Defense of termitaria by termitophilous ants

Summary Mounds of Amitermes laurensis are frequently faided by meat ants Iridomyrmex sanguineus. Of eight ant species which often cohabit with the termites, Camponotus sp. B and C were considerably dependent on the termintaria for their nest sites and effectively protected it from the attacks by meat ants. Many termite colonies cohabiting with those two ant species were vigorous, suggesting that this ant-termite relationship is mutualistic; thus, the ants were provided nest sites and probably even food and the termites were protected from destructive natural enemies.     

Functional structure of ant and termite assemblages in old growth forest,logged forest and oil palm plantation in Malaysian Borneo

Forested tropical landscapes around the world are being extensively logged and converted to agriculture, with serious consequences for biodiversity and potentially ecosystem functioning. Here we investigate associations between habitat disturbance and functional diversity of ants and termites—two numerically dominant and functionally important taxa in tropical rain forests that perform key roles in predation, decomposition, nutrient cycling and seed dispersal. We compared ant and termite occurrence and composition within standardised volumes of soil and dead wood in old growth forest, logged forest and oil palm plantation in Sabah, Malaysian Borneo. Termites occurred substantially less frequently in converted habitats than in old growth forest, whereas ant occurrences were highest in logged forest and lowest in old growth forest. All termite feeding groups had low occurrence in disturbed habitats, with soil feeders occurring even less frequently than wood feeders. Ant functional groups showed more variable associations, with some opportunist and behaviourally dominant groups being more abundant in degraded habitats. The importance of ants and termites in tropical ecosystems and such differing patterns of assemblage variation suggest that ecosystem functioning may be significantly altered in converted habitats.     

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.     

Protein marking reveals predation on termites by the woodland ant, <Emphasis Type="Italic">Aphaenogaster rudis</Emphasis>

Subterranean termites provide a major potential food source for forest-dwelling ants, yet the interactions between ants and termites are seldom investigated largely due to the cryptic nature of both the predator and the prey. We used protein marking (rabbit immunoglobin protein, IgG) and double antibody sandwich enzyme-linked immunosorbent assay (DAS-ELISA) to examine the trophic interactions between the woodland ant, Aphaenogaster rudis (Emery) and the eastern subterranean termite, Reticulitermes flavipes (Kollar). We marked the prey by feeding the termites paper treated with a solution of rabbit immunoglobin protein (IgG). Subsequently, we offered live, IgG-fed termites to ant colonies and monitored the intracolony distribution of IgG-marked prey. Laboratory experiments on the distribution of protein-marked termite prey in colonies of A. rudis revealed that all castes and developmental stages receive termite prey within 24 h. In field experiments, live, protein-marked termites were offered to foraging ants. Following predation, the marker was recovered from the ants, demonstrating that A. rudis preys on R. flavipes under field conditions. Our results provide a unique picture of the trophic-level interactions between predatory ants and subterranean termites. Furthermore, we show that protein markers are highly suitable to track trophic interactions between predators and prey, especially when observing elusive animals with cryptic food-web ecology. Received 19 January 2007; revised 23 March 2007; accepted 26 March 2007.     

Feeding habits of Hymenoptera and Isoptera in a tropical rain forest as revealed by nitrogen and carbon isotope ratios

