Intra- and Interspecific Comparisons of Bacterial Diversity and Community Structure Support Coevolution of Gut Microbiota and Termite Host

We investigated the bacterial gut microbiota from 32 colonies of wood-feeding termites, comprising four Microcerotermes species (Termitidae) and four Reticulitermes species (Rhinotermitidae), using terminal restriction fragment length polymorphism analysis and clonal analysis of 16S rRNA. The obtained molecular community profiles were compared statistically between individuals, colonies, locations, and species of termites. Both analyses revealed that the bacterial community structure was remarkably similar within each termite genus, with small but significant differences between sampling sites and/or termite species. In contrast, considerable differences were found between the two termite genera. Only one bacterial phylotype (defined with 97% sequence identity) was shared between the two termite genera, while 18% and 50% of the phylotypes were shared between two congeneric species in the genera Microcerotermes and Reticulitermes, respectively. Nevertheless, a phylogenetic analysis of 228 phylotypes from Microcerotermes spp. and 367 phylotypes from Reticulitermes spp. with other termite gut clones available in public databases demonstrated the monophyly of many phylotypes from distantly related termites. The monophyletic “termite clusters” comprised of phylotypes from more than one termite species were distributed among 15 bacterial phyla, including the novel candidate phyla TG2 and TG3. These termite clusters accounted for 95% of the 960 clones analyzed in this study. Moreover, the clusters in 12 phyla comprised phylotypes from more than one termite (sub)family, accounting for 75% of the analyzed clones. Our results suggest that the majority of gut bacteria are not allochthonous but are specific symbionts that have coevolved with termites and that their community structure is basically consistent within a genus of termites.     

Chemical constituents of tropical woods and resistance to the invasive drywood termite Cryptotermes brevis

Wood-feeding, nesting and production of secondary reproductives are key determinant traits of invasive species of drywood termites, and the West Indian drywood termite Cryptotermes brevis (Walker) is one of their major examples of worldwide concern as pest species of structural lumber, furniture and other wood products. The problem and losses by this species are determined by the prevailing wood characteristics. However, despite the current widespread occurrence of this species in the tropics, except Asia, tropical wood resistance and underlying mechanisms of resistance against this termite are scarcely known. Nonetheless, wood hardness and particularly wood density were recently recognized as important underlying traits for C. brevis resistance in tropical woods, but the chemical wood constituents were not considered. Here, we assessed tropical wood resistance to the invasive termite species C. brevis and tested the relevance of their holocellulose, lignin and (total) extractive contents preventing termite infestation. Free-choice and no-choice tests were carried out in parallel with wood chemical characterization. Resistance to the West Indian termite varied with wood species in terms of both colonization and consumption, but only under free-choice testing because without choice, no significant difference was detected among wood species. Regardless, none of these traits were significantly correlated with wood resistance to C. brevis. Therefore, wood physical resistance, particularly wood density, seems the main recognized determinant of tropical wood resistance against the West Indian drywood termite. The pattern of termite movement on the surface of soft, mid and hard wood was also consistent with this conclusion.     

Field Evaluations of Subterranean Termite Preference for Sap-Stain Inoculated Wood

Few studies have focused on interactions between subterranean termites and the ophiostomatoid fungal associates of pine bark beetles or root feeding weevils. Field stake tests were employed at four locations throughout Mississippi to determine the feeding preference of subterranean termites for blue-stained, unstained, and partially decayed southern pine sapwood stakes. This study also utilized wood decayed by Gloeophyllum trabeum, a fungus previously shown to elicit a positive subterranean termite feeding response, as a positive control. Stakes inoculated with G. trabeum received significantly more attacks than all other treatments after 16 weeks. Of the stakes attacked by subterranean termites, stakes inoculated with Ophiostoma minus were degraded faster than any other treatment. Subterranean termite preference for stakes treated with either of two Leptographium spp. and the untreated negative controls did not differ; however, each was fed upon less than all other treatments. The feeding rate on stakes inoculated with O. ips and G. trabeum being fed upon by subterranean termites was not significantly different. These results represent the first evidence of wood containing non-structurally degrading fungi (O. ips and O. minus) eliciting a feeding preference from subterranean termites greater than that of decayed wood. The implications of these results are particularly relevant to pine forest ecology, nutrient cycling, subterranean termite control, and the utilization of blue-stained southern pine building products in the southeastern U.S.     

