X‐ray computed tomography reveals that intraspecific competition promotes soldier differentiation in a one‐piece nesting termite

Investment in soldier production in eusocial lineages involves a trade‐off between maintenance costs and defense benefits. Termites are eusocial insects that live in colonies organized into three castes: primary reproductives, soldiers, and workers or pseudergates. Neotermes chilensis (Blanchard) (Isoptera: Kalotermitidae) is a one‐piece nesting termite that nests and forages in a single piece of wood. Two scenarios may be of importance in a defense context of one‐piece nesting termites: during swarms, when colonies may be invaded by winged termites (alates) in search of a place to found a new colony, and when colonies of conspecifics are present within the same substrate. It was hypothesized that the ratio of soldiers to non‐soldiers would be higher at the onset of the swarming period and in substrates bearing more than one termite colony. A method based on X‐ray computed tomography (CT) was developed to study gallery connectivity in colonies of N. chilensis and caste composition within colonies. Computed tomography allowed the digital reconstruction of the galleries within the substrate, even when they belonged to different colonies, and was effective in distinguishing termites from substrate, and soldiers from reproductives and pseudergates. Using CT, the ratio of soldiers to non‐soldiers was shown to be highest in colonies within multicolonial scapes (i.e., neighboring colonies were present in the same substrate) during the swarming season, thus supporting our initial hypotheses. These results constitute a unique example of induced defenses arising from intraspecific interactions in termites.     

Asexual queen succession in the higher termite Embiratermes neotenicus

Asexual queen succession (AQS), in which workers, soldiers and dispersing reproductives are produced sexually while numerous non-dispersing queens arise through thelytokous parthenogenesis, has recently been described in three species of lower termites of the genus Reticulitermes. Here, we show that AQS is not an oddity restricted to a single genus of lower termites, but a more widespread strategy occurring also in the most advanced termite group, the higher termites (Termitidae). We analysed the genetic structure in 10 colonies of the Neotropical higher termite Embiratermes neotenicus (Syntermitinae) using five newly developed polymorphic microsatellite loci. The colonies contained one primary king accompanied either by a single primary queen or by up to almost 200 neotenic queens. While the workers, the soldiers and most future dispersing reproductives were produced sexually, the non-dispersing neotenic queens originated through thelytokous parthenogenesis of the founding primary queen. Surprisingly, the mode of thelytoky observed in E. neotenicus is most probably automixis with central fusion, contrasting with the automixis with terminal fusion documented in Reticulitermes. The occurrence of AQS based on different mechanisms of ploidy restoration raises the hypothesis of an independent evolutionary origin of this unique reproductive strategy in individual lineages of lower and higher termites.     

Estimation of foraging territories of Reticulitermes grassei through mark–release–recapture

Subterranean termites have highly cryptic life habits and their foraging activities are as a rule confined to below-ground level gallery systems. Mark–release–recapture (MRR) using fat-soluble histological dyes is a candidate method for the study of foraging dynamics and territories, but has not hitherto been standardized experimentally. A wide range of dye types and concentrations is reported in the literature. In this study, six potential dyes were evaluated at different concentrations for marking workers of Reticulitermes grassei (Clément) (Isoptera: Rhinotermitidae), under laboratory and simulated field conditions. Neutral red (1% wt/wt) was considered the most effective while showing acceptably low toxicity. In a subsequent field trial using dye impregnated into a wooden bait, the MMR procedure was used to map the foraging territory perimeter of a single colony. Assumptions inherent in the interpretation of MMR data are reviewed. To map the foraging territory perimeter by this method, two theoretical approaches are defined (a conservative and a non-conservative hypothesis). We show that the approach adopted may affect the estimate obtained by as much as 100%. Results are discussed in the context of the ecology and behaviour of subterranean termite colonies.     

Disturbance‐induced changes in a high‐alpine cushionfield community,south‐central New Zealand

Long‐term effects of two nearby disturbances, mechanical blading 34 years ago and an approximately century‐old sheep graveyard, in high‐alpine cushionfield share certain vegetation features despite their contrasting soil nutrient status. Secondary succession following blading has continued beyond that recorded over the first 24 years with continued trends towards the adjacent undisturbed cushionfield: increased cover of several cushion species and the lichen Cetraria islandica ssp. antarctica. Several mid‐seral species remain prominent while bare soil and stone pavement continue to decrease. Additional shelter, facilitated by tall snow tussocks planted locally at near‐natural density on part of the bladed site, allowed establishment of some larger, thin‐leaved species. Soil factors remain little affected by the mechanical disturbance compared with the general enrichment persisting at the graveyard. Here three small indigenous grasses, which dominated 50 years earlier (Trisetum spicatum, Agrostis muelleriana, Poa colensoi), remain prominent among the relatively diverse community (50 species, including several calciphilic lichens on the degrading bones). By contrast, frequency of the cushionfield dominant, Dracophyllum muscoides and the lichen Alectoria nigricans, are negatively correlated with the presence of bones. The post‐disturbance temporal community changes are assessed in the context of the initial productivity of the habitat, competitive exclusion and the intermediate disturbance hypothesis.     

