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.     

Studies on a population of large termite mounds in Uganda

• 1 Two study-sites near Kampala were mapped and all of the termite mounds within them were measured. Changes were recorded in the three succeeding years. A method was developed for estimating mound volumes from height and diameter.
• 2 Macrotermes bellicosus mounds reached a volume of about 2.4 m³ in 3 years after which they grew more slowly to a maximum from 4 to 6 m³. Growth in volume during the first 3 years was approximately linear and its rate did not vary seasonally, probably because growth is related to temperature which was nearly constant throughout the year.
• 3 M.bellicosus mounds had an average length of life of about 10 years, although colonies survived on average only 4 years. The corresponding figures for Pseudacanthotermes spp. were 5 and 2 1/2 years. Some mounds were recolonized after the death of the original colony.
• 5 Fresh earth was brought up at a minimum rate of about 10 m³ ha^-1 yr^-1 but more was lost by erosion and destruction and the populations of mounds were decreasing.
• 4 The number of ventilation shafts in M.bellicosus mounds increased with the volume of the mound. Many mounds, both dead and alive, supported vegetation.
• 6 Local opinion is that Pseudacanthotermes are harmless but M.bellicosus is regarded as a pest and attempts are made to destroy its mounds.

     

Spatial variability and abiotic determinants of termite mounds throughout a savanna catchment

Termite mounds contribute to the spatial heterogeneity of ecological processes in many savannas, but the underlying patterns and determinants of mound distributions remain poorly understood. Using the Carnegie Airborne Observatory (CAO), we mapped the distribution of termite mounds across a rainfall gradient within a river catchment (～ 27 000 ha) of the Kruger National Park, South Africa. We assessed how different factors were associated with the distribution and height of termite mounds at three spatial scales: the entire catchment, among three broad vegetation types, and on individual hillslope crests. Abiotic factors such as the underlying geology and mean annual precipitation shaped mound densities at broad scales, while local hillslope morphology strongly influenced mound distribution at finer scales, emphasising the importance of spatial scale when assessing mound densities. Fire return period had no apparent association with mound densities or height. Mound density averaged 0.46 mounds ha^?1, and exhibited a clustered pattern throughout the landscape, occurring at relatively high densities (up to 2 mounds ha^?1) on crests, which are nutrient‐poor elements of the landscape. Mounds exhibited significant over‐dispersion (even spacing) at scales below 60 m so that evenly spaced aggregations of termite mounds are embedded within a landscape of varying mound densities. The tallest mounds were found in dry savanna (500 mm yr^?1) and were positively correlated with mound density, suggesting that dry granitic savannas are ideal habitat for mound‐building termites. Mound activity status also varied significantly across the rainfall gradient, with a higher proportion of active (live) mounds in the drier sites. The differential spacing of mounds across landscapes provides essential nutrient hotspots in crest locations, potentially sustaining species that would otherwise not persist. The contribution to biodiversity and ecosystem functioning that mounds provide is not uniform throughout landscapes, but varies considerably with spatial scale and context.     

Differences between bacterial communities in the gut of a soil-feeding termite (Cubitermes niokoloensis) and its mounds

In tropical ecosystems, termite mound soils constitute an important soil compartment covering around 10% of African soils. Previous studies have shown (S. Fall, S. Nazaret, J. L. Chotte, and A. Brauman, Microb. Ecol. 28:191-199, 2004) that the bacterial genetic structure of the mounds of soil-feeding termites (Cubitermes niokoloensis) is different from that of their surrounding soil. The aim of this study was to characterize the specificity of bacterial communities within mounds with respect to the digestive and soil origins of the mound. We have compared the bacterial community structures of a termite mound, termite gut sections, and surrounding soil using PCR-denaturing gradient gel electrophoresis (DGGE) analysis and cloning and sequencing of PCR-amplified 16S rRNA gene fragments. DGGE analysis revealed a drastic difference between the genetic structures of the bacterial communities of the termite gut and the mound. Analysis of 266 clones, including 54 from excised bands, revealed a high level of diversity in each biota investigated. The soil-feeding termite mound was dominated by the Actinobacteria phylum, whereas the Firmicutes and Proteobacteria phyla dominate the gut sections of termites and the surrounding soil, respectively. Phylogenetic analyses revealed a distinct clustering of Actinobacteria phylotypes between the mound and the surrounding soil. The Actinobacteria clones of the termite mound were diverse, distributed among 10 distinct families, and like those in the termite gut environment lightly dominated by the Nocardioidaceae family. Our findings confirmed that the soil-feeding termite mound (C. niokoloensis) represents a specific bacterial habitat in the tropics.     

