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.     

Scale-dependent effects of landscape context on urban bee diversity

Journal of Insect Conservation - As urbanization continues throughout much of the world, there is great interest in better understanding the value of urban and residential environments to...     

Effects of Rhododendron removal and prescribed fire on bees and plants in the southern Appalachians

Rhododendron maximum is an evergreen shrub native to the Appalachian Mountains of North America that has expanded in recent decades due to past disturbances and land management. The purpose of this study was to explore how bees and plants were affected by the experimental removal of R. maximum followed by a prescribed fire in one watershed compared to a neighboring reference watershed. Bees and plants were sampled for three years in both watersheds. Comparisons were based on the rarefaction and extrapolation sampling curves of Hill numbers as well as multivariate methods to assess effects on community composition. Bee richness, Shannon''s diversity, and Simpson''s diversity did not differ between watersheds in the year after removal but were all significantly higher in the removal watershed in year two, following the prescribed fire. Bee Shannon''s diversity and Simpson''s diversity, but not richness, remained significantly higher in the removal watershed in the third year. Similar but weaker patterns were observed for plants. Comparisons of community composition found significant differences for bees in the second and third year and significant differences for plants in all three years. For both groups, significant indicator taxa were mostly associated with the removal watershed. Because bees appeared to respond more strongly to the prescribed fire than to the removal of R. maximum and these benefits weakened considerably one year after the fire, clearing R. maximum does not appear to dramatically improve pollinator habitat in the southern Appalachians. This conclusion is underscored by the fact that about one quarter of the bee species in our study area were observed visiting R. maximum flowers. The creation of open areas with wildflowers may be a better way to benefit bees in this region judging from the high diversity of bees captured in the small roadside clearings in this study.     

Feeding Preference of Native Subterranean Termites (Isoptera: Rhinotermitidae: Reticulitermes) for Wood Containing Bark Beetle Pheromones and Blue-Stain Fungi

Surprisingly little research has been conducted to investigate interactions between subterranean termites and bark beetles. Facilitative interactions between these organisms could easily alter stand dynamics and impact wood utilization strategies. A series of American Wood Protection Association Standard E1-09 “choice tests” were carried out to determine the feeding preference of Reticulitermes flavipes Kollar (Isoptera: Rhinotermitidae) for blue-stained sapwood and sapwood impregnated with various bark beetle pheromones. Reticulitermes flavipes exhibited a feeding preference for both air-dried and kiln-dried blue-stained sapwood, unstained sapwood treated with frontalin, and air-dried blue-stained sapwood treated with a 0.02% solution of both frontalin and endo-brevicomin. The implications of these results are far reaching, with particular relevance to forest health, ecology, and utilization.     

The importance of streamside sandbars to ground beetle (Coleoptera, Carabidae) communities in a deciduous forest

We used pitfall traps to sample ground beetles on sandbars along a small woodland stream and in the adjacent floodplain forest (Oglethorpe Co., GA, USA). We captured a total of 1,477 ground beetles representing 41 species. Twenty-two species were exclusive to sandbars, while eight were found only in the forested habitat. Ground beetles were captured in significantly greater numbers from sandbars, especially Brachinus janthinipennis and Omophron americanum. The B. janthinipennis record represents a new state record for the species. This study demonstrates that many unique species can be found in specialized microhabitats and emphasizes the need for biodiversity assessment surveys to include a wide range of these microhabitats within a survey area. In addition, it appears that many generalist forest species might use sandbars seasonally to exploit available resources. The U.S. Government's right to retain a non-exclusive, royalty-free license in and to any copyright is acknowledged.     

Abundance of volatile organic compounds in white ash phloem and emerald ash borer larval frass does not attract Tetrastichus planipennisi in a Y‐tube olfactometer

Many natural enemies employ plant‐ and/or herbivore‐derived signals for host/prey location. The larval parasitoid Tetrastichus planipennisi Yang (Hymenoptera: Eulophidae) is 1 of 3 biocontrol agents currently being released in an effort to control the emerald ash borer (EAB), Agrilus planipennis Fairmaire (Coloeptera: Burprestidae) in North America. To enhance its efficiency, allelochemicals that attract it need to be assessed. In this study, ash phloem volatile organic compounds (VOCs) of black, green, and white ash, and EAB larval frass were compared. Foraging behavior of T. planipennisi females in response to VOCs of white ash or frass from EAB larvae feeding on white ash phloem was tested using a Y‐tube olfactometer. Results indicated that the 3 ash species had similar VOC profiles. EAB larval frass generally contained greater levels of VOCs than phloem. Factor analysis indicated that the 11 VOCs could be broadly divided into 2 groups, with α‐bisabolol, β‐caryophyllene, (E)‐2‐hexenal, (Z)‐3‐hexenal, limonene, methyl benzoate, methyl indole‐3‐acetic acid, methyl jasmonate, methyl salicylate as the first group and the rest (i.e., methyl linoleate and methyl linolenate) as a second. Abundance of VOCs in white ash phloem tissue and frass, nevertheless, did not attract T. planipennisi females. The concealed feeding of EAB larvae might explain the selection for detectable and reliable virbrational signals, instead of undetectable and relatively unreliable VOC cues from phloem and frass, in short‐range foraging by T. planipennisi. Alternatively, it is possible that T. planipennisi is not amenable to the Y‐tube olfactometer assay employed.     