Despite the recognition of the functional role of Hymenoptera (ants, bees and wasps) and Isoptera (termites) in tropical ecosystems, their detailed feeding habits are not well known. To examine the feeding habits of these groups, we measured nitrogen (N) and carbon (C) stable isotope ratios (δ¹⁵N and δ¹³C) of hymenopterans (12 families, ≥16 genera and ≥32 species) and isopterans (one family and 10 species) collected in a tropical rain forest, Sarawak, Malaysia. We compared the isotopic signatures of these insects to those previously reported for other consumers collected in the same forest. The δ¹⁵N and δ¹³C values of these insects overlapped with those of the other consumers, indicating that they have access to diverse C and N sources in the forest. The δ¹⁵N values of ants and termites indicated that their feeding habits range along a continuum from herbivory (i.e. dependent on honeydew and nectar) to predation and from wood-feeders to soil-feeders, respectively. In addition, the δ¹⁵N values of wasps varied greatly from −0.1‰ (Braconidae sp.) to 8.6‰ (Bembix sp.), suggesting that their feeding habits also range from omnivory to predation. The ant species Camponotus gigas had δ¹³C values similar to those of invertebrate detritivores and omnivores rather than to those of invertebrate herbivores, although the diet of this species consists mostly of honeydew. This discrepancy suggests that the ant uses carbohydrates as an energy source, the isotopic signatures of which are not well retained in the body tissues. Values of both δ¹⁵N and δ¹³C of the predatory army ant Leptogenys diminuta and the soil-feeding termite Dicuspiditermes nemorosus did not differ significantly, indicating that both trophic level and the humification of feeding substrates can increase the isotopic signatures of terrestrial consumers.     

Behavioral Interactions Between <Emphasis Type="Italic">Aphaenogaster rudis</Emphasis> (Hymenoptera: Formicidae) and <Emphasis Type="Italic">Reticulitermes flavipes</Emphasis> (Isoptera: Rhinotermitidae): The Importance of Physical Barriers

Predation pressure from ants is a major driving force in the adaptive evolution of termite defense strategies and termites have evolved elaborate chemical and physical defenses to protect themselves against ants. We examined predator–prey interactions between the woodland ant, Aphaenogaster rudis (Emery) and the eastern subterranean termite, Reticulitermes flavipes (Kollar), two sympatric species widely distributed throughout deciduous forests in eastern North America. To examine the behavioral interactions between A. rudis and R. flavipes we used a series of laboratory behavioral assays and predation experiments where A. rudis and R. flavipes could interact individually or in groups. One-on-one aggression tests revealed that R. flavipes are vulnerable to predation by A. rudis when individual termite workers or soldiers are exposed to ant attacks in open dishes and 100% of termite workers and soldiers died, even though the soldiers were significantly more aggressive towards the ants. The results of predation experiments where larger ant and termite colony fragments interacted provide experimental evidence for the importance of physical barriers for termite colony defense. In experiments where the termites nested within artificial nests (sand-filled containers), A. rudis was aggressive at invading termite nests and inflicted 100% mortality on the termites. In contrast, termite mortality was comparable to controls when termite colonies nested in natural nests comprised of wood blocks. Our results highlight the importance of physical barriers in termite colony defense and suggest that under natural field conditions termites may be less susceptible to attacks by ants when they nest in solid wood, which may offer more structural protection than sand alone.     

Driver Ants Invading a Termite Nest: Why Do the Most Catholic Predators of All Seldom Take This Abundant Prey?

Driver ants ( i.e. , epigaeic species in the army ant genus Dorylus , subgenus Anomma ) are among the most extreme polyphagous predators, but termites appear to be conspicuously absent from their prey spectrum and attacks by driver ants on termite nests have not yet been described. Here, we report a Dorylus ( Anomma ) rubellus attack on a colony of the fungus-growing termite Macrotermes subhyalinus that was observed during the dry season in a savannah habitat in Nigeria's Gashaka National Park. It was estimated that several hundred thousand termites (probably more than 2.4 kg dry mass) were retrieved. The apparent rarity of driver ant predation on Macrotermes nests may be explained by different habitat requirements, by the fact that these ants mostly forage aboveground, by efficient termite defense behavior and nest architecture that make entry into the nest difficult, and finally by driver ant worker morphology, which differs remarkably from that of subterranean Dorylus species that regularly invade and destroy termite colonies.     