Decay and termite resistance of medium density fiberboard (MDF) made from different wood species

Medium density fiberboard (MDF) production worldwide is increasing due to the development of new manufacturing technologies. As a result, MDF products are increasingly utilized in traditional wood applications that require fungal and insect resistance. This study evaluated the ability of white and brown rot fungi and termites to decompose MDF consisting of different wood species by measuring weight loss. Furnish in the boards was prepared from heart and sapwood portions of pine (Pinus nigra Arnold var. pallasiana), beech (Fagus orientalis Lipsky), and European oak (Quercus robur L.) species. Fungal decay resistance tests were performed according to ASTM D 2017-81 standard method using two brown-rot fungi, Gloeophyllum trabeum (Pers. ex Fr.) Murr. (Mad 617), Postia placenta (Fries) M. Larsen et Lombard (Mad 698), and one white-rot fungus, Trametes versicolor (L. ex Ft.) Pilat (Mad 697). MDF and wood specimens were also bioassayed against the eastern subterranean termite, Reticulitermes flavipes (Kollar) in order to determine termite resistance of the specimens. MDF specimens containing oak and mixed furnish demonstrated increased durability against decay fungi. Only pine, oak, and mixed MDF specimens met the 25% or less weight loss limit to be classified resistant according to ASTM D 2017-81 standard method. Overall, MDF specimens made from oak showed better performance than oak solid wood specimens. Accelerated aging according to ASTM D 1037-96a standard method before fungal bioassay decreased fungal resistance of the specimens. In contrast to the fungal bioassay, MDF specimens made from beech and mixed furnish showed decreased weight losses from termite attack after 4 weeks. However, none of the MDF specimens were resistant to termite attack. In severe conditions, the MDFs may require the incorporation of chemical biocides prior to board production for increasing the resistance of MDF to termite attack.     

Nitrous Oxide (N2O) Emissions by Termites: Does the Feeding Guild Matter?

In the tropics, termites are major players in the mineralization of organic matter leading to the production of greenhouse gases including nitrous oxide (N₂O). Termites have a wide trophic diversity and their N-metabolism depends on the feeding guild. This study assessed the extent to which N₂O emission levels were determined by termite feeding guild and tested the hypothesis that termite species feeding on a diet rich in N emit higher levels of N₂O than those feeding on a diet low in N. An in-vitro incubation approach was used to determine the levels of N₂O production in 14 termite species belonging to different feeding guilds, collected from a wide range of biomes. Fungus-growing and soil-feeding termites emit N₂O. The N₂O production levels varied considerably, ranging from 13.14 to 117.62 ng N₂O-N d^-1 (g dry wt.)^-1 for soil-feeding species, with Cubitermes spp. having the highest production levels, and from 39.61 to 65.61 ng N₂O-N d^-1 (g dry wt.)^-1 for fungus-growing species. Wood-feeding termites were net N₂O consumers rather than N₂O producers with a consumption ranging from 16.09 to 45.22 ng N₂O-N d^-1 (g dry wt.)^-1. Incubating live termites together with their mound increased the levels of N₂O production by between 6 and 13 fold for soil-feeders, with the highest increase in Capritermes capricornis, and between 14 and 34 fold for fungus-growers, with the highest increase in Macrotermes muelleri. Ammonia-oxidizing (amoA-AOB and amoA-AOA) and denitrifying (nirK, nirS, nosZ) gene markers were detected in the guts of all termite species studied. No correlation was found between the abundance of these marker genes and the levels of N₂O production from different feeding guilds. Overall, these results support the hypothesis that N₂O production rates were higher in termites feeding on substrates with higher N content, such as soil and fungi, compared to those feeding on N-poor wood.     