Response of beetles (Coleoptera) at three heights to the experimental removal of an invasive shrub, Chinese privet (Ligustrum sinense), from floodplain forests

Chinese privet (Ligustrum sinense Lour.), an invasive shrub from Asia, is well established in the southeastern United States where it dominates many floodplain forests. We used flight intercept traps to sample beetles at three heights (0.5, 5 and 15 m) in ~2 ha plots in which L. sinense had (by chainsaws or mulching machine) or had not been removed from forests in northeastern Georgia. Removing L. sinense, particularly by machine, increased the richness and diversity of beetles and affected composition near the ground (0.5 m) but not in the forest canopy (15 m). There were no differences among treatments above the L. sinense canopy (5 m) aside from Xylosandrus crassiusculus, an exotic ambrosia beetle from Asia, dominating the beetle community at that height in control plots. Removing the L. sinense layer greatly increased beetle richness near the ground, resulting in vertical distribution patterns more similar to those observed in areas of forest devoid of L. sinense. We suspect that even organisms in the canopy will benefit from privet removal in the long term given that tree regeneration is nearly impossible in forests in which L. sinense has become well established.     

The Response of Ground Beetles (Coleoptera: Carabidae) to Selection Cutting in a South Carolina Bottomland Hardwood Forest

We compared the response of ground beetles (Coleoptera: Carabidae) to the creation of canopy gaps of different size (0.13, 0.26, and 0.50 ha) and age (1 and 7 years) in a bottomland hardwood forest (South Carolina, USA). Samples were collected four times in 2001 by malaise and pitfall traps placed at the center and edge of each gap, and 50 m into the surrounding forest. Species richness was higher at the center of young gaps than in old gaps or in the forest, but there was no statistical difference in species richness between old gaps and the forests surrounding them. Carabid abundance followed the same trend, but only with the exclusion of Semiardistomis viridis (Say), a very abundant species that differed in its response to gap age compared to most other species. The carabid assemblage at the gap edge was very similar to that of the forest, and there appeared to be no distinct edge community. Species known to occur in open or disturbed habitats were more abundant at the center of young gaps than at any other location. Generalist species were relatively unaffected by the disturbance, but one species (Dicaelus dilatatus Say) was significantly less abundant at the centers of young gaps. Forest inhabiting species were less abundant at the centers of old gaps than in the forest, but not in the centers of young gaps. Comparison of community similarity at various trapping locations showed that communities at the centers of old and young gaps had the lowest similarity (46.5%). The community similarity between young gap centers and nearby forest (49.1%) and old gap centers and nearby forest (50.0%) was similarly low. These results show that while the abundance and richness of carabids in old gaps was similar to that of the surrounding forest, the species composition between the two sites differed greatly.     

Role of emerald ash borer (Coleoptera: Buprestidae) larval vibrations in host-quality assessment by Tetrastichus planipennisi (Hymenoptera: Eulophidae)

The biological control agent Tetrastichus planipennisi Yang (Hymenoptera: Eulophidae) is a gregarious larval endoparasitoid of the emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), an invasive cambium-feeding species responsible for recent, widespread mortality of ash (Fraxinus spp.) in North America. T. planipennisi is known to prefer late-instar emerald ash borer, but the cues used to assess host size by this species and most other parasitoids of concealed hosts remain unknown. We sought to test whether vibrations produced by feeding emerald ash borer vary with larval size and whether there are any correlations between these cues and T. planipennisi progeny number (i.e., brood size) and sex ratio. The amplitudes and rates of 3-30-ms vibrational impulses produced by emerald ash borer larvae of various sizes were measured in the laboratory before presenting the larvae to T. planipennisi. Impulse-rate did not vary with emerald ash borer size, but vibration amplitude was significantly higher for large larvae than for small larvae. T. planipennisi produced a significantly higher proportion of female offspring from large hosts than small hosts and was shown in previous work to produce more offspring overall from large hosts. There were no significant correlations, however, between the T. planipennisi progeny data and the emerald ash borer sound data. Because vibration amplitude varied significantly with host size, however, we are unable to entirely reject the hypothesis that T. planipennisi and possibly other parasitoids of concealed hosts use vibrational cues to assess host quality, particularly given the low explanatory potential of other external cues. Internal chemical cues also may be important.     

Differential utilization of ash phloem by emerald ash borer larvae: ash species and larval stage effects

• 1 Two experiments were performed to determine the extent to which ash species (black, green and white) and larval developmental stage (second, third and fourth instar) affect the efficiency of phloem amino acid utilization by emerald ash borer (EAB) Agrilus planipennis Fairmaire (Coleoptera: Buprestidae) larvae.
• 2 EAB larvae generally utilized green ash amino acids more efficiently than those of the other two species. For example, the concentrations of only six (two essential) and seven (two essential) amino acids were lower in frass from EAB that fed upon black and white ash than in the corresponding phloem, respectively. By contrast, concentrations of 16 (eight essential) amino acids were lower in the frass from EAB that fed upon green ash than in the phloem. In addition, in green ash, the frass : phloem ratios of 13 amino acids were lower than their counterparts in black and white ash.
• 3 The concentrations of non‐essential amino acids glycine and hydroxylproline were greater in frass than in phloem when EAB fed on black ash, although not when EAB fed on green or white ash.
• 4 The concentration of total phenolics (a group of putative defensive compounds to EAB, expressed as antioxidant activity of acetone extraction) was high in EAB frass but even higher in the phloem samples when the data were pooled across ash species and EAB larval stages. This suggests EAB larvae may eliminate phenolics through a combination of direct excretion and enzymatic conversion of phenolics to nonphenolics before excretion. Because the ratio of frass total phenolics to phloem total phenolics in white ash was lower than the ratios in black and green ash, the ability to destroy phenolics or convert them to nonphenolics was greater when EAB larvae fed on white ash.
• 5 Fourth‐instar EAB extracted phloem amino acids, including threonine, more efficiently than third‐instar EAB. The different larval developmental stages of EAB did not differ in their apparent ability to destroy phenolics or convert them to nonphenolics.