Ant colonization and coarse woody debris decomposition in temperate forests

Ants are ubiquitous, abundant and have widespread impacts on ecological communities and ecosystem processes. However, ant effects on coarse woody debris decomposition are unexplored. Several ant species colonize coarse woody debris for nesting, and this puts them in contact with fauna and microbes that utilize coarse woody debris as habitat and food, potentially influencing nutrient cycling and, ultimately, forest productivity. We report results from a field experiment employing 138 artificial ant nests (routed pine blocks) across five locations in southeastern US deciduous forests. We examine the correspondence between ant, termite and wood-eating fungi colonization and variation in coarse woody debris decomposition. After 1 year, nests colonized by ants had 5% more mass than those not colonized. Ant colonization corresponded with significantly less termite- and fungal-mediated decomposition of the nests. Without ants, termites removed 11.5% and fungi removed 4% more wood biomass. Ants, termites and wood-eating fungi all colonized pine nests where temperatures were highest, and ants also preferred higher soil moisture whereas termites and fungi responded negatively to high soil moisture when temperatures were higher. Ants reduce termite colonies through predation, and may inhibit fungi through the secretion of antimicrobial compounds. Our results indicate that interactions between forest understory ants, termites and fungi may influence the rate of coarse woody debris decomposition—biotic interactions that potentially influence forest structure and function.     

Formyltetrahydrofolate synthetase gene diversity in the guts of higher termites with different diets and lifestyles

In this study, we examine gene diversity for formyl-tetrahydrofolate synthetase (FTHFS), a key enzyme in homoacetogenesis, recovered from the gut microbiota of six species of higher termites. The "higher" termites (family Termitidae), which represent the majority of extant termite species and genera, engage in a broader diversity of feeding and nesting styles than the "lower" termites. Previous studies of termite gut homoacetogenesis have focused on wood-feeding lower termites, from which the preponderance of FTHFS sequences recovered were related to those from acetogenic treponemes. While sequences belonging to this group were present in the guts of all six higher termites examined, treponeme-like FTHFS sequences represented the majority of recovered sequences in only two species (a wood-feeding Nasutitermes sp. and a palm-feeding Microcerotermes sp.). The remaining four termite species analyzed (a Gnathamitermes sp. and two Amitermes spp. that were recovered from subterranean nests with indeterminate feeding strategies and a litter-feeding Rhynchotermes sp.) yielded novel FTHFS clades not observed in lower termites. These termites yielded two distinct clusters of probable purinolytic Firmicutes and a large group of potential homoacetogens related to sequences previously recovered from the guts of omnivorous cockroaches. These findings suggest that the gut environments of different higher termite species may select for different groups of homoacetogens, with some species hosting treponeme-dominated homoacetogen populations similar to those of wood-feeding, lower termites while others host Firmicutes-dominated communities more similar to those of omnivorous cockroaches.     

Strong but taxon‐specific responses of termites and wood‐nesting ants to forest regeneration in Borneo

Land use change is accelerating globally at the expense of biodiversity and ecosystem functioning. Invertebrates are numerically dominant and functionally important in old growth tropical rain forests but highly susceptible to the adverse effects of forest degradation and fragmentation. Ants (Formicidae) and termites (Blattodea: Termitoidae) perform crucial ecosystem services. Here, the potential effects of anthropogenic disturbance on ant and termite communities in dead wood are investigated. Community composition, generic richness, and occupancy rates of ants and termites were compared among two old growth sites (Danum Valley and Maliau Basin) and one twice‐logged site (the Stability of Altered Forest Ecosystems’ (SAFE) Project), in Sabah, Malaysian Borneo. Occupancy was measured as the number of ant or termite encounters (1) per deadwood items, and (2) per deadwood volume, and acts as surrogates for relative abundance (or generic richness). Termites had a lower wood‐occupancy per volume in logged forest. In contrast, there were more ant encounters, and more ant genera, in logged sites and there was a community shift (especially, there were more Crematogaster encounters). The disruption of soil and canopy structure in logged forest may reduce both termite and fungal decay rates, inducing increased deadwood residence times and therefore favoring ants that nest in dead wood. There is an anthropogenic‐induced shift of dead wood in ants and termites in response to disturbance in tropical rain forests and the nature of that shift is taxon‐specific.     

