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The higher classification of termites requires substantial revision as the Neoisoptera, the most diverse termite lineage, comprise many paraphyletic and polyphyletic higher taxa. Here, we produce an updated termite classification using genomic-scale analyses. We reconstruct phylogenies under diverse substitution models with ultraconserved elements analyzed as concatenated matrices or within the multi-species coalescence framework. Our classification is further supported by analyses controlling for rogue loci and taxa, and topological tests. We show that the Neoisoptera are composed of seven family-level monophyletic lineages, including the Heterotermitidae Froggatt, Psammotermitidae Holmgren, and Termitogetonidae Holmgren, raised from subfamilial rank. The species-rich Termitidae are composed of 18 subfamily-level monophyletic lineages, including the new subfamilies Crepititermitinae, Cylindrotermitinae, Forficulitermitinae, Neocapritermitinae, Protohamitermitinae, and Promirotermitinae; and the revived Amitermitinae Kemner, Microcerotermitinae Holmgren, and Mirocapritermitinae Kemner. Building an updated taxonomic classification on the foundation of unambiguously supported monophyletic lineages makes it highly resilient to potential destabilization caused by the future availability of novel phylogenetic markers and methods. The taxonomic stability is further guaranteed by the modularity of the new termite classification, designed to accommodate as-yet undescribed species with uncertain affinities to the herein delimited monophyletic lineages in the form of new families or subfamilies., (© 2024. The Author(s).)

In the past, when scientists encountered and studied 'new' environmental phenomena, they rarely considered the existing knowledge of First Peoples (also known as Indigenous or Aboriginal people). The scientific debate over the regularly spaced bare patches (so-called fairy circles) in arid grasslands of Australian deserts is a case in point. Previous researchers used remote sensing, numerical modelling, aerial images and field observations to propose that fairy circles arise from plant self-organization. Here we present Australian Aboriginal art and narratives, and soil excavation data, that suggest these regularly spaced, bare and hard circles in grasslands are pavement nests occupied by Drepanotermes harvester termites. These circles, called linyji (Manyjilyjarra language) or mingkirri (Warlpiri language), have been used by Aboriginal people in their food economies and for other domestic and sacred purposes across generations. Knowledge of the linyji has been encoded in demonstration and oral transmission, ritual art and ceremony and other media. While the exact origins of the bare circles are unclear, being buried in deep time and Jukurrpa, termites need to be incorporated as key players in a larger system of interactions between soil, water and grass. Ecologically transformative feedbacks across millennia of land use and manipulation by Aboriginal people must be accounted for. We argue that the co-production of knowledge can both improve the care and management of those systems and support intergenerational learning within and across diverse cultures., (© 2023. The Author(s).)

Termites sense tiny substrate-borne vibrations through subgenual organs (SGOs) located within their legs' tibiae. Little is known about the SGOs' structure and physical properties. We applied high-resolution (voxel size 0.45 μm) micro-computed tomography (μCT) to Australian termites, Coptotermes lacteus and Nasutitermes exitiosus (Hill) to test two staining techniques. We compared the effectiveness of a single stain of Lugol's iodine solution (LS) to LS followed by Phosphotungstic acid (PTA) solutions (1% and 2%). We then present results of a soldier of Nasutitermes exitiosus combining μCT with LS + 2%PTS stains and scanning electron microscopy to exemplify the visualisation of their SGOs. The termite's SGO due to its approximately oval shape was shown to have a maximum diameter of 60 μm and a minimum of 48 μm, covering 60 ± 4% of the leg's cross-section and 90.4 ± 5% of the residual haemolymph channel. Additionally, the leg and residual haemolymph channel cross-sectional area decreased around the SGO by 33% and 73%, respectively. We hypothesise that this change in cross-sectional area amplifies the vibrations for the SGO. Since SGOs are directly connected to the cuticle, their mechanical properties and the geometric details identified here may enable new approaches to determine how termites sense micro-vibrations., Competing Interests: Declaration of competing interest The authors declare no conflict of interest., (Copyright © 2022 Elsevier Ltd. All rights reserved.)

Termites are major decomposers in terrestrial ecosystems and the second most diverse lineage of social insects. The Kalotermitidae form the second-largest termite family and are distributed across tropical and subtropical ecosystems, where they typically live in small colonies confined to single wood items inhabited by individuals with no foraging abilities. How the Kalotermitidae have acquired their global distribution patterns remains unresolved. Similarly, it is unclear whether foraging is ancestral to Kalotermitidae or was secondarily acquired in a few species. These questions can be addressed in a phylogenetic framework. We inferred time-calibrated phylogenetic trees of Kalotermitidae using mitochondrial genomes of ∼120 species, about 27% of kalotermitid diversity, including representatives of 21 of the 23 kalotermitid genera. Our mitochondrial genome phylogenetic trees were corroborated by phylogenies inferred from nuclear ultraconserved elements derived from a subset of 28 species. We found that extant kalotermitids shared a common ancestor 84 Ma (75-93 Ma 95% highest posterior density), indicating that a few disjunctions among early-diverging kalotermitid lineages may predate Gondwana breakup. However, most of the ∼40 disjunctions among biogeographic realms were dated at <50 Ma, indicating that transoceanic dispersals, and more recently human-mediated dispersals, have been the major drivers of the global distribution of Kalotermitidae. Our phylogeny also revealed that the capacity to forage is often found in early-diverging kalotermitid lineages, implying the ancestors of Kalotermitidae were able to forage among multiple wood pieces. Our phylogenetic estimates provide a platform for critical taxonomic revision and future comparative analyses of Kalotermitidae., (© The Author(s) 2022. Published by Oxford University Press on behalf of Society for Molecular Biology and Evolution.)