The architecture of termite mounds: a result of a trade-off between thermoregulation and gas exchange?

We examined the influence of gas exchange on the architectureof termite mounds. In Comoé National Park (Côted'Ivoire), Macrotermes bellicosus builds, as an adaptation toambient temperature conditions, differently shaped mounds inthe shrub savanna and the gallery forest. Previous studies suggestedthat there might be a constraint that limits the degree of thermalinsulation of the interior (i.e., nest) of the mounds in environmentswith relatively low ambient temperatures. This factor causes,in proximate terms, suboptimal low nest temperatures and ultimatelyleads to reduced reproductive success in the gallery forest.In this study, we examined whether the necessity for gas exchangemight constrain mound architecture. We measured CO₂ concentrationsin the air channels of mounds in different habitats and undermanipulated temperature regimes. During both the dry and therainy season we found higher CO₂ concentrations in mounds ofthe gallery forest than in mounds of the savanna. Additional measurementsin forest mounds, architecturally resembling those of the savanna dueto an experimental increase in ambient temperatures, revealedlower CO₂ concentrations than unmanipulated mounds in this habitat. Generally,concentrations were higher during the rainy season comparedto the dry season and lower during day than during night. Summarizingthese results we present a model that illustrates this trade-offbetween thermoregulation and gas exchange under different temperatureregimes. Both factors together result in different mound architecturesunder different environmental temperatures and may finally limitthe distribution of this species.     

Effects of controlled livestock grazing and annual prescribed fire on epigeal termite mounds in a savannah woodland in Burkina Faso

The diversity and abundance of epigeal termite mounds were investigated in response to controlled livestock grazing and annual prescribed fire in a Sudanian savannah-woodland in Tiogo State Forest. Sampling of termite mounds was carried out in 4×4 subplots of 0.25 ha in a split-plot experimental design during the rainy season in 2002. There were two main plots of which one was fenced to exclude livestock grazing and the second exposed to grazing. Each of the main plots included 4 subplots with annual prescribed fire since 1992 and 4 subplots without fire. Data were collected on the number and characteristics of termite mounds. A mean density of 698 mounds ha⁻¹ was recorded. Mounds built by Trinervitermes spp. were the most abundant followed by Cubitermes spp., Macrotermes subhyalinus and M. bellicosus. The large mound-builders Macrotermes spp. dominated the community interms of basal area (96% of the total) and above-ground volume (99%). The diversity of mound types was notaffected by livestock grazing and annual early prescribed fire (P>0.05). There was no statistical effect of livestock grazing on mound density, whereas a strong depressive effect of annual fire was observed for Trinervitermes spp. mound density (P=0.012). In this ecosystem, annual prescribed fire appeared to be the major determinant for termite mound abundance. Received 2 February 2007; revised 23 October 2007 and 21 January 2008; accepted 22 Feburary 2008.     