Ant and termite mound coinhabitants in the wetlands of Santo Antonio da Patrulha, Rio Grande do Sul, Brazil.

This paper reports on ant and termite species inhabiting the mounds (murundus) found in three wetland sites in Santo Antonio da Patrulha. Ants and termites were found in 100% of the mounds of two sites and in 20% of those in the third site. Colonies of Camponotus fastigatus were found inhabiting all the mounds, while colonies of Brachymyrmex sp., Linepithema sp., Pheidole sp., and/or Solenopsis sp. were collected in less than 30% of the mounds. In the mounds of the three sites, colonies of Anoplotermes sp. and/or Aparatermes sp. termites were found together with the ant colonies. Another cohabiting termite species, Cortaritermes sp., was found only in the mounds of one site. The results suggest that C. fastigatus is the species building the mounds, with the other species, whether ants or termites, being the inquilines.     

成都市白蚁天敌资源——蚂蚁种类调查研究

部分蚂蚁种类是白蚁的重要天敌。为了探明成都市白蚁天敌——蚂蚁的常见种类及其生境,2011—2015年通过对成都市房屋建筑、园林树木、古建筑以及安装在住宅小区绿地的地下型白蚁监测装置等生境中调查白蚁与蚂蚁种类及数量,共鉴定白蚁9种,隶属于3科4属,黑胸散白蚁Reticulitermes chinensis占比最高(83.9%);蚂蚁29种,隶属于4亚科15属,中国短猛蚁Brachyponera chinensis占比最高(38.2%)。相关性分析表明,黑毛蚁Lasius niger与尖唇散白蚁R.aculabialis、新中华散白蚁R.neochinensis之间,莱曼氏蚁Formica lemani与黑胸散白蚁、尖唇散白蚁之间,敏捷扁头猛蚁Pachycondyla astuta、铺道蚁Tetramorium caesputum与尖唇散白蚁之间均呈显著负相关。黑毛蚁、莱曼氏蚁、敏捷扁头猛蚁、铺道蚁的种群数量大,在白蚁生物防治中具有一定的应用潜力。鉴于蚂蚁对白蚁的控制作用受环境、人为等多种因素的影响,建议进一步加强蚂蚁对白蚁自然控制作用的系统研究。     

Community interactions between ants and arboreal-nesting termites in New Guinea coconut plantations

Summary: We investigated the incidence of inquiline ants and of arboreal-nesting ants on a community of three arboreal-nesting termites living in New Guinea coconut plantations. Inquiline ants were present in 10 % of Microcerotermes biroi nests and in 4 % of Nasutitermes princeps nests. Live termite nests inhabited by the most common inquiline ant, Camponotus sp. A, were generally left by the ant after several months. In some nests, Camponotus sp. A was observed coexisting with its host during the whole observation period (3 years). Therefore, Camponotus sp. A was apparently an opportunistic inquiline which did not affect significantly the mortality of termite colonies. The arboreal-nesting ant, Crematogaster irritabilis, was locally found occupying up to 99 % of the trees present in 1 ha plots. In such hot spots, the overall abundance of termites was approximately half that of plots devoid of Crematogaster irritabilis. The high density of Crematogaster irritabilis may be an important limiting factor for the termite assemblage, by hastening the death or hindering the establishment of arboreal termite colonies.     