Comparative natural durability of five wood species from Mozambique

The objective of this study was to evaluate the natural durability of five lesser-utilized wood species from Mozambique. Both laboratory methods and field tests were applied for assessing wood decay of muanga (Pericopsis angolensis), metil (Sterculia appendiculata), namuno (Acacia nigrescens), ncurri (Icuria dunensis), and ntholo (Pseudolachnostylis maprounaefolia). Laboratory tests involved soft-, brown-, and white-rot fungi and termites. Heart- and sapwood of ncurri and ntholo were exposed in above-ground field tests; additionally, all species were exposed to in-ground contact tests. The results indicated that namuno, muanga, ncurri, and ntholo are resistant to soft-, brown- and white-rot fungi and the termite species Reticulitermes grassea and Mastotermes darwiniensis. Comparatively, soft-rot caused more severe decay on the studied wood species than did basidiomycete fungi. The brown-rot fungi Coniophora puteana, Gloeophiyllum trabeum, and Postia placenta caused less decay on the tested species than did the white-rot Trametes versicolor. Metil was not resistant to any of the mentioned hazards. Therefore, this species is not recommendable for exterior use if untreated.     

Molecular Identification of Sibling Species of Sclerodermus (Hymenoptera: Bethylidae) That Parasitize Buprestid and Cerambycid Beetles by Using Partial Sequences of Mitochondrial DNA Cytochrome Oxidase Subunit 1 and 28S Ribosomal RNA Gene

The species belonging to Sclerodermus (Hymenoptera: Bethylidae) are currently the most important insect natural enemies of wood borer pests, mainly buprestid and cerambycid beetles, in China. However, some sibling species of this genus are very difficult to distinguish because of their similar morphological features. To address this issue, we conducted phylogenetic and genetic analyses of cytochrome oxidase subunit I (COI) and 28S RNA gene sequences from eight species of Sclerodermus reared from different wood borer pests. The eight sibling species were as follows: S. guani Xiao et Wu, S. sichuanensis Xiao, S. pupariae Yang et Yao, and Sclerodermus spp. (Nos. 1–5). A 594-bp fragment of COI and 750-bp fragment of 28S were subsequently sequenced. For COI, the G-C content was found to be low in all the species, averaging to about 30.0%. Sequence divergences (Kimura-2-parameter distances) between congeneric species averaged to 4.5%, and intraspecific divergences averaged to about 0.09%. Further, the maximum sequence divergences between congeneric species and Sclerodermus sp. (No. 5) averaged to about 16.5%. All 136 samples analyzed were included in six reciprocally monophyletic clades in the COI neighbor-joining (NJ) tree. The NJ tree inferred from the 28S rRNA sequence yielded almost identical results, but the samples from S. guani, S. sichuanensis, S. pupariae, and Sclerodermus spp. (Nos. 1–4) clustered together and only Sclerodermus sp. (No. 5) clustered separately. Our findings indicate that the standard barcode region of COI can be efficiently used to distinguish morphologically similar Sclerodermus species. Further, we speculate that Sclerodermus sp. (No. 5) might be a new species of Sclerodermus.     

Microbial Communities in Sunken Wood Are Structured by Wood-Boring Bivalves and Location in a Submarine Canyon

The cornerstones of sunken wood ecosystems are microorganisms involved in cellulose degradation. These can either be free-living microorganisms in the wood matrix or symbiotic bacteria associated with wood-boring bivalves such as emblematic species of Xylophaga, the most common deep-sea woodborer. Here we use experimentally submerged pine wood, placed in and outside the Mediterranean submarine Blanes Canyon, to compare the microbial communities on the wood, in fecal pellets of Xylophaga spp. and associated with the gills of these animals. Analyses based on tag pyrosequencing of the 16S rRNA bacterial gene showed that sunken wood contained three distinct microbial communities. Wood and pellet communities were different from each other suggesting that Xylophaga spp. create new microbial niches by excreting fecal pellets into their burrows. In turn, gills of Xylophaga spp. contain potential bacterial symbionts, as illustrated by the presence of sequences closely related to symbiotic bacteria found in other wood eating marine invertebrates. Finally, we found that sunken wood communities inside the canyon were different and more diverse than the ones outside the canyon. This finding extends to the microbial world the view that submarine canyons are sites of diverse marine life.     