Chemical warfare in termites

The rapid development of analytical methods in the last four decades has led to the discovery of a fascinating diversity of defensive chemicals used by termites. The last exhaustive review on termite defensive chemicals was published by G.D. Prestwich in 1984. In this text, we aim to fill the gap of the past 25 years and overview all of the relevant primary sources about the chemistry of termite defense (126 original papers, see Fig. 1 and online supplementary material) along with related biological aspects, such as the anatomy of defensive glands and their functional mechanisms, alarm communication, and the evolutionary significance of these defensive elements.     

Elevational environmental stress modulating species cohabitation in nests of a social insect

1. Termite nests may offer shelter to a number of species, alleviating the effects of environmental harshness. Certain elevational gradients provide variation on edaphoclimatic features, possibly generating harsh environmental conditions and boosting the number of immigrants seeking shelter within termitaria. Therefore, it is expected that metrics describing the community of termitaria cohabitants would correlate with elevation.
2. To test this hypothesis, we surveyed the termitophiles inhabiting 20 nests of Nasutitermes coxipoensis along an elevational gradient in a tropical mountain in Brazil. We assessed the richness, abundance, and composition (β‐diversity) of termite nests' cohabitants, testing nest volume and elevational position as explanatory covariates.
3. We found a positive correlation between the elevation at which termitaria were located and the richness and abundance of cohabiting termitophiles. Additionally, no correlation was found between elevational distance and dissimilarity of cohabitant communities between termitaria. Hence, the understanding that termitaria work as an ‘oasis’ of favorable microclimate is reinforced by our findings that the composition changed but was not correlated to elevation.
4. In short, environmental harshness boosts the establishment of distinct species of termitophiles in termitaria and it does so regardless of the invading species identity.

     

Microsatellite markers in soil‐feeding termites (Cubitermes subarquatus,Isoptera, Termitidae,Termitinae)

Seven microsatellite markers were isolated from Cubitermes subarquatus belonging to the soil‐feeding termite trophic group that plays a key role in tropical rain forests. A variability study performed by using a population of C. subarquatus (n = 73) sampled in the La Lopé forest reserve (Gabon) from 42 nests revealed from 4 to 12 alleles per locus and an heterozygosity from 0.28 to 0.84. Tests for cross‐species amplification realized in the sympatric C. intercalatus and in eight other sympatric termite genera indicated a wider application of the primers isolated from Cubitermes in soil‐feeding termites.     

Disturbance‐driven variation in abundance–occupancy relationships over time in tropical forest fragments

A central problem in ecology is to understand spatial–temporal variation in abundance–occupancy relationship (AOR) and to identify the biological and anthropogenic drivers behind this variation. How AOR is influenced by ecological traits and anthropogenic disturbances is poorly understood. A data set of woody plants from eleven forest fragments around Kampala area, Uganda, recorded in 1990 and resampled in 2010 was used to analyse spatial variation in AORs at regional scales, variation between tree size classes and temporal change in these patterns. Slopes of the AORs for each forest were related to data on forest disturbances and effects of species traits on AOR at regional scales assessed. There were statistically significant positive interspecific AORs at regional scales. Over two decades, the strength of AOR and slopes increased at the regional scale. At local scale, slopes decreased, in correspondence with increasing disturbance. Species traits interacted with abundance to explain statistically significant variation in occupancy. Species successional status best explained occupancy variability. Incorporating species' traits and anthropogenic disturbance over time may lead to better understanding of the variation in interspecific AOR, and these results suggest that anthropogenic exploitation could be responsible for the changes in slope over time.     