Nectar quality affects ant aggressiveness and biotic defense provided to plants

Ant–plant mutualisms are useful models for investigating how plant traits mediate interspecific interactions. As plant‐derived resources are essential components of ant diets, plants that offer more nutritious food to ants should be better defended in return, as a result of more aggressive behavior toward natural enemies. We tested this hypothesis in a field experiment by adding artificial nectaries to individuals of the species Vochysia elliptica (Vochysiaceae). Ants were offered one of four liquid foods of different nutritional quality: amino acids, sugar, sugar + amino acids, and water (control). We used live termites (Nasutitermes coxipoensis) as herbivore competitors and observed ant behavior toward them. In 88 hr of observations, we recorded 1,009 interactions with artificial nectaries involving 1,923 individual ants of 26 species. We recorded 381 encounters between ants and termites, of which 38% led to attack. Sixty‐one percent of these attacks led to termite exclusion from the plants. Recruitment and patrolling were highest when ants fed upon nectaries providing sugar + amino acids, the most nutritious food. This increase in recruitment and patrolling led to higher encounter rates between ants and termites, more frequent attacks, and faster and more complete termite removal. Our results are consistent with the hypothesis that plant biotic defense is mediated by resource quality. We highlight the importance of qualitative differences in nectar composition for the outcome of ant–plant interactions. Abstract in Portuguese is available with online material.     

Iron reduction in the metal-rich guts of wood-feeding termites

Termites play important roles in lignocellulose and humus turnover in diverse terrestrial ecosystems, and are significant sources of global atmospheric methane and carbon dioxide. All known termite species engage in obligate, complex nutritional symbioses with their gut microbes to carry out such processes. Several hundred microbial species, representing a broad phylogenetic and physiological diversity, are found within the well‐bounded, microliter‐in‐scale gut ecosystem of a given termite. However, most of these species have never been obtained in laboratory culture, and little can be said about their functional roles in the gut community or symbiosis. Herein, an unappreciated facet of the gut chemistry and microbiology of wood‐feeding termites is revealed: the redox metabolism of iron. Gut fluids from field‐collected termites contained millimolar amounts of ferrous iron and other heavy metals. When iron(III) hydroxides were amended to a filter paper diet of Zootermopsis nevadensis, a dampwood termite collected in the San Gabriel Mountains of Southern California, the specimens accumulated high levels of iron(II) in their guts. Additionally, iron was reduced at rapid initial rates in anoxic gut homogenates prepared from field‐collected Z. nevadensis specimens. A Clostridium sp. and a Desulfovibrio sp. were isolated from dilution‐to‐extinction enrichments of Z. nevadensis gut contents and were found to reduce iron(III), as did the termite gut spirochete Treponema primitia. The iron in the guts of wood‐feeding termites may influence the pathways of carbon‐ and electron‐flow, as well as microbial community composition in these tiny ecosystems of global importance.     

Cryptic termites avoid predatory ants by eavesdropping on vibrational cues from their footsteps

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.     

Foraging,recruitment and predation by Decamorium uelense (Sanstchi) (Formicidae: Myrmicinae) on termites in Southern Guinea Savanna,Nigeria

Summary The myrmicine ant Decamorium uelense is a predator of small termites which feed inside their food sources of roots and plant debris. The ant uses a specialized recruitment regime to secure its prey. A scout ant searches for foraging termites and returns to the nest where it recruits a column of 10–30 ants which attack and immobilize the termites. A mass recruitment phase is then instigated, with larger numbers of ants retrieving the prey. The major prey item is Microtermes (consumed at an annual rate of 632 termites m^-2), with other small termites accounting for 0.5% of the total predation. The annual predation of Microtermes in primary savanna woodland by D. uelense removes 74% of the standing population.     