Wood-Destroying Soft Rot Fungi in the Historic Expedition Huts of Antarctica

Three expedition huts in the Ross Sea region of Antarctica, built between 1901 and 1911 by Robert F. Scott and Ernest Shackleton, sheltered and stored the supplies for up to 48 men for 3 years during their explorations and scientific investigation in the South Pole region. The huts, built with wood taken to Antarctica by the early explorers, have deteriorated over the past decades. Although Antarctica has one of the coldest and driest environments on earth, microbes have colonized the wood and limited decay has occurred. Some wood in contact with the ground contained distinct microscopic cavities within secondary cell walls caused by soft rot fungi. Cadophora spp. could be cultured from decayed wood and other woods sampled from the huts and artifacts and were commonly associated with the soft rot attack. By using internal transcribed spacer sequences of ribosomal DNA and morphological characteristics, several species of Cadophora were identified, including C. malorum, C. luteo-olivacea, and C. fastigiata. Several previously undescribed Cadophora spp. also were found. At the Cape Evans and Cape Royds huts, Cadophora spp. commonly were isolated from wood in contact with the ground but were not always associated with soft rot decay. Pure cultures of Cadophora used in laboratory decay studies caused dark staining of all woods tested and extensive soft rot in Betula and Populus wood. The presence of Cadophora species, but only limited decay, suggests there is no immediate threat to the structural integrity of the huts. These fungi, however, are widely found in wood from the historic huts and have the capacity to cause extensive soft rot if conditions that are more conducive to decay become common.     

Do the unique properties of nanometals affect leachability or efficacy against fungi and termites?

Nanotechnology has the potential to affect the field of wood preservation through the creation of new and unique metal biocides with improved properties. This study evaluated leachability and efficacy of southern yellow pine wood treated with copper, zinc, or boron nanoparticles against mould fungi, decay fungi, and Eastern subterranean termites. Results showed that nanocopper with and without surfactant, nanozinc, and nanozinc plus silver with surfactant resisted leaching compared with metal oxide controls. Nearly all nanoboron and boric acid was released from the treated wood specimens during leaching. Mould fungi were moderately inhibited by nanozinc oxide with surfactant, but the other nanometal preparations did not significantly inhibit mould fungi. Mass loss from Gloeophyllum trabeum was significantly inhibited by all copper preparations, while Antrodia sp. was not inhibited by nanometal treatments. Nanometals imparted high resistance in southern yellow pine to the white-rot fungus, Trametes versicolor. Unleached specimens treated with nanoboron or nanozinc plus surfactant caused 100% and 31% mortality, respectively. All specimens treated with nanozinc or nanozinc plus silver inhibited termite feeding, but the copper treatments were less effective against termites. Nanozinc possessed the most favorable properties: leach resistance, termite mortality, and inhibition of termite feeding and decay by the white-rot fungus.     

Changes in the termite assemblage across a sequence of land-use systems in the rural area around Lamto Reserve in central Côte d’Ivoire

Termites are major decomposers in tropical regions and play an important role in soil processes. This study investigated the termite assemblage structure across a sequence of differing land-use systems. With a standardized method, data were collected on termites from the following habitats: semi-deciduous forest, teak plantation, cocoa plantation, Jatropha plantation, food crop field and 4-years old fallow. Termite species richness declined from the semi-deciduous forest to 4-years old fallow through teak plantation, food crop field, cocoa plantation and Jatropha plantation. The relative abundance of fungus-growers was the highest in all land-use types while that of soil-feeders steeply declined in all man-modified sites. The wood-feeding species showed clear responses to disturbance, with low abundances in monospecific- and modified sites without high trees. Comparisons with other studies suggest that changes in the termite assemblage structure result from forest conversion to agricultural systems. To help mitigate the loss of termites when forests are disturbed or cleared, we recommend to: (1) promote the association of cropping and silvicultural systems that reduces changes in microclimate and maintains the original termite assemblage with the associated ecosystem services; (2) leave dead wood on the ground after forest disturbance to accelerate the recovery of the termite assemblage; and (3) increase forest and silvicultural patch size and reduce length of forest edges to sustain the survival of forest-dependent species.     

Fuel Characteristics of Solid Biofuel Derived from Oil Palm Biomass and Fast Growing Timber Species in Malaysia

