight, but this relationship was stronger when J larvae and prepupae were excluded from the data set. Fourth instars yielded the most parasitoid progeny, followed by, in approximately equal numbers, J larvae, prepupae, and third instars. Second instars yielded too few parasitoid progeny to benefit rearing efforts.

Effect of Trap Type, Trap Position, Time of Year, and Beetle Density on Captures of the Redbay Ambrosia Beetle (Coleoptera: Curculionidae: Scolytinae)

ABSTRACT The exotic redbay ambrosia beetle, Xyleborus glabratus Eichhoff (Coleoptera: Curculionidae: Scolytinae), and its fungal symbiont Raffaellea lauricola Harrington, Fraedrich, and Aghayeva are responsible for widespread redbay, Persea borbonia (L.) Spreng., mortality in the southern United States. Effective traps and lures are needed to monitor spread of the beetle and for early detection at ports-of-entry, so we conducted a series of experiments to find the best trap design, color, lure, and trap position for detection of X. glabratus. The best trap and lure combination was then tested at seven sites varying in beetle abundance and at one site throughout the year to see how season and beetle population affected performance. Manuka oil proved to be the most effective lure tested, particularly when considering cost and availability. Traps baited with manuka oil lures releasing 5 mg/d caught as many beetles as those baited with lures releasing 200 mg/d. Distributing manuka oil lures from the top to the bottom of eight-unit funnel traps resulted in similar numbers of X. glabratus as a single lure in the middle. Trap color had little effect on captures in sticky traps or cross-vane traps. Funnel traps caught twice as many beetles as cross-vane traps and three times as many as sticky traps but mean catch per trap was not significantly different. When comparing height, traps 1.5 m above the ground captured 85% of the beetles collected but a few were caught at each height up to 15 m. Funnel trap captures exhibited a strong linear relationship (r2 = 0.79) with X. glabratus attack density and they performed well throughout the year. Catching beetles at low densities is important to port of entry monitoring programs where early detection of infestations is essential. Our trials show that multiple funnel traps baited with a single manuka oil lure were effective for capturing X. glabratus even when no infested trees were visible in the area.

Evidence of cue synergism in termite corpse response behavior

Subterranean termites of the genus Reticulitermes are known to build walls and tubes and move considerable amounts of soil into wood but the causes of this behavior remain largely unexplored. In laboratory assays, we tested the hypothesis that Reticulitermes virginicus (Banks) would carry more sand into wooden blocks containing corpses compared to corpse-free controls. We further predicted that the corpses of predatory ants would elicit a stronger response than those of a benign beetle species or nestmates. As hypothesized, significantly more sand was carried into blocks containing corpses and this material was typically used to build partitions separating the dead from the rest of the colony. Contrary to expectations, however, this behavior did not vary among corpse types. We then tested the hypothesis that oleic acid, an unsaturated fatty acid released during arthropod decay and used by ants and other arthropod taxa in corpse recognition, would induce a similar building response in R. virginicus. To additionally determine the role of foreign objects in giving rise to this behavior, the experiment was carried out with and without imitation corpses (i.e., small glass beads). As predicted, oleic acid induced building (a tenfold increase) but only when applied to beads, suggesting strong synergism between tactile and chemical cues. Oleic acid also significantly reduced the amount of wood consumed by R. virginicus and may possess useful repellent properties.

Biology and Life History of Tetrastichus planipennisi (Hymenoptera: Eulophidae), a Larval Endoparasitoid of the Emerald Ash Borer (Coleoptera: Buprestidae)

ABSTRACT Tetrastichus planipennisi (Hymenoptera: Eulophidae) is a gregarious larval endoparasitoid from China that is being released in the United States as a biocontrol agent of the emerald ash borer (Agrilus planipennis Fairmaire), an exotic beetle responsible for widespread ash mortality. The developmental time of immature stages, adult longevity, reproductive age, oviposition rate, and realized fecundity were determined in the laboratory under normal rearing conditions (25 ± 2 °C, 65 ± 10% RH, and L:D 16:8 hr photoperiod). It takes approximately four weeks to complete a single generation (from egg to adult) under normal rearing conditions. While male wasps lived for a median of 5 weeks, females lived significantly longer with a median survival time of 6 weeks and a maximum survival time of 9 weeks. Newly emerged females appeared to be as capable of producing progeny as older females, indicating that T. planipennisi may be synovigenic. The average number of progeny per reproductive female remained relatively constant through the first 6 weeks of the trial with each reproductively active female producing 23–26 progeny each week. Lifetime realized fecundity of the adults averaged 57 progeny per reproductively active female with a female-tomale sex ratio of approximately 3:1; the maximum number of progeny produced by a single female was 108. These results suggest that T. planipennisi may have several generations in the mid-Atlantic and Midwestern regions of United States, where normal growing seasons (with average temperature >25 °C) are normally four to five months (May–Sep). Because of the gregarious nature, long life span and oviposition period of adults, T. planipennisi is likely to have multiple overlapping generations and has the potential to be an effective biocontrol agent against EAB.