Repeated copulation and testes enlargement in Macrotermes michaelseni

ABSTRACT. The minimal copulation frequency of the reproductives of Macrotermes michaelseni (Sjöstedt) was investigated by removing the king from incipient laboratory colonies and from mature field colonies. Laboratory females stopped producing fertile eggs 56–154 days, and physogastric queens from field colonies less than 13 days after the removal of the king. Compared with young laboratory male reproductives, kings from mature field colonies are reported to have enormously enlarged testes, larger corpora allata (CA) and a higher juvenile hormone (JH) titre. However, JH application to young males did not increase the volume of the testes. The testes volumes only increased if the males were kept with additional workers or if their incipient colonies were allowed to grow continuously under unrestricted space conditions. These males also had bigger CA, and the soil in their midgut was replaced by food from the workers. It is concluded that in a growing colony the king is differently fed by the increasing number of workers and that trophic change in combination with higher CA activity induce enlargement of the testes. These enlarged testes are able to produce the sperm to fertilize the large number of eggs laid by a physogastric queen.     

Short‐term changes in the structure of termite assemblages associated with slash‐and‐burn agriculture in Côte d'Ivoire

Termites are major decomposers in tropical regions and play critical roles in many soil‐related processes. Studies conducted in Asia and the Neotropics suggest that habitat modification can strongly affect termite assemblages, but data on termite communities from forests in Africa, especially West Africa, are scarce. Here, we measured the short‐term impact of slash‐and‐burn agriculture on termite assemblages in an agricultural region of central Côte d'Ivoire. We assessed termite diversity and relative abundance in four habitat types: secondary forest, cleared forest, burned forest, and crop fields. The secondary forest had higher species richness compared with the other habitats, but all habitat types had similar assemblage structures. Fungus‐growing termites were the most abundant feeding group in all habitats. Soil feeders were most abundant in secondary forest, intermediately abundant in cleared and burned forests, and almost entirely absent in crop fields. Wood‐feeding species showed clear responses to burning; their abundances decreased after fire. We conclude that slash‐and‐burn agriculture does not appear to severely erode the diversity of termite assemblages. This could be due to the dominance of ecologically versatile fungus growers or to the relatively long time between clearing and burning. However, forest clearing negatively affects soil feeders, with the Apicotermitinae most affected by canopy loss.     

Male‐specific alleles in the Ryukyu drywood termite Neotermes sugioi

Sex‐determination systems often show remarkable diversity in upstream signals, although downstream genes are broadly conserved. Therefore, the downstream genes have been investigated in various taxa, but the most upstream signals determining sex in insects have been well‐described mainly in model organisms, including fruit flies and honey bees, and not in hemimetabolous insects such as termites. Identification of sex‐linked genetic markers in termites is important to the survey of primary sex‐determination signals. Here, we report male‐specific alleles at the microsatellite locus NK12‐1 in the Ryukyu drywood termite Neotermes sugioi (Kalotermitidae). This study provides the third example of a genetic marker linked with sexual phenotype in termites, which is a small but important step to elucidate the evolutionary process of the sex‐determination system in termites.     

Interspecific variation in the defence secretions of Nasutitermes soldiers

Termite soldiers of the genus Nasutitermes (Isoptera, Termitidae, Nasutitermitinae) eject a viscous, sticky defense secretion composed of non-polar monoterpene hydrocarbons and polyoxygenated diterpenes from the frontal gland. Chemical compositions are described in detail for the East African species, N. kempae and N. infuscatus, and for the Neotropical species N. ephratae. Additional comparisons with the new world species N. costalis, N. rippertii and N. octopilis are presented. The structure of a new monohydroxykempene from N. ephratae is described, and a physicochemical model for the stickiness of the glue is presented. Although the structures of the individual secretion components vary within the genus, the glue-like nature of the secretion remains essentially unchanged. Chemical analysis of soldier defense secretions may be useful in studying the systematic biology of termites.     

Soil‐dwelling insect pests of tree crops in Sub‐Saharan Africa,problems and management strategies—A review

This review aims to draw the attention of researchers, ecologists and farmers to the threats of soil‐dwelling insect pests on important tree crops in sub‐Saharan Africa, with a special focus on termites. It synthesizes the information on the effects of various factors affecting soil pest occurrence and damage, suggesting that the resultant undesirable effects of soil pests in this region are largely as a result of indiscriminate tree cutting, slash‐and‐burn agriculture and indiscriminate use of pesticides. Major insect orders, their host ranges and the nature of damage on selected tree crops are described. This study further critiques existing soil pest management practices, showing that majority of soil pest management practices are ineffective. Thus, management strategies like “attract and kill” approach based on entomopathogenic fungi need to be studied, developed and emphasized for the management of soil insect pests in sub‐Saharan Africa. A conclusion section attempts to offer suggestions for ways in which future work on soil pests in sub‐Saharan Africa could proceed.     