The fuel characteristics of solid biofuels derived from biomass that is abundantly available in Malaysia are presented. The objectives of the study were to characterize fuel properties of oil palm biomass (empty fruit bunch (EFB) and oil palm trunk (OPT)) and wood from a range of fast growing timber species (Albizia falcataria, Acacia spp., Endospermum spp. and Macaranga spp.), inclusive and exclusive of bark. Among the fast-growing timber species, the higher heating values ranged from 4288 cal g^-1 to 4383 cal g^-1 for wood inclusive of bark, and 4134 cal g^-1 to 4343 cal g^-1 for wood exclusive of bark. The inclusive of bark portion in the biomass sample generally increased the heating value except for Macaranga spp. Empty fruit bunch and oil palm trunk had heating values of 4315 cal g^-1 and 4104 cal g^-1, respectively. Ash-forming elements and trace elements were much higher in the timber species samples inclusive of bark than samples exclusive of bark. On the other hand, oil palm biomass contained higher ash-forming elements and trace elements than the wood from the fast growing timber species. The European energy crops show higher HHV, Cl and Si content but lower K, Mg, Na and P compared to the local biomass used in this study. The data obtained from this study can serve as a foundation for the selection of suitable biomass to be used as solid fuel, or as a reference on the fabrication of conversion systems for the selection of biomass solid fuel.     

Taxonomic Identification of Mediterranean Pines and Their Hybrids Based on the High Resolution Melting (HRM) and trnL Approaches: From Cytoplasmic Inheritance to Timber Tracing

Fast and accurate detection of plant species and their hybrids using molecular tools will facilitate the assessment and monitoring of local biodiversity in an era of climate and environmental change. Herein, we evaluate the utility of the plastid trnL marker for species identification applied to Mediterranean pines (Pinus spp.). Our results indicate that trnL is a very sensitive marker for delimiting species biodiversity. Furthermore, High Resolution Melting (HRM) analysis was exploited as a molecular fingerprint for fast and accurate discrimination of Pinus spp. DNA sequence variants. The trnL approach and the HRM analyses were extended to wood samples of two species (Pinus nigra and Pinus sylvestris) with excellent results, congruent to those obtained using leaf tissue. Both analyses demonstrate that hybrids from the P. brutia (maternal parent) × P. halepensis (paternal parent) cross, exhibit the P. halepensis profile, confirming paternal plastid inheritance in Group Halepensis pines. Our study indicates that a single one-step reaction method and DNA marker are sufficient for the identification of Mediterranean pines, their hybrids and the origin of pine wood. Furthermore, our results underline the potential for certain DNA regions to be used as novel biological information markers combined with existing morphological characters and suggest a relatively reliable and open taxonomic system that can link DNA variation to phenotype-based species or hybrid assignment status and direct taxa identification from recalcitrant tissues such as wood samples.     

Influence of Soil Properties on Soldierless Termite Distribution

In tropical rainforests, termites constitute an important part of the soil fauna biomass, and as for other soil arthropods, variations in soil composition create opportunities for niche partitioning. The aim of this study was twofold: first, we tested whether soil-feeding termite species differ in the foraging substrate; second, we investigated whether soil-feeding termites select their foraging sites to enhance nutrients intake. To do so, we collected termites and analysed the composition and structure of their feeding substrates. Although Anoplotermes-group members are all considered soil-feeders, our results show that some species specifically feed on abandoned termite nests and very rotten wood, and that this substrate selection is correlated with previous stable isotope analyses, suggesting that one component of niche differentiation among species is substrate selection. Our results show that the composition and structure of bare soils on which different termite species foraged do not differ, suggesting that there is no species specialization for a particular type of bare soil. Finally, the bare soil on which termites forage does not differ from random soil samples. Overall, our results suggest that few species of the Anoplotermes-group are specialized toward substrates rich in organic matter, but that the vast majority forage on soil independently of its structural and chemical composition, being ecologically equivalent for this factor.     

Distribution patterns of four Termitomyces species cultivated by a fungus-growing termite,Odontotermes formosanus,in Taiwan

Fungus-growing termites inoculate the obligate symbiotic fungus Termitomyces into the fungus comb in their colonies. In this study, Taiwanese Termitomyces species were determined by diagnostic PCR using the metagenome of the body of Odontotermes formosanus (the only host termite living in Taiwan) as a template. Phylogenetic analyses revealed that four different Termitomyces species are cultivated by O. formosanus in Taiwan. Three have previously been registered in the DNA database, but one was first recorded in Taiwan. Only Termitomyces sp. Type C was distributed in all areas investigated in Taiwan, whereas the other three species were distributed regionally. Field observations indicated that the flush period and the number of fruit bodies in each colony varied between species. The distribution patterns of Termitomyces spp. in Taiwan may be related to the Taiwanese climate and/or the fruiting pattern.     