There are 28 invasive termite species, most belong to two families, the Kalotermitidae (esp. Cryptotermes spp.) and Rhinotermitidae (esp. Coptotermes spp.). Six invasive termite species are known to have spread into natural habitats, but little direct research has been conducted into their ecological impacts. Predictions based on indirect research (natural durability of commercial wood species) suggest fast-growing, pioneer tree species with low density wood, perhaps notably legumes, are most vulnerable to invasive termites, but even slow growing climax tree species may succumb. Cryptotermes will likely have less ecological impact, due to small colonies attacking dead branch stubs in the canopy. Coptotermes will likely have greater impact, due to large colony sizes and nesting in living trees, which they hollow out and can kill. There are no studies of invasive termites on native termites, other wood-eating insects, or predators, such as ants, showing considerable scope for future research., (Copyright © 2021 Elsevier Inc. All rights reserved.)

Termites inhabit complex underground mounds of intricate stigmergic labyrinthine designs with multiple functions as nursery, food storage and refuge, while maintaining a homeostatic microclimate. Past research studied termite building activities rather than the actual material structure. Yet, prior to understanding how multi-functionality shaped termite building, a thorough grasp of submillimetre mechanistic architecture of mounds is required. Here, we identify for Nasutitermes exitiosus via granulometry and Fourier transform infrared spectroscopy analysis, preferential particle sizes related to coarse silts and unknown mixtures of organic/inorganic components. High-resolution micro-computed X-ray tomography and microindentation tests reveal wall patterns of filigree laminated layers and sub-millimetre porosity wrapped around a coarse-grained inner scaffold. The scaffold geometry, which is designed of a lignin-based composite and densely biocementitious stercoral mortar, resembles that of trabecula cancellous bones. Fractal dimension estimates indicate multi-scaled porosity, important for enhanced evaporative cooling and structural stability. The indentation moduli increase from the outer to the inner wall parts to values higher than those found in loose clays and which exceed locally the properties of anthropogenic cementitious materials. Termites engineer intricately layered biocementitious composites of high elasticity. The multiple-scales and porosity of the structure indicate a potential to pioneer bio-architected lightweight and high-strength materials.

Animals use cues to find their food, in microhabitats within their physiological tolerances. Termites build and modify their microhabitat, to transform hostile environments into benign ones, which raises questions about the relative importance of cues. Termites are desiccation intolerant and foraging termites are attracted to water, so most research has considered moisture to be a cue. However, termites can also transport water to food, and so moisture may play other roles than previously considered. To examine the role of moisture, we compared Coptotermes acinaciformis termite foraging decisions in laboratory experiments when they were offered dry and moist wood, with and without load. Without load, termites preferred moist wood and ate it without any building, whereas they moistened dry wood after wrapping it in a layer of clay. For the 'With load' units, termites substituted some of the wood for load-bearing clay walls, and kept the wood drier than on the unloaded units. As drier wood has higher compressive strength and higher rigidity, it allows more of the wood to be consumed. These results suggest that moisture plays a more important role in termite ecology than previously thought. Termites manipulate the moisture content according to the situational context and use it for multiple purposes: increased moisture levels soften the fibre, which facilitates foraging, yet keeping the wood dry provides higher structural stability against buckling which is especially important when foraging on wood under load.

As eusocial and wood-dwelling insects, termites have been shown to use vibrations to assess their food, to eavesdrop on competitors and predators and to warn nest-mates. Bioassay choice experiments used to determine food preferences in animals often consider single factors only but foraging decisions can be influenced by multiple factors such as the quantity and quality of the food and the wood as a medium for communication. A statistical analysis framework is developed here to design a single bioassay experiment to study multifactorial foraging choice ( Pinus radiata) in the basal Australian termite species Coptotermes ( C.) acinaciformis (Isoptera: Rhinotermitidae). By employing a correlation analysis, 17 measured physical properties of 1417 Pinus radiata veneer discs were reduced to five key material properties: density, moisture absorption, early wood content, first resonance frequency and damping. By applying a fuzzy c-means clustering technique, these veneer discs were optimally paired for treatment and control trials to study food preference by termites based on these five key material properties. A multifactorial analysis of variance was compared to a permutation analysis of the results indicating for the first time that C. acinaciformis takes into account multiple factors when making foraging decisions. C. acinaciformis prefer denser wood with large early wood content, preferably humid and highly damped. Results presented here have practical implications for food choice experiments and for studies concerned with communication in termites as well as their ecology and coevolution with trees as their major food source.

