Diversity of fungus‐growing termites (Macrotermes) and their fungal symbionts (Termitomyces) in the semiarid Tsavo Ecosystem,Kenya

Fungus‐growing termites of the subfamily Macrotermitinae together with their highly specialized fungal symbionts (Termitomyces) are primary decomposers of dead plant matter in many African savanna ecosystems. The termites provide crucial ecosystem services also by modifying soil properties, translocating nutrients, and as important drivers of plant succession. Despite their obvious ecological importance, many basic features in the biology of fungus‐growing termites and especially their fungal symbionts remain poorly known, and no studies have so far focused on possible habitat‐level differences in symbiont diversity across heterogeneous landscapes. We studied the species identities of Macrotermes termites and their Termitomyces symbionts by excavating 143 termite mounds at eight study sites in the semiarid Tsavo Ecosystem of southern Kenya. Reference specimens were identified by sequencing the COI region from termites and the ITS region from symbiotic fungi. The results demonstrate that the regional Macrotermes community in Tsavo includes two sympatric species (M. subhyalinus and M. michaelseni) which cultivate and largely share three species of Termitomyces symbionts. A single species of fungus is always found in each termite mound, but even closely adjacent colonies of the same termite species often house evolutionarily divergent fungi. The species identities of both partners vary markedly between sites, suggesting hitherto unknown differences in their ecological requirements. It is apparent that both habitat heterogeneity and disturbance history can influence the regional distribution patterns of both partners in symbiosis.     

Strong positive effects of termites on savanna bird abundance and diversity are amplified by large herbivore exclusion

Vast areas of the African savanna landscapes are characterized by tree‐covered Macrotermes termite mounds embedded within a relatively open savanna matrix. In concert with termites, large herbivores are important determinants of savanna woody vegetation cover. The relative cover of woody species has considerable effects on savanna function. Despite the potentially important ecological relationships between termite mounds, woody plants, large herbivores, and birds, these associations have previously received surprisingly little attention. We experimentally studied the effects of termites and large herbivores on the avian community in Lake Mburo National Park, Uganda, where woody vegetation is essentially limited to termite mounds. Our experiment comprised of four treatments in nine replicates; unfenced termite mounds, fenced mounds (excluding large mammals), unfenced adjacent savanna, and fenced savanna. We recorded species identity, abundance, and behavior of all birds observed on these plots over a two‐month period, from late dry until wet season. Birds used termite mounds almost exclusively, with only 3.5% of observations occurring in the treeless intermound savanna matrix. Mean abundance and species richness of birds doubled on fenced (large herbivores excluded) compared to unfenced mounds. Feeding behavior increased when large mammals were excluded from mounds, both in absolute number of observed individuals, and relative to other behaviors. This study documents the fundamental positive impact of Macrotermes termites on bird abundance and diversity in an African savanna. Birds play crucial functional roles in savanna ecosystems, for example, by dispersing fruits or regulating herbivorous insect populations. Thus, the role of birds in savanna dynamics depends on the distribution and abundance of termite mounds.     

Prospects for Bonobo Insectivory: Lui Kotal, Democratic Republic of Congo

Chimpanzees (Pan troglodytes) are well-known to eat invertebrates, especially social insects, across Africa, but allopatric bonobos (P. paniscus) are not. Bonobo insectivory is sparsely documented and apparently sporadic. However, the availability to bonobos of social insect prey and raw materials with which to make tools to exploit them is unknown. Here, we test a set of hypotheses that relates to questions of presence, abundance, density, and distribution of taxa that Pan consume and of vegetation suitable for making extractive foraging tools. We worked at Lui Kotal, Democratic Republic of Congo, where unprovisioned bonobos live in intact forest, far from villages. We collected insect and fecal specimens, transected for prey and assessed raw materials, and monitored mounds of Macrotermes. All but 1 of the major taxa of relevant termites, ants, and (stinging) honey bees were present. The 3 main taxa of insects that chimpanzees elsewhere eat —Macrotermes (fungus-growing termites), Dorylus (Anomma; army or driver ants), and Apis (honey bees)— were abundant and widespread, and usually at densities exceeding those at well-known chimpanzee study-sites. Similarly, woody and nonwoody vegetation suitable for making fishing probes was common at mounds of Macrotermes. There is no obvious ecological reason why bonobos should not use elementary technology in extractive foraging, e.g., termite-fish, ant-fish, ant-dip, honey-dip, to obtain social insects.     