Termite diversity in Neotropical dry forests of Colombia and the potential role of rainfall in structuring termite diversity

Termites are ecosystem engineers that play an important role in the biotransformation and re‐distribution of nutrients in soil. The dry forests are endemic repositories, but at same time, they are most threatened by extensive livestock and crop farming, fires, and climate change. In Colombia, the best‐protected dry forests are located in the north. The termite fauna of dry forests are poorly known. The aim was to identify the termite species occurring in tropical dry forests of the Colombian Caribbean coast in relation to diet and precipitation, temperature, elevation, and soil properties. A total of 32 species in 1,103 occurrences were found. Termitidae accounted for 78% of the species richness with the Anoplotermes‐group, Microcerotermes, and Nasutitermes being the dominant genera. Differences in species composition and abundance were found across sites. These differences may be linked to anthropogenic disturbance and polygyny and polydomy. Strikingly, our highest elevation site (334 m) had the highest species richness much higher than the two lower elevation sites. This implies an inversion of the common elevation‐diversity gradient, also found for termites which can be explained by increasing precipitation with elevation in the dry forest. An analysis of termite species richness at the global scale confirms that termite species richness correlates positively with rainfall. Hence, rainfall seems to positively affect termite diversity. In line, the studied Colombian tropical dry forests had low diversity compared to rain forests. A decline of species‐rich soil‐feeding termites with increasing aridity may explain why the highest termite diversity occurs in humid tropical rain forests. Abstract in Spanish is available with online material.     

Termite diversity along a land use intensification gradient in a semi-arid savanna

Disturbance, particularly agricultural expansion is one of the major threats to the biodiversity and ecological functions of tropical and sub-tropical ecosystems. In this regard, we examined changes in the species richness, abundance, and diversity of termites across different disturbance treatments in a sub-tropical semi-arid savanna in south eastern Zimbabwe. Nine transects (100?×?2 m) representing three habitat disturbance treatments (primary woodland; grazing area; agricultural field) were sampled for termites using a rapid biodiversity assessment protocol. Termites were more abundant and species-rich in primary woodland and grazing area than in the agricultural field. Twelve termite species from three sub-families were present, with Microtermes sp. constituting 35% of the identified termite species. Termite feeding group structure differed significantly among land-use types, and of all termites present, wood-feeding termites were the most abundant while soil-feeders were rare in the agricultural field. In conclusion the observed pattern in termite species richness and relative abundance indicates that termites are very resilient to natural disturbance and might actually benefit from some natural disturbances like they did in the grazing area of this study, but they are not resilient to extreme anthropogenic disturbance. Although there was no notable difference in termite species richness and relative abundance between agricultural field and primary woodland, the pattern observed across the three sites may be potential support for the IDH suggesting that intermediate levels of physical disturbance intensity influence the structure and functioning of termite assemblages in semi-arid savanna.     

Drought and presence of ants can influence hemiptera in tropical leaf litter

Climate change is predicted to impact tropical rain forests, with droughts becoming more frequent and more severe in some regions. We currently have a poor understanding of how increased drought will change the functioning of tropical rain forest. In particular, tropical rain forest invertebrates, which are numerous and biologically important, may respond to drought in different ways across trophic levels. Ants are a diverse group that carry out important ecosystem processes, shaping ecosystem structure and function through predation and competition, which can influence multiple trophic levels. Hemiptera are a mega-diverse order, abundant in tropical rain forests and are ecologically important. To understand the roles of ants in exerting predation and competition pressure on invertebrates in tropical rain forests during drought and a post-drought period, we established a large-scale ecosystem manipulation experiment in Maliau Basin Conservation Area in Malaysian Borneo, suppressing the activity of ants on four 0.25 ha plots over a two-year period. We sampled hemipterans found in the leaf litter during a drought (July 2015) and a post-drought period (September 2016) period. We found significant shifts in the assemblage of hemipterans sampled from the leaf litter following ant suppression. Specifically, for ant-suppression plots, the species richness and abundance of herbivorous hemipterans increased only during the post-drought period. For predatory hemipterans, abundance increased with ant-suppression regardless of drought conditions, and we found marginal evidence for a species richness increase during the post-drought period with little or no change in the drought period. These results illustrate how ants in tropical forests structure invertebrate communities and how these effects may vary with climatic variation.