Faunal makeup of wild bees and their flower utilization in a semi-urbanized area in central Japan

The species composition of wild bees and their flower utilization patterns were surveyed from April to November in 1996 in a semi‐urbanized area adjacent to Sugao Marsh, Ibaraki, central Japan. A total of 750 individuals belonging to 43 species in six families were collected. The most dominant family was Halictidae, for which 13 species and 251 individuals were collected. The most dominant species was Colletes patellatus (120 individuals) of the Colletidae. The results at Sugao were compared with those obtained from three other areas of Ibaraki Prefecture, which have similar climatic conditions, yet have different environmental characteristics in terms of human impact. The four sites in Ibaraki can be classified into two groups: the first comprising Sugao and Mito in cultivated and/or human‐dwelling areas, and the second comprising Yamizo and Gozen’yama, in forest areas with more natural elements. The number of species at Sugao was the smallest among the four study sites. On the other hand, the values for species evenness at Sugao were the second‐highest of the four study sites. These findings show that the different characteristics of different bee communities reflect their local environmental conditions, including their floral compositions. The bees visited 36 flower species in 20 families, and 70.7% of all individuals studied visited Compositae flowers. The heavy utilization of composite flowers is possibly because of the existence of a simplified flora consisting of a few dominant composite plant species. Among these plants, Solidago altissima and Lactuca indica made large contributions to supporting autumn bees, especially Colletes patellatus and Colletes perforator, which are solitary and oligolectic on Compositae.     

The causes of spatial patterning of mounds of a fungus-cultivating termite: results from nearest-neighbour analysis and ecological studies

Little is known about processes regulating population dynamics in termites. We investigated the distribution of mound-colonies of the fungus-cultivating termite Macrotermes bellicosus (Smeathman) in two habitats in the Comoé National Park (Côte d'Ivoire) with nearest-neighbour analysis differentiating between different age classes. These results were compared with ecological data on processes influencing population dynamics. High mound densities were recorded in shrub savannah while only a few mounds were found in gallery forest. Mounds were distributed randomly in both habitats when all mounds were considered together, and when inhabited and uninhabited mounds were treated separately. However, distinctive non-random patterns were revealed in the savannah when we distinguished between different age classes. Small, young colonies were aggregated when they coexisted with larger, older colonies, which were more regularly distributed. This indicates that the distribution of older colonies is influenced by intraspecific competition whereas that of younger colonies is influenced by opposing factors that lead to aggregation. This is in accordance with ecological data. Food is a limiting resource for large colonies, while patchily distributed appropriate microclimatic conditions seem to be more important for young colonies. Colonies that had formerly coexisted (i.e. living colonies and recently dead colonies) showed aggregated, random and regular distribution patterns, suggesting several causes of mortality. Colonies that had never had contact with each other were randomly distributed and no specific regulation mechanism was implicated. These results show that different age classes seem to be regulated by different processes and that separation between age classes is necessary to reveal indicative spatial patterns in nearest-neighbour analysis.     

Thermoregulation of termite mounds: what role does ambient temperature and metabolism of the colony play?

Summary: Termites are well known for their ability to regulate the environment of their nest such as temperature and humidity. The influence of fluctuating ambient temperature and mound characteristics on mean nest temperature and daily fluctuation of nest temperature was analysed quantitatively in the fungus-cultivating, mound-building termite Macrotermes bellicosus (Macrotermitinae) in the savanna of the Comoé National Park (Côte d'Ivoire). Additionally, the nest temperatures of inhabited and uninhabited mounds were compared to analyse the contribution of ambient temperature to nest temperature in relation to metabolic heat production of the termites and their fungi. Mound structure alone resulted in a relatively constant nest temperature. Abiotic heat production via solar radiation alone yielded nest temperatures that corresponded to mean ambient temperatures. However, only the production of metabolic heat by the termites and the fungi increased these temperatures to the actual nest temperature. Therefore, and due to the high heat capacities of the mounds, large colonies (mound height above 2.0 m) had higher nest temperatures than smaller ones. Only large colonies attain constant nest temperatures of 30 °C that are largely independent from ambient temperatures and optimal for the growth and development of the termites and their fungi.     