The gut microbiota of animals exert major effects on host biology [1]. Although horizontal transfer is generally considered the prevalent route for the acquisition of gut bacteria in mammals [2], some bacterial lineages have co-speciated with their hosts on timescales of several million years [3]. Termites harbor a complex gut microbiota, and their advanced social behavior provides the potential for long-term vertical symbiont transmission, and co-evolution of gut symbionts and host [4-6]. Despite clear evolutionary patterns in the gut microbiota of termites [7], a consensus on how microbial communities were assembled during termite diversification has yet to be reached. Although some studies have concluded that vertical transmission has played a major role [8, 9], others indicate that diet and gut microenvironment have been the primary determinants shaping microbial communities in termite guts [7, 10]. To address this issue, we examined the gut microbiota of 94 termite species, through 16S rRNA metabarcoding. We analyzed the phylogeny of 211 bacterial lineages obtained from termite guts, including their closest relatives from other environments, which were identified using BLAST. The results provided strong evidence for rampant horizontal transfer of gut bacteria between termite host lineages. Although the majority of termite-derived phylotypes formed large monophyletic groups, indicating high levels of niche specialization, numerous other clades were interspersed with bacterial lineages from the guts of other animals. Our results indicate that "mixed-mode" transmission, which combines colony-to-offspring vertical transmission with horizontal colony-to-colony transfer, has been the primary driving force shaping the gut microbiota of termites., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

The higher termites (Termitidae) are keystone species and ecosystem engineers. They have exceptional biomass and play important roles in decomposition of dead plant matter, in soil manipulation, and as the primary food for many animals, especially in the tropics. Higher termites are most diverse in rainforests, with estimated origins in the late Eocene (∼54 Ma), postdating the breakup of Pangaea and Gondwana when most continents became separated. Since termites are poor fliers, their origin and spread across the globe requires alternative explanation. Here, we show that higher termites originated 42-54 Ma in Africa and subsequently underwent at least 24 dispersal events between the continents in two main periods. Using phylogenetic analyses of mitochondrial genomes from 415 species, including all higher termite taxonomic and feeding groups, we inferred 10 dispersal events to South America and Asia 35-23 Ma, coinciding with the sharp decrease in global temperature, sea level, and rainforest cover in the Oligocene. After global temperatures increased, 23-5 Ma, there was only one more dispersal to South America but 11 to Asia and Australia, and one dispersal back to Africa. Most of these dispersal events were transoceanic and might have occurred via floating logs. The spread of higher termites across oceans was helped by the novel ecological opportunities brought about by environmental and ecosystem change, and led termites to become one of the few insect groups with specialized mammal predators. This has parallels with modern invasive species that have been able to thrive in human-impacted ecosystems., (© The Author 2016. Published by Oxford University Press on behalf of the Society for Molecular Biology and Evolution. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Eavesdropping has evolved in many predator-prey relationships. Communication signals of social species may be particularly vulnerable to eavesdropping, such as pheromones produced by ants, which are predators of termites. Termites communicate mostly by way of substrate-borne vibrations, which suggest they may be able to eavesdrop, using two possible mechanisms: ant chemicals or ant vibrations. We observed termites foraging within millimetres of ants in the field, suggesting the evolution of specialised detection behaviours. We found the termite Coptotermes acinaciformis detected their major predator, the ant Iridomyrmex purpureus, through thin wood using only vibrational cues from walking, and not chemical signals. Comparison of 16 termite and ant species found the ants-walking signals were up to 100 times higher than those of termites. Eavesdropping on passive walking signals explains the predator detection and foraging behaviours in this ancient relationship, which may be applicable to many other predator-prey relationships., (© 2017 John Wiley & Sons Ltd/CNRS.)

Termite mounds built by representatives of the family Termitidae are among the most spectacular constructions in the animal kingdom, reaching 6-8 m in height and housing millions of individuals. Although functional aspects of these structures are well studied, their evolutionary origins remain poorly understood. Australian representatives of the termitid subfamily Nasutitermitinae display a wide variety of nesting habits, making them an ideal group for investigating the evolution of mound building. Because they feed on a variety of substrates, they also provide an opportunity to illuminate the evolution of termite diets. Here, we investigate the evolution of termitid mound building and diet, through a comprehensive molecular phylogenetic analysis of Australian Nasutitermitinae. Molecular dating analysis indicates that the subfamily has colonized Australia on three occasions over the past approximately 20 Myr. Ancestral-state reconstruction showed that mound building arose on multiple occasions and from diverse ancestral nesting habits, including arboreal and wood or soil nesting. Grass feeding appears to have evolved from wood feeding via ancestors that fed on both wood and leaf litter. Our results underscore the adaptability of termites to ancient environmental change, and provide novel examples of parallel evolution of extended phenotypes., (© 2017 The Author(s).)