The use of kairomones by Megaponera foetens (Fab.) (Hymenoptera: Formicidae) in the detection of its termite prey

Scout ants of the obligate termite predator, Megaponera foetens, respond to cues in fresh soil sheeting laid down by foraging termites by returning to their nests laying a recruitment trail. Effective cues are found in the sheeting of Macrotermes and Odontotermes, but not Microtermes and Ancistrotermes. They can be extracted using organic solvent, but have disappeared from the sheeting after about. 24 h. Solvent extracts of minor worker termites have the same effect on scout ants as fresh sheeting. The ability of M. foetens to detect fresh soil sheeting may increase the efficiency of predation on Macrotermes and Odontotermes species.     

You eat what you find – Local patterns in vegetation structure control diets of African fungus‐growing termites

Fungus‐growing termites and their symbiotic Termitomyces fungi are critically important carbon and nutrient recyclers in arid and semiarid environments of sub‐Saharan Africa. A major proportion of plant litter produced in these ecosystems is decomposed within nest chambers of termite mounds, where temperature and humidity are kept optimal for the fungal symbionts. While fungus‐growing termites are generally believed to exploit a wide range of different plant substrates, the actual diets of most species remain elusive. We studied dietary niches of two Macrotermes species across the semiarid savanna landscape in the Tsavo Ecosystem, southern Kenya, based on carbon (C) and nitrogen (N) stable isotopes in Termitomyces fungus combs. We applied Bayesian mixing models to determine the proportion of grass and woody plant matter in the combs, these being the two major food sources available for Macrotermes species in the region. Our results showed that both termite species, and colonies cultivating different Termitomyces fungi, occupied broad and largely overlapping isotopic niches, indicating no dietary specialization. Including laser scanning derived vegetation cover estimates to the dietary mixing model revealed that the proportion of woody plant matter in fungus combs increased with increasing woody plant cover in the nest surroundings. Nitrogen content of fungus combs was positively correlated with woody plant cover around the mounds and negatively correlated with the proportion of grass matter in the comb. Considering the high N demand of large Macrotermes colonies, woody plant matter seems to thus represent a more profitable food source than grass. As grass is also utilized by grazing mammals, and the availability of grass matter typically fluctuates over the year, mixed woodland‐grasslands and bushlands seem to represent more favorable habitats for large Macrotermes colonies than open grasslands.     

Effects of Erosion from Mounds of Different Termite Genera on Distinct Functional Grassland Types in an African Savannah

A key aspect of savannah vegetation heterogeneity is mosaics formed by two functional grassland types, bunch grasslands, and grazing lawns. We investigated the role of termites, important ecosystem engineers, in creating high-nutrient patches in the form of grazing lawns. Some of the ways termites can contribute to grazing lawn development is through erosion of soil from aboveground mounds to the surrounding soil surface. This may alter the nutrient status of the surrounding soils. We hypothesize that the importance of this erosion varies with termite genera, depending on feeding strategy and mound type. To test this, we simulated erosion by applying mound soil from three termite genera (Macrotermes, Odontotermes, and Trinervitermes) in both a field experiment and a greenhouse experiment. In the greenhouse experiment, we found soils with the highest macro nutrient levels (formed by Trinervitermes) promoted the quality and biomass of both a lawn (Digitaria longiflora) and a bunch (Sporobolus pyramidalis) grass species. In the field we found that soils with the highest micro nutrient levels (formed by Macrotermes) showed the largest increase in cover of grazing lawn species. By linking the different nutrient availability of the mounds to the development of different grassland states, we conclude that the presence of termite mounds influences grassland mosaics, but that the type of mound plays a crucial role in determining the nature of the effects.     

The distribution,abundance, and the effects of fire on mound building termites (Trinervitermes and Cubitermes spp., Isoptera: Termitidae) in northern guinea savanna West Africa

Summary Termite mound densities in typical guinea savanna, Detarium, and grassland (boval) habitats in northern guinea savanna were determined by random quadratting of 2–3 sites in each habitat (100, 10x10 m quadrats per habitat). Dominant species in guinea savanna were T. geminatus (46 mounds ha^-1) and T. oeconomus (21 mounds ha^-1), in Detarium T. geminatus (59 mounds ha^-1) and C. curtatus (45 mounds ha^-1) and in boval C. curtatus (72 mounds ha^-1) and T. geminatus (22 mounds ha^-1). Only C. curtatus densities and total densities differed significantly between sites within habitats, but all species differed significantly in abundance between habitats. The composition of each community was related to general environment but no particular environmental variable was shown to be a major determinant of termite distribution. Evidence for the limitation of termite populations was obtained from indirect evidence of competition between colonies in Detarium, and by experimental manipulation of fire regimes in the typical guinea savanna habitat. Harvester termites increased four-five fold over two years in fire-protected plots as a result of increased food supplies. Total termite densities in the fire-protected community equilibrated to the new population density (100 mounds ha^-1) after only two-three years.     