中国野生稻遗传资源的保护及其在育种中的利用

我国有三种野生稻,即普通野生稻(Oryza rufipogon)、药用野生稻(O.officinalis)和瘤粒野生稻(O.meyeriana)。这三种野生稻均被列为国家二级保护植物(渐危种)。调查结果表明,野生稻由于其自然群落大量丧失而濒危,濒危程度为普通野生稻>药用野生稻>瘤粒野生稻。造成濒危的主要原因是人为的破坏活动。人类的经济活动导致了野生稻生境丧失、生境质量不断恶化、栖息地越来越少;人类的活动也导致了外来种的入侵。目前,对野生稻的保护措施主要有就地保护(原地保护或原位保护)和迁地保护(易地保护或异位保护)。易地保护包括以种子保存的种质厍、以种茎保存的种质圃和以器官培养物作为材料的超低温保存。野生稻具有许多优良特性,如特强的耐寒性、高的抗病虫性、优质蛋白质含量高、功能叶片耐衰老的特异性、特强的再生性、良好的繁茂性及生长优势等等,这些优良特性已被广泛用于水稻常规育种和杂交育种中,并取得了巨大的社会效益和经济效益。有关野生稻生物技术方面的研究,如花药培养、原生质培养、体细胞杂交和基因工程等方面已取得了较大的进展。野生稻将在水稻育种中发挥越来越重要的作用。     

Soil particle accumulation in termite (Macrotermes bellicosus) mounds and the implications for soil particle dynamics in a tropical savanna Ultisol

This study investigated the influence of mound-building termites on soil particle dynamics on the land surface and in soil-forming processes by examining the amount of soil particles in mound structures of Macrotermes bellicosus in a highly weathered Ultisol of tropical savanna. Soil particle turnover via the mounds was estimated using particle stock data and soil turnover data from previous studies. A 4-ha study plot with six mounds of relatively uniform shape and size was investigated. Soil mass constituting the mounds was 6,166 ± 1,581 kg mound⁻¹ within which the mound wall and nest body accounted for 5,002 ± 1,289 and 1,164 ± 293 kg, respectively. The mound wall contained a significantly larger amount of clay (252 ± 9.97 g kg⁻¹) balanced with a lower sand content (676 ± 26.5 g kg⁻¹) than in the adjacent surface (Ap1) horizon, (46.4 ± 12.8 g clay kg⁻¹; 866 ± 83.2 g sand kg⁻¹); the nest body had much higher clay content (559 ± 51.0 g kg⁻¹) but less sand (285 ± 79.2 g kg⁻¹) than the mound wall. As a result, the mounds of M. bellicosus accumulated clay of 2,874 ± 781 kg ha⁻¹ (corresponding to 2.52% of clay stock in the Ap1 horizon) along with an estimated clay turnover rate of 169 kg ha⁻¹ year⁻¹. These findings suggest a positive feedback effect from termite mound-building activity on soil particle dynamics in tropical savanna ecosystems: M. bellicosus preferentially use subsoil material for mound construction, resulting in relocation of illuvial clay in the subsoil to the land surface where clay eluviation from the surface soil and its illuviation in the subsoil are major soil-forming processes.     

Geophagic termite mounds as one of the resources for African elephants in Ugalla Game Reserve,Western Tanzania

Knowledge of the distribution and nutrient values of key resources supporting the survival of wildlife species is integral for an effective conservation planning and management of the species. In the Miombo ecosystem of the Ugalla Game Reserve, African elephants (Loxodonta africana Blumenbach 1797), eat soil, that is geophagy, from certain termite mounds. We mapped that all the geophagic termite mounds are exclusively situated in the flood plain. To understand why soils from some termite mounds are eaten, we collected and analysed soil samples from 10 geophagic termite mounds, seven nongeophagic termite mounds and 13 samples from the surrounding flood plain. Percentage of clay content did not differ significantly among the soil samples. Soils from geophagic termite mounds were richer in mineral elements compared with other soil samples. The results demonstrate that the driver for geophagic behaviour is related to rich mineral element contents found in geophagic termite mounds made of the mineral‐enriching termites (Macrotermes). Thus, geophagic termite mounds play a role in elephant's dietary needs and possibly influence their movement patterns in Ugalla, as the elephants cannot obtain enough minerals from their feeds. Geophagic termite mounds should be protected from potential destructive land uses, such as airstrip construction.     