Reticulitermes, Heterotermes and Coptotermes form a small termite clade with partly overlapping distributions. Although native species occur across all continents, the factors influencing their distribution are poorly known. Here, we reconstructed the historical biogeography of these termites using mitochondrial genomes of species collected on six continents. Our analyses showed that Reticulitermes split from Heterotermes + Coptotermesat 59.5 Ma (49.9-69.5 Ma 95% CI), yet the oldest split within Reticulitermes(Eurasia and North America) is 16.1 Ma (13.4-19.5 Ma) and the oldest split within Heterotermes + Coptotermesis 36.0 Ma (33.9-40.5 Ma). We detected 14 disjunctions between biogeographical realms, all of which occurred within the last 34 Ma, not only after the break-up of Pangaea, but also with the continents in similar to current positions. Land dispersal over land bridges explained four disjunctions, oceanic dispersal by wood rafting explained eight disjunctions, and human introduction was the source of two recent disjunctions. These wood-eating termites, therefore, appear to have acquired their modern worldwide distribution through multiple dispersal processes, with oceanic dispersal and human introduction favoured by the ecological traits of nesting in wood and producing replacement reproductives., (© 2016 The Author(s).)

Termites are eusocial insects that evolved from solitary cockroaches. It is not known precisely what factors drove the evolution of termite eusociality, that is, skewed reproduction with distinct winged reproductive and wingless worker phenotypes. In other eusocial insects (bees and wasps), reproductive skew evolved first and phenotype differences evolved second. We propose that the reverse pattern occurred in termites, that is, that the winged-wingless diphenism evolved before eusociality. We discuss existing phylogenetic and pheromonal evidence supporting our hypothesis. We provide new experimental evidence from the most basal termite species (Mastotermes darwiniensis), suggesting that the ancestral state was indeed diphenic but presocial. We propose that the mechanism promoting a winged-wingless diphenism-in the absence of eusociality-was greater predation of aerial than terrestrial dispersers, and we support this with a game theoretic model. We augment our hypothesis with a novel explanation for the evolution of the developmental pathways leading to winged and wingless phenotypes in termites. An added benefit of our hypothesis is that it neatly explains the origin of termite eusociality itself: in the pre-eusocial ancestral species, the poor dispersal ability of the wingless phenotype would have led to clustering of relatives around shared resources-a prerequisite for nonparental care of close relatives.

Many termite species use clay to build foraging galleries and mound-nests. In some cases clay is placed within excavations of their wooden food, such as living trees or timber in buildings; however the purpose for this clay is unclear. We tested the hypotheses that termites can identify load bearing wood, and that they use clay to provide mechanical support of the load and thus allow them to eat the wood. In field and laboratory experiments, we show that the lower termite Coptotermes acinaciformis, the most basal species to build a mound-nest, can distinguish unloaded from loaded wood, and use clay differently when eating each type. The termites target unloaded wood preferentially, and use thin clay sheeting to camouflage themselves while eating the unloaded wood. The termites attack loaded wood secondarily, and build thick, load-bearing clay walls when they do. The termites add clay and build thicker walls as the load-bearing wood is consumed. The use of clay to support wood under load unlocks otherwise unavailable food resources. This behaviour may represent an evolutionary step from foraging behaviour to nest building in lower termites.

Termite baiting is now one of the two main management tools in developed countries after 20 years of commercial release. It has two main goals: to use small amounts of active ingredient and 'colony elimination', i.e. death of all individuals in the colony. We consider how well baiting has been evaluated from 100 studies in the scientific literature. Studies have included 15 active ingredients, 23 termite species and 16 countries, yet most studies have focused on the chitin synthesis inhibitor hexaflumuron, Reticulitermes and the United States. Baiting has mostly met its goals: typically about 0.5 g of active ingredient was used, and colony elimination achieved, albeit with rates varying from 0 to 100%, and sometimes supplemented with liquid insecticide. Baiting was most successful using chitin synthesis inhibitors against Reticulitermes and Coptotermes (Rhinotermitidae), in temperate locations, although colony elimination was usually inferred indirectly - mostly by termite absence from baits - and was often slow, from 25 to 450 days. Baiting has been less tested and less successful against higher termites in tropical locations, where they are most diverse and abundant. Future research may have to consider greater termite species diversity and other active ingredients to reduce control times in order to fulfil the potential of baiting., (© 2014 Society of Chemical Industry.)

Termites have colonized many habitats and are among the most abundant animals in tropical ecosystems, which they modify considerably through their actions. The timing of their rise in abundance and of the dispersal events that gave rise to modern termite lineages is not well understood. To shed light on termite origins and diversification, we sequenced the mitochondrial genome of 48 termite species and combined them with 18 previously sequenced termite mitochondrial genomes for phylogenetic and molecular clock analyses using multiple fossil calibrations. The 66 genomes represent most major clades of termites. Unlike previous phylogenetic studies based on fewer molecular data, our phylogenetic tree is fully resolved for the lower termites. The phylogenetic positions of Macrotermitinae and Apicotermitinae are also resolved as the basal groups in the higher termites, but in the crown termitid groups, including Termitinae + Syntermitinae + Nasutitermitinae + Cubitermitinae, the position of some nodes remains uncertain. Our molecular clock tree indicates that the lineages leading to termites and Cryptocercus roaches diverged 170 Ma (153-196 Ma 95% confidence interval [CI]), that modern Termitidae arose 54 Ma (46-66 Ma 95% CI), and that the crown termitid group arose 40 Ma (35-49 Ma 95% CI). This indicates that the distribution of basal termite clades was influenced by the final stages of the breakup of Pangaea. Our inference of ancestral geographic ranges shows that the Termitidae, which includes more than 75% of extant termite species, most likely originated in Africa or Asia, and acquired their pantropical distribution after a series of dispersal and subsequent diversification events., (© The Author 2014. Published by Oxford University Press on behalf of the Society for Molecular Biology and Evolution. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