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).     

Termite mound density and distribution patterns across three land-use types in the Vhembe District of the Limpopo Province,South Africa

This study investigated the effect of land-use on density and distribution patterns of termite mounds. A total area of 12 ha was investigated using four 1 ha plots from each of three land-use types (mango orchards, maize fields and communal rangelands). A total of 297 mounds from four termite species were recorded. Plotted GIS coordinates for each mound in ArcMap showed a random distribution pattern in all land-use types. The mean number of mounds per hectare was significantly higher (p < 0.001) in communal rangelands (52.5 ± 1.21), than in maize fields (14.75 ± 3.15) and mango orchards (7.5 ± 0.87), and dominated by small-sized mounds of Trinervitermes sp. Few mounds of Odontotermes sp. were found. Mounds of the edible termites, Macrotermes natalensis and M. falciger, were found in all land-use types, with the highest density for both species being in maize fields. Although the mound height for both species was similar, mound circumference for M. falciger was significantly larger (p < 0.001) which may limit land available for agricultural use. Density of mounds was influenced by land-use which may lead to changes in termite ecosystem functioning and availability of termites as a free source of protein.     

Termite Mounds Increase Functional Diversity of Woody Plants in African Savannas

Fine-scale spatial heterogeneity influences biodiversity and ecosystem productivity at many scales. In savanna systems, Macrotermes termites, through forming spatially explicit mounds with unique woody plant assemblages, emerge as important sources of such heterogeneity. Despite a growing consensus regarding the importance of functional diversity (FD) to ecosystem processes, no study has quantified how termite mounds affect woody plant FD. We address whether termite mounds alter the distribution of functional traits, and increase FD of woody plant communities within Africa’s largest savanna woodland, the 2.7 million km² miombo system. Using plant traits that change according to soil resources (for example, water and nutrients), and disturbance (for example, fire and elephant herbivory), we identified response functional groups and compared relative representation of these groups between mound and matrix habitats. We also asked whether mound and matrix habitats differed in their contribution to FD within the system. Although species representing most functional groups were found in both mound and matrix habitats, relative abundance of functional groups differed between mound and matrix. Mound plant assemblages had greater response diversity to soil resources than matrix plots, but there was no difference in response diversity to disturbance. High trait values on mounds included tree height, leaf nitrogen, phosphorus, and palatability. Species with root ectomycorrhizae dominated the matrix. In conclusion, these small patches of nutrient-enriched substrate emerge as drivers of FD in above-ground woody plant communities.     

Prevalence of a koinobiont endoparasitoid Misotermes mindeni (Diptera: Phoridae) in colonies of the fungus-growing termite Macrotermes gilvus (Blattodea: Termitidae) in Malaysia

A survey of the infestation rate of colonies of Macrotermes gilvus (Hagen) (Termitidae: Macrotermitinae) with the koinobiont endoparasitoid Misotermes mindeni Disney & Neoh (Diptera: Phoridae) was conducted in Malaysia from September 2009 to January 2011 in the states of Kedah, Penang, Perak, Selangor, Kuala Lumpur, Johor, Terengganu, and Sarawak. Of the 1,125 M. gilvus mounds surveyed, 12.4% contained termites parasitized by M. mindeni and these mounds occurred only in the states of Penang and Perak. High frequencies of mounds containing parasitized termites were found at sites in Penang: Bayan Lepas (21.1%), Minden Campus of Universiti Sains Malaysia ([USM]; 24.5%), Teluk Bahang (28.0%), and Bukit Mertajam (35.0%); the lowest frequency (4.0%) was recorded from Gelugor. The parasitized colonies at all sites were classified as healthy, with exception of several from the Minden Campus of USM (96.4% healthy) and Ayer Itam (87.5% healthy). Most parasitized colonies (71.2%) had a low level of M. mindeni infestation. Only 16.7 and 12.1% of the infested colonies had moderate or high parasite infestation levels, respectively. The height of infected mounds was significantly higher than that of the healthy mounds, but there was no difference between the mound diameters of infested and uninfested mounds. Parasite infestation level was not significantly correlated with mound height or mound diameter. The ambient light intensity at sites with infested mounds was significantly lower than that of uninfested mounds. There was also a significant negative relationship between light intensity and degree of parasitism.     