Expanding the population genetic perspective of cnidarian‐Symbiodinium symbioses

The modern synthesis was a seminal period in the biological sciences, establishing many of the core principles of evolutionary biology that we know today. Significant catalysts were the contributions of R.A. Fisher, J.B.S. Haldane and Sewall Wright (and others) developing the theoretical underpinning of population genetics, thus demonstrating adaptive evolution resulted from the interplay of forces such as natural selection and mutation within groups of individuals occupying the same space and time (i.e. a population). Given its importance, it is surprising that detailed population genetic data remain lacking for numerous organisms vital to many ecosystems. For example, the coral reef ecosystem is well recognized for its high biodiversity and productivity, numerous ecological services and significant economic and societal values (Moberg & Folke 1999; Cinner 2014). Many coral reef invertebrates form symbiotic relationships with single‐celled dinoflagellates within the genus Symbiodinium Freudenthal (Taylor 1974), with hosts providing these (typically) intracellular symbionts with by‐products of metabolism and in turn receiving photosynthetically fixed carbon capable of meeting hosts’ respiratory demands (Falkowski et al. 1984; Muscatine et al. 1984). Unfortunately, the health and integrity of the coral reef ecosystem has been significantly and negatively impacted by onslaughts like anthropogenic eutrophication and disease in addition to global climate change, with increased incidences of ‘bleaching’ events (characterized as the loss of photosynthetic pigments from the algal cell or massive reduction of Symbiodinium density from hosts’ tissue) and host mortality leading to staggering declines in geographic coverage (Bruno & Selig 2007) that have raised questions on the viability of this ecosystem as we know it (Bellwood et al. 2004; Parmesan 2006). One avenue towards anticipating the future of the coral reef ecosystem is by developing a broader and deeper understanding of the current genotypic diversity encompassed within and between populations of their keystone species, the scleractinian corals and dinoflagellate symbionts, as they potentially possess functional variation (either singularly or in combination) that may come under selection due to the ongoing and rapid environmental changes they are experiencing. However, such studies, especially for members of the genus Symbiodinium, are sparse. In this issue, Baums et al. (2014) provide a significant contribution by documenting the range‐wide population genetics of Symbiodinium ‘fitti’ (Fig. 1 ) in the context of complementary data from its host, the endangered Caribbean elkhorn coral Acropora palmata (Fig. 1 ). Notable results of this study include a single S. ‘fitti’ genotype typically dominates an individual A. palmata colony both spatially and temporally, gene flow among coral host populations is a magnitude higher to that of its symbiont populations, and the partners possess disparate patterns of genetic differentiation across the Greater Caribbean. The implications of such findings are discussed herein.     

Beta‐Diversity of Termite Assemblages Among Primary French Guiana Rain Forests

Termites are an important component of tropical rain forests, and have been included in many studies focusing on the influence of human disturbance. Their distribution among primary rain forest has, however, rarely been investigated. Here we studied the termite fauna in seven mostly undisturbed forest sites, representing several rain forest types. Overall, approximately 70 percent of species were soil‐feeders and 25 percent were wood‐feeders, the remaining 5 percent being classified here as litter‐feeders. Termite species richness did not differ significantly among sites, but sites differed in termite abundance. The palm swamp and the low forest situated on the foothills of an inselberg, hosted different termite communities to the other sites. These two sites presented a singular physiognomy suggesting that forest type is an important factor influencing species composition. We found no correlation between termite species composition and distance between sites, highlighting that at the scale of our study (about 100 km), forest sites share a similar species pool.     

Effects of large‐scale host plant addition and removal on parasitoid‐mediated associational resistance in the gall midge Asphondylia borrichiae

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Long‐term effects of repeated fuel‐reduction burning and logging on bats in south‐eastern Australia