Effects of two locust control methods on wood-eating termites in arid Australia

Termites are ubiquitous detritivores and are a key influence on soil function and nutrient cycles, particularly in arid and semi-arid ecosystems. Locust control presents a unique hazard to termites and the effective functioning of ecosystems as a consequence of the overlap between pesticide applications and termite populations in grassland and desert landscapes. We monitored the effects of locust control methods using ultra-low-volume (ULV) barrier application of a chemical pesticide, fipronil, and a blanket application of a fungal biopesticide, Metarhizium acridum, on wood-eating termites in arid western New South Wales, Australia. We tested the hypothesis that spray applications decrease termite activity at wood baits using a BACI designed field experiment over 2 years. Our replicated control and treatment sites represented the spatial scale of Australian locust control activities. There was no detectable impact of either locust control treatment on termite activity, bait mass loss or termite community composition measures. Non-significant differences in termite survey measures among sites suggested that climate and environmental conditions were stronger drivers of our termite measures than the single, localized and unreplicated application of pesticides more commonly used in locust control operations in arid Australia. A lack of evidence for an impact of our fipronil or Metarhizium application methods supports their use as low hazard locust control options with minimal large scale and longer-term effects on termites in Australian arid rangelands. Future research would be necessary to determine the probable short-term impacts of treatments on individual termite colonies and the possible impacts on non-wood eating termite species in the arid-zone.     

Colony structure and caste distribution in living trees of the Ryukyu drywood termite,Neotermes sugioi (Blattodea: Kalotermitidae) in Okinawa Island

Termites are major pests of houses and buildings, and also living plants such as agricultural crops, trees in forests, urban areas and gardens. However, in Japan, the basic ecology of termites nesting in living trees is not fully understood. In this study, we observed 255 colonies (nests) of the drywood termite Neotermes sugioi, collected in the field on Okinawa Island, and reported the frequency composition of the reproductive castes, the size of wood with termite gallery, the population size of colonies, and the relative position of the reproductive and non-reproductive castes within nests. Most colonies were headed by a primary queen and a primary king. However, colonies headed only by primary queens, primary kings, or neotenic kings, each accounted for approximately 5% of the colonies. A colony size of 1,000– 4,000 individuals (2058.2 ± 1695.0 [mean ± SD]) was common and the average length of colony branches was less than 100 cm. Queens and kings were found in the same or nearby nest areas, and more predominantly in the central or root side of nest wood areas. The termites may experience colony fragmentation and reproductive loss as a consequence of typhoons. Incipient colonies (i.e., colonies at an early stage of development) were found on 11.3% of branches of Leucaena leucocephala that did not show any obvious signs of infestation. In future research, it will be necessary to update the list of trees damaged by this termite species, compare the damage by tree species, and evaluate the economic impact.     

Analysis of Extensive [FeFe] Hydrogenase Gene Diversity Within the Gut Microbiota of Insects Representing Five Families of Dictyoptera

We have designed and utilized degenerate primers in the phylogenetic analysis of [FeFe] hydrogenase gene diversity in the gut ecosystems of roaches and lower termites. H₂ is an important free intermediate in the breakdown of wood by termite gut microbial communities, reaching concentrations in some species exceeding those measured for any other biological system. The primers designed target with specificity the largest group of enzymatic H domain proteins previously identified in a termite gut metagenome. “Family 3” hydrogenase sequences were amplified from the guts of lower termites, Incisitermes minor, Zootermopsis nevadensis, and Reticulitermes hesperus, and two roaches, Cryptocercus punctulatus and Periplaneta americana. Subsequent analyses revealed that all termite and Cryptocercus sequences were phylogenetically distinct from non-termite-associated hydrogenases available from public databases. The abundance of unique sequence operational taxonomic units (as many as 21 from each species) underscores the previously demonstrated physiological importance of H₂ to the gut ecosystems of these wood-feeding insects. The diversity of sequences observed might be reflective of multiple niches that the enzymes have been evolved to accommodate. Sequences cloned from Cryptocercus and the lower termite samples, all of which are wood feeding insects, clustered closely with one another in phylogenetic analyses to the exclusion of alleles from P. americana, an omnivorous cockroach, also cloned during this study. We present primers targeting a family of termite gut [FeFe] hydrogenases and provide results that are consistent with a pivotal role for hydrogen in the termite gut ecosystem and point toward unique evolutionary adaptations to the gut ecosystem.