The termite genus Coptotermes (Rhinotermitidae) is found in Asia, Africa, Central/South America and Australia, with greatest diversity in Asia. Some Coptotermes species are amongst the world's most damaging invasive termites, but the genus is also significant for containing the most sophisticated mound-building termites outside the family Termitidae. These mound-building Coptotermes occur only in Australia. Despite its economic and evolutionary significance, the biogeographic history of the genus has not been well investigated, nor has the evolution of the Australian mound-building species. We present here the first phylogeny of the Australian Coptotermes to include representatives from all described species. We combined our new data with previously generated data to estimate the first phylogeny to include representatives from all continents where the genus is found. We also present the first estimation of divergence dates during the evolution of the genus. We found the Australian Coptotermes to be monophyletic and most closely related to the Asian Coptotermes, with considerable genetic diversity in some Australian taxa possibly representing undescribed species. The Australian mound-building species did not form a monophyletic clade. Our ancestral state reconstruction analysis indicated that the ancestral Australian Coptotermes was likely to have been a tree nester, and that mound-building behaviour has arisen multiple times. The Australian Coptotermes were found to have diversified ∼13million years ago, which plausibly matches with the narrowing of the Arafura Sea allowing Asian taxa to cross into Australia. The first diverging Coptotermes group was found to be African, casting doubt on the previously raised hypothesis that the genus has an Asian origin., (Copyright © 2014 Elsevier Inc. All rights reserved.)

A comparative field study was conducted to evaluate the ability of subterranean termites to damage a set of four different plastic materials (cable sheathings) exposed below- and above-ground. Eight pest species from six countries were included, viz., Coptotermes formosanus (Shiraki) in China, Japan, and the United States; Coptotermes gestroi (Wasmann) in Thailand and Malaysia; Coptotermes curvignathus (Holmgren) and Coptotermes kalshoveni (Kemner) in Malaysia; Coptotermes acinaciformis (Froggatt) with two forms of the species complex and Mastotermes darwiniensis (Froggatt) in Australia; and Reticulitermes flavipes (Kollar) in the United States. Termite species were separated into four tiers relative to decreasing ability to damage plastics. The first tier, most damaging, included C. acinaciformis, mound-building form, and M. darwiniensis, both from tropical Australia. The second tier included C. acinaciformis, tree-nesting form, from temperate Australia and C. kalshoveni from Southeast Asia. The third tier included C. curcignathus and C. gestroi from Southeast Asia and C. formosanus from China, Japan, and the United States, whereas the fourth tier included only R. flavipes, which caused no damage. A consequence of these results is that plastics considered resistant to termite damage in some locations will not be so in others because of differences in the termite fauna, for example, resistant plastics from the United States and Japan will require further testing in Southeast Asia and Australia. However, plastics considered resistant in Australia will be resistant in all other locations.

The number of recognized invasive termite species has increased from 17 in 1969 to 28 today. Fourteen species have been added to the list in the past 44 years; 10 have larger distributions and 4 have no reported change in distribution, and 3 species are no longer considered invasive. Although most research has focused on invasive termites in urban areas, molecular identification methods have answered questions about certain species and found that at least six species have invaded natural forest habitats. All invasive species share three characteristics that together increase the probability of creating viable propagules: they eat wood, nest in food, and easily generate secondary reproductives. These characteristics are most common in two families, the Kalotermitidae and Rhinotermitidae (which make up 21 species on the invasive termite list), particularly in three genera, Cryptotermes, Heterotermes, and Coptotermes (which together make up 16 species). Although it is the largest termite family, the Termitidae (comprising 70% of all termite species) have only two invasive species, because relatively few species have these characteristics. Islands have double the number of invasive species that continents do, with islands in the South Pacific the most invaded geographical region. Most invasive species originate from Southeast Asia. The standard control methods normally used against native pest termites are also employed against invasive termites; only two eradication attempts, in South Africa and New Zealand, appear to have been successful, both against Coptotermes species.