Termites, Large Herbivores, and Herbaceous Plant Dominance Structure Small Mammal Communities in Savannahs

Large herbivores and termites are important functional groups in African savannahs. Both groups affect small mammals, which are also important determinants for savannah structure and function. Because vegetation on Macrotermes mounds are preferentially grazed by large herbivores, and mounds represent resource-rich distinct habitat patches for small mammals in relatively resource-poor savannahs, termite mounds are ideal sites for studies of how grazing by large mammals and productivity affect communities of small mammals. We conducted an experiment in Lake Mburo National Park, Uganda, with four treatments: large vegetated Macrotermes mounds (with and without large herbivores) and adjacent savannah areas (with and without large herbivores). We replicated the treatment blocks nine times and trapped small mammals regularly over a period of almost 2 years. Small mammal species assemblages differed considerably between mounds and savannah areas. Grazing had a substantial effect on small mammal species assemblages in the resource-poor savannah, but not in the relatively resource-rich termitaria. Small mammal species abundance, biomass, and richness were higher on termite mounds than adjacent savannah areas. Excluding large herbivores caused a major increase in species abundance, biomass, and richness both on savannah and termitaria. Herbaceous plant species evenness was an important determinant of the small mammal community. Small mammal biomass increased with high plant dominance, indicating that a few dominant plant species are important for biomass production of small mammals. Small mammal diversity was not related to any of the treatments, but increased with plant species evenness as well as richness. Fencing increased species dominance in the small mammal community on both savannah and termitaria, probably because competitive patterns shift from inter-guild (that is, between large and small mammals) to intra-guild (that is, between small mammals) when large mammals are excluded. The study highlights the complex interactions among large herbivores, termites, herbaceous plants, and small mammals in African savannahs. When studying the structure and function of small mammal communities it is therefore important to consider several coexisting functional groups.     

Species abundance and habitat differences in biomass of subterranean termites (Isoptera) in the wheatbelt of Western Australia

The distribution and abundance, and habitat differences in biomass of subterranean termites, were assessed through soil trenching for woodland, mallee and heath habitats in the central wheatbelt of Western Australia. Over an 11 month sampling period, there were no significant habitat differences in biomass. Mean dry biomass in the surficial layer of soil (per 5000 cm³) averaged 46 mg in woodland, 28 mg in mallee, and 23 mg in heath. Termite biomass peaked in September, with 141 mg for woodland, 83 mg for mallee and 47 mg for heath (per 5000 cm³). Soil moisture and termite activity near the surface were positively correlated. A total of 36 species of termites, comprising 11 genera, were identified, and species abundance within and across habitats differed significantly.     

Relation between termite numbers and the size of their mounds

This article provides a meta-analysis of quantitative data available in literature regarding the relation between termite numbers and the volume of their mounds for 24 species belonging to 13 genera. The leading question behind this analysis is: “how do the respiratory gas exchanges regulate the size of termite mounds?” This question is answered through the analysis of the log–log regression between the volume of the mound and the number of inhabitants. The most confident data support the hypothesis of a respiratory regulation that can be achieved through a relation between the termite numbers and (1) the volume of their mounds (slope of the regression near 1, Noditermes), (2) the surface of the outer walls of their mounds (slope of the regression near 0.67, Termitinae and Nasutitermes) or (3) a compromise between the surface of the outer walls of their mounds and some linear structures of their nests (slope of the regression between 0.67 and 0.33, Trinervitermes and Macrotermes). The way this is achieved is linked with the architecture of the mound. A confident relation was found between the number of individuals and the epigeal volume of their mounds for 18 species for which the most reliable data were provided. Three more accurate models are proposed for estimating the termite population based on the nest material and architecture and on the size of the termites.     

The role of termites in an equatorial rain forest ecosystem of West Malaysia

Summary To estimate the rate of consumption of leaf litter by termites on the forest floor of Pasoh Forest Reserve, Negeri Sembilan, West Malaysia, newly fallen leaves were marked and distributed on the ground. The loss of leaf area due to termites was determined either photometrically or visually. An average of 1.70% of the total surface area of the leaf litter disappeared per week in experiment 1 and 1.25% in experiment 2 in Plot 1, and 2.9% per day in other plots located near the mounds of Macrotermes carbonarius. The amount of leaf litter accumulation in the Ao layer was estimated at about 2.3 t/ha at Plot 1, so it was likely that an amount equivalent to about 32% of the daily leaf-litter fall was transported by M. carbonarius to their mounds in experiment 1 and 22% in experiment 2. It was considered that the termites had an important role in the detritus food chain of the ecosystem.Japanese Contribution No. 13, IBP Pasoh Project     