Fire regimes have a major influence on biodiversity in many ecosystems around the globe, yet our understanding of the longer‐term response of fauna is typically poor. We sampled bats with ultrasonic detectors in three different years in dry sclerophyll forests of south‐eastern Australia in a long‐term, management‐scale experiment. Frequent low‐intensity burning (every 2 or 4 years plus unburnt) and logging (with 33% retention of the original unlogged tree basal area) were manipulated to assess their effects on bats. We found that both the fire regime and regrowth after logging influenced the local bat community. The routine burning treatment (burnt every 4 years) in unlogged forest was consistently related to higher total bat activity (2–3 times) and species richness when compared to unburnt controls and logging treatments. Foraging activity was more variable, but it was typically lowest in Unlogged Unburnt Controls. These patterns were evident at both the detector site scale and the block scale and were probably due to a reduction in understorey stem density with burning, especially in unlogged forest. Bat activity was significantly lower across the entire study area (including controls) in 1 year, when sampling occurred within 6 months of burning. When pooled across burning treatments, unlogged forest supported higher bat activity (1.5 times) and species richness than logged forest (12‐ to 17‐year‐old regrowth), again most likely because of a negative association with high stem density in regrowth after logging. We conclude that low‐intensity burning had positive benefits for echolocating bats, most notably in unlogged forest. However, careful planning is required to generate heterogeneous vegetation patterns that are likely to be most suitable for a range of taxa.     

Choice of hydrogen uptake (Hup) status in legume‐rhizobia symbioses

The H₂ is an obligate by-product of N-fixation. Recycling of H₂ through uptake hydrogenase (Hup) inside the root nodules of leguminous plants is often considered an advantage for plants. However, many of the rhizobium-legume symbioses found in nature, especially those used in agriculture are shown to be Hup⁻, with the plants releasing H₂ produced by nitrogenase activity from root nodules into the surrounding rhizosphere. Recent studies have suggested that, H₂ induces plant-growth-promoting rhizobacteria, which may explain the widespread of Hup⁻ symbioses in spite of the low energy efficiency of such associations. Wild legumes grown in Nova Scotia, Canada, were surveyed to determine if any plant-growth characteristics could give an indication of Hup choice in leguminous plants. Out of the plants sampled, two legumes, Securigera varia and Vicia cracca, showed Hup⁺ associations. Securigera varia exhibited robust root structure as compared with the other plants surveyed. Data from the literature and the results from this study suggested that plants with established root systems are more likely to form the energy-efficient Hup⁺ symbiotic relationships with rhizobia. Conversely, Hup⁻ associations could be beneficial to leguminous plants due to H₂-oxidizing plant-growth-promoting rhizobacteria that allow plants to compete successfully, early in the growing season. However, some nodules from V. cracca tested Hup⁺, while others were Hup⁻. This was similar to that observed in Glycine max and Pisum sativum, giving reason to believe that Hup choice might be affected by various internal and environmental factors.     

Abundance and diversity of termites (Isoptera) in imburnt versus burnt vegetation at the Barrens in Mediterranean Western Australia

Abstract Two years after an intense fire burnt large parts of the Barrens in Mediterranean Western Australia, its effects on wood-eating and litter-harvesting termites were investigated. Nine vegetation types varying in height, structure and floristics, were used for paired unburnt/burnt comparisons. Wood-eating termites were significantly less abundant and diverse in burnt stands, apparently by perishing in the fire rather than through food limitation. The harvester Tumulitermes westraliensis was not significantly affected by fire. Strategies enhancing persistence despite intense fire were construction of hard, protective clay mounds (Coptotermes frenchi, Amitermes obeuntis, T. westraliensis) and flexibility to site nests in diverse microhabitats, including the mounds of other termite species (Heterotermes).     

Ant‐termite interactions: an important but under‐explored ecological linkage

Animal interactions play an important role in understanding ecological processes. The nature and intensity of these interactions can shape the impacts of organisms on their environment. Because ants and termites, with their high biomass and range of ecological functions, have considerable effects on their environment, the interaction between them is important for ecosystem processes. Although the manner in which ants and termites interact is becoming increasingly well studied, there has been no synthesis to date of the available literature. Here we review and synthesise all existing literature on ant–termite interactions. We infer that ant predation on termites is the most important, most widespread, and most studied type of interaction. Predatory ant species can regulate termite populations and subsequently slow down the decomposition of wood, litter and soil organic matter. As a consequence they also affect plant growth and distribution, nutrient cycling and nutrient availability. Although some ant species are specialised termite predators, there is probably a high level of opportunistic predation by generalist ant species, and hence their impact on ecosystem processes that termites are known to provide varies at the species level. The most fruitful future research direction will be to evaluate the impact of ant–termite predation on broader ecosystem processes. To do this it will be necessary to quantify the efficacy both of particular ant species and of ant communities as a whole in regulating termite populations in different biomes. We envisage that this work will require a combination of methods, including DNA barcoding of ant gut contents along with field observations and exclusion experiments. Such a combined approach is necessary for assessing how this interaction influences entire ecosystems.