Despite their ecological significance as decomposers and their evolutionary significance as the most speciose eusocial insect group outside the Hymenoptera, termite (Blattodea: Termitoidae or Isoptera) evolutionary relationships have yet to be well resolved. Previous morphological and molecular analyses strongly conflict at the family level and are marked by poor support for backbone nodes. A mitochondrial (mt) genome phylogeny of termites was produced to test relationships between the recognised termite families, improve nodal support and test the phylogenetic utility of rare genomic changes found in the termite mt genome. Complete mt genomes were sequenced for 7 of the 9 extant termite families with additional representatives of each of the two most speciose families Rhinotermitidae (3 of 7 subfamilies) and Termitidae (3 of 8 subfamilies). The mt genome of the well supported sister-group of termites, the subsocial cockroach Cryptocercus, was also sequenced. A highly supported tree of termite relationships was produced by all analytical methods and data treatment approaches, however the relationship of the termites+Cryptocercus clade to other cockroach lineages was highly affected by the strong nucleotide compositional bias found in termites relative to other dictyopterans. The phylogeny supports previously proposed suprafamilial termite lineages, the Euisoptera and Neoisoptera, a later derived Kalotermitidae as sister group of the Neoisoptera and a monophyletic clade of dampwood (Stolotermitidae, Archotermopsidae) and harvester termites (Hodotermitidae). In contrast to previous termite phylogenetic studies, nodal supports were very high for family-level relationships within termites. Two rare genomic changes in the mt genome control region were found to be molecular synapomorphies for major clades. An elongated stem-loop structure defined the clade Polyphagidae + (Cryptocercus+termites), and a further series of compensatory base changes in this stem-loop is synapomorphic for the Neoisoptera. The complicated repeat structures first identified in Reticulitermes, composed of short (A-type) and long (B-type repeats) defines the clade Heterotermitinae+Termitidae, while the secondary loss of A-type repeats is synapomorphic for the non-macrotermitine Termitidae., (Copyright © 2012 Elsevier Inc. All rights reserved.)

Agricultural intensification has increased crop yields, but at high economic and environmental cost. Harnessing ecosystem services of naturally occurring organisms is a cheaper but under-appreciated approach, because the functional roles of organisms are not linked to crop yields, especially outside the northern temperate zone. Ecosystem services in soil come from earthworms in these cooler and wetter latitudes; what may fulfill their functional role in agriculture in warmer and drier habitats, where they are absent, is unproven. Here we show in a field experiment that ants and termites increase wheat yield by 36% from increased soil water infiltration due to their tunnels and improved soil nitrogen. Our results suggest that ants and termites have similar functional roles to earthworms, and that they may provide valuable ecosystem services in dryland agriculture, which may become increasingly important for agricultural sustainability in arid climates.

The efficacy of bistrifluron, a chitin synthesis inhibitor, in cellulose bait pellets was evaluated on the mound-building subterranean termite, Coptotermes acinaciformis (Froggatt). Three concentrations of the bistrifluron were used: 0 (untreated control), 0.5, and 1.0% over an 8 wk period. Both doses of bistrifluron bait eliminated (viz. termites absent from nest or mound) termite colonies: 83% of colonies (10 of 12) were either eliminated or moribund (viz. colony had no reproductive capacity and decreased workforce) after 8 wk, compared with none of the control colonies. The remaining two treated colonies were deemed to be in decline. Early signs that bistrifluron was affecting the colonies included: 3 wk after baiting mound temperatures showed a loss of metabolic heat, 4 wk after baiting foraging activity in feeding stations was reduced or absent, and dissection of two mounds at 4 wk showed they were moribund. Colony elimination was achieved in around half or less the time, and with less bait toxicant, than other bait products tested under similar conditions in the field, because of either the active ingredient, the high surface area of the pellets, or a combination of both. This suggests the sometimes long times reported for control using baits may be reduced significantly. The use of a mound building species demonstrated clearly colony level effects before and after termites stopped foraging in bait stations.

In contrast to the majority of the Order, the dampwood termites of the family Termopsidae found in colder regions can experience frost and snow, either in cool temperate areas at high latitudes (45 degrees ), or alpine areas at high elevations (>1000m). This suggests that dampwood termites are adapted to cold climates. We investigated this hypothesis in two dampwood termites, Porotermes adamsoni Froggatt and Stolotermes victoriensis Hill. We measured nest temperatures and atmospheric temperatures of their alpine habitat during winter, and measured survival and recovery at subzero temperatures. We also determined the minimum temperature at which these species remain active and the LT50 values. We used a novel gas chromatographic strategy to examine eight metabolites from individuals of both species collected in winter and summer to identify possible cryoprotectants. Both P. adamsoni and S. victoriensis had significantly higher levels of trehalose, a known cryoprotectant, in winter than in summer; in addition S. victoriensis also had higher levels of unsaturated fatty acid ligands in winter than in summer, consistent with patterns observed for cold adaptation in other organisms. These results are the first to reveal that dampwood termites are adapted to cold climates and use trehalose and unsaturated lipids as cryoprotectants.