Contribution of the Western Australian wheatbelt termite,Drepanotermes tamminensis (Hill), to the soil nutrient budget

The role of soil modification by the mound-building termite,Drepanotermes tamminensis (Hill), was studied during 1991 in the Durokoppin Nature Reserve, Western Australia. Soil chemical parameters were quantified for ‘soils’ in nests and for surrounding soil in both a Wandoo (Eucalyptus capillosa) woodland and a Casuarina (Allocasuarina campestris) shrubland plot. All ‘soils’ in nests were more acidic than the surrounding soil within each study plot. Generally, nutrient levels in the nested soils were higher than the un-nested soil within each study plot and were also higher in the woodland than in the shrubland plot. Depending on the nuttient concerned, the nested soil contained between 0.3 and 21.9% of the total nutrient load per hectare within each study plot. The quantities of nutrients per hectare in termite mounds were higher in the woodland than in the shrubland plot. It is concluded that mounds of this species of termite form a significant bank of nutrients, although time for release of such nutrients depends on the degree of erosion and on the longevity of mounds.     

Revision of the termite-parasitizing genus Dicranopteron (Diptera: Phoridae)

Dicranopteron Schmitz is removed from the Metopininae and three new species are described. All species are represented by females only and come from Odontolermes and Macrotermes nests in the Orient. A key:to species) is provided. One species was observed ovipositing in the abdomens of the young stages of O. takensis.     

A model of litter harvesting by the Western Australian wheatbelt termite,Drepanotermes tamminensis (Hill), with particular reference to nutrient dynamics

A series of papers have been published which describe the influence of vegetation and soil type on the Western Australian wheatbelt termite,Drepanotermes tamminensis (Hill), and also on its litter harvesting levels and contribution to the soil nutrient budget. This paper integrates these findings by means of a computer simulation model. The model consists of three modules which respectively describe the dynamics of litter on the ground, the dynamics of litter within termite mounds and how these in turn influence nutrient loads within the habitat. The outputs of the model suggest that this litter harvesting termite plays an important role in the nutrient dynamics of the area and it provides an estimate of the unmeasured variable, litter consumed in mounds by termites, which is consistent with measurements for other termite species with similar feeding habits.     

Use of termites, Reticulitermes virginicus, as a springboard in the invasive success of a predatory ant, Pachycondyla (=Brachyponera) chinensis

Invasive ant species have general diet and nest requirements, which facilitate their establishment in novel habitats and their dominance over many native ants. The Asian needle ant, Pachycondyla chinensis, native throughout Australasia was introduced to the southeastern US where it has become established in woodland habitats, nests in close proximity to and consumes subterranean termites (Rhinotermitidae). P. chinensis do not occur in habitats lacking Rhinotermitidae. We suggest that subterranean termites are critical for P. chinensis success in new habitats. We demonstrate that P. chinensis is a general termite feeder, retrieving Reticulitermes virginicus five times more often than other potential prey near P. chinensis colonies. Odors produced by R. virginicus workers, as well as other potential prey, attract P. chinensis. Furthermore, P. chinensis occupy R. virginicus nests in the lab and field and display behaviors that facilitate capture of R. virginicus workers and soldiers. Termites are an abundant, high quality, renewable food supply, in many ways similar to the hemipteran honeydew exploited by most other invasive ant species. We conclude that the behavior of P. chinensis in the presence of termites increases their competitive abilities in natural areas where they have been introduced.     

Chimpanzees' (Pan troglodytes) choice of prey among termites (macrotermitinae) in western Tanzania

This study examines factors affecting chimpanzees' preying on termites, especially their choice of prey species. Regular surveys of a large sample of termite-mounds in the Mahale Mountains, Tanzania, confirmed that chimpanzees of B Group obtainedMacrotermes, which were relatively uncommon, by the use of fishing-tools, but ignored the much more abundantOdontotermes. This preference appeared to reflectMacrotermes' more extended swarming period, greater accessibility, larger size, and less noxious taste than the soldiers ofOdontotermes. Similar factors probably underly the chimpanzees' choice ofMacrotermes at Gombe, butPseudacanthotermes spiniger are also common there, and their absence from the diet is less easily explained. In contrast, the chimpanzees of K Group at Mahale prey mainly onP. spiniger;Macrotermes are apparently absent, andP. spiniger appear to be more abundant, accessible, and palatable than the alternativeOdontotermes.