Competition exclusion, when a single species dominates resources due to superior competitiveness, is seldom observed in nature. Termites compete for resources with deadly consequences, yet more than one species can be found feeding in the same wooden resource. This is especially surprising when drywood species, with colonies of a few hundred, are found cohabiting with subterranean species, with colonies of millions. Termites communicate vibro-acoustically and, as these signals can travel over long distances, they are vulnerable to eavesdropping. We investigated whether drywood termites could eavesdrop on vibration cues from subterranean species. We show, using choice experiments and recordings, that the drywood termite Cryptotermes secundus can distinguish its own species from the dominant competitor in the environment, the subterranean termite Coptotermes acinaciformis. The drywood termite was attracted to its own vibration cues, but was repelled by those of the subterranean species. This response increased with decreasing wood size, corresponding with both increased risk and strength of the cue. The drywood termites appear to avoid confrontation by eavesdropping on the subterranean termites; these results provide further evidence that vibro-acoustic cues are important for termite sensory perception and communication.

A year-long field experiment showed that Reticulitermes flavipes attacked and used single stakes and wooden stake bundles differently in two habitats that varied in alternative forage. In both habitats, the number of termites present in single stakes and stake-bundles increased with stake resource size, and the number of larvae recorded from feeding sites was a close reflection of the number of workers attracted to a given food source. Over 12 mo, more single stakes and stake bundles were heavily attacked and abandoned in the pine woods compared with grassy clearings. Paradoxically, the number of termites present in single stakes and stake bundles was greater in the food-poor habitat (grassy clearings), yet the wood was less consumed compared with the food-rich habitat (pine woods). Use of stake bundles as nests and the slower rate of wood consumption in grassy clearings indicated that termites seemed to value food resources in relative terms, i.e., those termites with abundant alternative food resources consumed quickly and departed, whereas those lacking extra resources ate slowly and settled in. These field results confirm earlier laboratory results that showed that termites modified wood consumption rates dependent on the amount of food available to the colony as a whole. The appearance of mature (physogastric) reproductives (>> yr old) in stake bundles in both habitats further indicated that termites track resources and opportunistically shift colony activities and possibly territory boundaries depending on the nature of the resources and other environmental factors. Implications for managing termites with a baiting strategy are discussed.

A phylogenetic hypothesis of termite relationships was inferred from DNA sequence data. Seven gene fragments (12S rDNA, 16S rDNA, 18S rDNA, 28S rDNA, cytochrome oxidase I, cytochrome oxidase II and cytochrome b) were sequenced for 40 termite exemplars, representing all termite families and 14 outgroups. Termites were found to be monophyletic with Mastotermes darwiniensis (Mastotermitidae) as sister group to the remainder of the termites. In this remainder, the family Kalotermitidae was sister group to other families. The families Kalotermitidae, Hodotermitidae and Termitidae were retrieved as monophyletic whereas the Termopsidae and Rhinotermitidae appeared paraphyletic. All of these results were very stable and supported with high bootstrap and Bremer values. The evolution of worker caste and foraging behavior were discussed according to the phylogenetic hypothesis. Our analyses suggested that both true workers and pseudergates ("false workers") were the result of at least two different origins. Our data support a traditional hypothesis of foraging behavior, in which the evolutionary transition from a one-piece type to a separate life type occurred through an intermediate behavioral form.

We formulate a mathematical model for food collection and production of workers and nymphs in 2 species of mound building termites. We maximise the number of nymphs (reproductives) produced by each colony over its lifetime with respect to the proportion of eggs that hatch as nymphs as opposed to workers. The results predict that food storage has a very important influence on the pattern of nymph and worker production. Food storage affects the part of the year that nymph production dominates, whether nymphs and workers are produced at the same time or not, and the existence of a final phase in the colony's life when a very large number of nymphs but no workers are produced.

Drywood termites are able to assess wood size using vibratory signals, although the exact mechanism behind this assessment ability is not known. Important vibratory characteristics such as the modal frequencies of a wooden block depend on its geometry and boundary conditions; however, they are also dependent on the material characteristics of the block, such as mass, density and internal damping. We report here on choice experiments that tested the ability of the drywood termite Cryptotermes secundus to assess wooden block size using a solid wooden block paired with a composite block, the latter made of either wood and aluminium or wood and rubber. Each composite block was constructed to match mass or low-frequency vibratory modes (i.e. fundamental frequency) of the solid wooden block. The termites always chose the blocks with more wood; they moved to the solid wooden blocks usually within a day and then tunnelled further into the solid wooden block by the end of the experiment. Termites offered composite blocks of wood and rubber matched for mass were the slowest to show a preference for the solid wooden block and this preference was the least definitive of any treatment, which indicated that mass and/or damping may play a role in food assessment. This result clearly shows that the termites were not fooled by composite blocks matched for mass or frequency, which implies that they probably employ more than a single simple measure in their food assessment strategy. This implies a degree of sophistication in their ability to assess their environment hitherto unknown. The potential importance of alternative features in the vibrational signals is discussed.

Contrary to the common perception that termites are indiscriminant eaters, termites choose their food carefully; however, the methods by which they choose food are not well understood. Using choice experiments and recordings of termites feeding on wooden blocks of different sizes, we show that worker drywood termites (Cryptotermes domesticus) use the resonant frequency of a block of wood to assess its size. Drywood termites showed differences in their response to vibration recordings of termites compared with artificially generated signals, suggesting that they can discriminate the source of vibration. Furthermore, fewer workers matured into neotenic reproductives when recorded termite signals were played, suggesting that vibration signals play an important role in termite communication.

Aim The Australian Coptotermes (Family: Rhinotermitidae) are a small monophyletic group of termites, some of which build mounds. In this study, we construct predicted distributions based on environmental data (niche models) for all Australian species of Coptotermes to test whether specific environmental factors have contributed to the evolution of mound-building behaviour and whether the degree of niche similarity and degree of phylogenetic similarity are correlated. Location The Australian mainland, including the known native ranges of all Australian species of Coptotermes. Methods We estimated the phylogenetic relationships between the species of Australian Coptotermes. We then generated and compared environmental niche models in a phylogenetic framework for all study species to test niche conservation. Our analyses were based on location data from our own sampling and from the Atlas of Living Australia, genetic data from a previous study of Australian Coptotermes, and environmental data from WorldClim and ASRIS. Results We found that no environmental variable differed consistently between mound-building and non-mound-building taxa and that the differences in niches between pairs of Australian species of Coptotermes are uncorrelated with time since divergence. The environmental tolerances of the Australian Coptotermes termites are more restricted by rainfall than they are by soil or temperature. Main conclusions Our results show that mound-building behaviour has not necessarily evolved in response to similar abiotic conditions. Our results are consistent with ecological speciation leading to niche divergence since Coptotermes first arrived in Australia ~12.5 million years ago. [ABSTRACT FROM AUTHOR]

Reticulitermes, Heterotermes and Coptotermes form a small termite clade with partly overlapping distributions. Although native species occur across all continents, the factors influencing their distribution are poorly known. Here, we reconstructed the historical biogeography of these termites using mitochondrial genomes of species collected on six continents. Our analyses showed that Reticulitermes split from Heterotermes + Coptotermes at 59.5 Ma (49.9-69.5 Ma 95% CI), yet the oldest split within Reticulitermes (Eurasia and North America) is 16.1 Ma (13.4-19.5 Ma) and the oldest split within Heterotermes + Coptotermes is 36.0 Ma (33.9-40.5 Ma). We detected 14 disjunctions between biogeographical realms, all of which occurred within the last 34 Ma, not only after the break-up of Pangaea, but also with the continents in similar to current positions. Land dispersal over land bridges explained four disjunctions, oceanic dispersal by wood rafting explained eight disjunctions, and human introduction was the source of two recent disjunctions. These wood-eating termites, therefore, appear to have acquired their modern worldwide distribution through multiple dispersal processes, with oceanic dispersal and human introduction favoured by the ecological traits of nesting in wood and producing replacement reproductives. [ABSTRACT FROM AUTHOR]

We developed 10 microsatellite loci in the primitive termite Mastotermes darwiniensis. The number of alleles per locus ranged from four to 15, and the expected heterozygosites spanned from 0.21 to 0.90, in a sample of 40 workers collected from the Northern Territory, Australia. We also determined that only two loci amplified in five other termite species. The low frequency of cross-amplification probably resulted from the high level of phylogenetic divergence between M. darwiniensis and the other taxa. Thus, although the loci are not widely applicable, they should prove effective in elucidating the genetic structure of M. darwiniensis populations. [ABSTRACT FROM AUTHOR]

Graphical abstract Mitochondrial genome phylogenetic tree of the termite subfamily Rhinotermitinae showing divergence time among the main geographic clades. The map depicts the biogeographic realms in which Rhinotermitinae occur, and the arrows show the most likely scenario of dispersal. Highlights • Mitochondrial genome phylogenetic supports the monophyly of Rhinotermitinae and all genera of Rhinotermitinae. • Rhinotermitinae arose about 50 million years ago, after the final phase of the Pangaea breakup. • Rhinotermitinae dispersed worldwide through at least four dispersal events. • Dispersals were both transoceanic dispersals, and dispersals across land bridges. Abstract Termites are the principal decomposers in tropical and subtropical ecosystems around the world. Time-calibrated molecular phylogenies show that some lineages of Neoisoptera diversified during the Oligocene and Miocene, and acquired their pantropical distribution through transoceanic dispersal events, probably by rafting in wood. In this paper, we intend to resolve the historical biogeography of one of the earliest branching lineages of Neoisoptera, the Rhinotermitinae. We used the mitochondrial genomes of 27 species of Rhinotermitinae to build two robust time-calibrated phylogenetic trees that we used to reconstruct the ancestral distribution of the group. Our analyses support the monophyly of Rhinotermitinae and all genera of Rhinotermitinae. Our molecular clock trees provided time estimations that diverged by up to 15.6 million years depending on whether or not 3rd codon positions were included. Rhinotermitinae arose 50.4–64.6 Ma (41.7–74.5 Ma 95% HPD). We detected four disjunctions among biogeographic realms, the earliest of which occurred 41.0–56.6 Ma (33.0–65.8 Ma 95% HPD), and the latest of which occurred 20.3–34.2 Ma (15.9–40.4 Ma 95% HPD). These results show that the Rhinotermitinae acquired their distribution through a combination of transoceanic dispersals and dispersals across land bridges. [ABSTRACT FROM AUTHOR]