Spatial distribution of ant mounds and effects on soil physical properties in wetlands of the Sanjiang plain, China

Ants constitute a dominant element of soil mesofauna due to their biomass, abundance, richness of species and distribution within terrestrial ecosystems. They are important regulators of soil aggregate structure as they translocate large amounts of soil from the bottom to the soil surface. In doing so, they form biogenic structures (BS) made up of aggregates of different sizes and characteristics, i.e. ant mounds. These BS have varying characteristics according to the ant species and the soil where they carry their activities. Ants are considered soil engineers because of their effects on soil properties, availability of resource and flow of energy and nutrients in soil. Thus, it is important to gain information on their distribution and abundance. Relatively little is known about the spatial distribution of mounds and their role in the soil physical properties in wetlands of the Sanjiang plain, China. We conducted a survey of ant mounds and measured the density, height, and diameter and material composition of different ant mounds. The ecological characteristics of wetlands that ant mounds wide occurrence were also investigated, including soil type, hydrology characters and plant composition. Differences in soil particle composition, bulk density and soil moisture between ant mound and natural meadow were measured to assess the influences of ant mounds on soil physical properties. We also studied the effects of ant mounds on the microtopography of meadows. Ant mounds were found mainly in the transition zone between terrestrial and aquatic habitats, with wetland type, including Calamagrostis augustifolia wet meadow, C.augustifolia marsh meadow, shrubs marsh meadow and Carexmeyeriana–Carexappendiculata wetland, being a significant factor. Most of the mounds detected were inhabited by Lasius flavus Fabricius, Lasius niger Linnaeus and Formica sanguinea Latreille, which occupied 52.9%, 26.5% and 20.6% of the mounds surveyed, respectively. The density, height, diameter and mound composition were significantly different among the mounds of F. sanguinea Latreille, L. flavus Fabricius and L. niger Linnaeus. The average density and diameter of L. flavus mounds was significantly higher than those of other ant mounds. The average height of F. sanguinea mounds was highest among the mounds detected. Mound building activities changed soil particle size distribution, with the silt and clay content of mounds higher than for non-mound soil. Compared with adjacent, non-mound soil, the bulk density (0–30 cm) and water content (0–25 cm) of mound soil were significantly lower, but there were no significant differences between the mound soil of F. sanguinea Latreille and L. flavus Fabricius. The spatial distribution of ant mounds with different height and diameter also changed the micro-geomorphology of the soil surface, increasing the degree of fluctuation of the microtopography. The ant distribution characteristics and their ecological roles respond to a wide range of environmental alterations. The biogenic structures of ant and the specific environment associated with them have been defined as the “functional domain”, a sphere of influence that may significantly affect soil processes at certain spatial and temporal scales. Our results suggest that the distribution and structure of ant mounds can indicate wetland environmental changes, with mounds influencing ecosystem functions and enhancing wetland degradation.     

Spatial distribution of ant mounds and effects on soil physical properties in wetlands of the Sanjiang plain, China

Ants constitute a dominant element of soil mesofauna due to their biomass, abundance, richness of species and distribution within terrestrial ecosystems. They are important regulators of soil aggregate structure as they translocate large amounts of soil from the bottom to the soil surface. In doing so, they form biogenic structures (BS) made up of aggregates of different sizes and characteristics, i.e. ant mounds. These BS have varying characteristics according to the ant species and the soil where they carry their activities. Ants are considered soil engineers because of their effects on soil properties, availability of resource and flow of energy and nutrients in soil. Thus, it is important to gain information on their distribution and abundance. Relatively little is known about the spatial distribution of mounds and their role in the soil physical properties in wetlands of the Sanjiang plain, China. We conducted a survey of ant mounds and measured the density, height, and diameter and material composition of different ant mounds. The ecological characteristics of wetlands that ant mounds wide occurrence were also investigated, including soil type, hydrology characters and plant composition. Differences in soil particle composition, bulk density and soil moisture between ant mound and natural meadow were measured to assess the influences of ant mounds on soil physical properties. We also studied the effects of ant mounds on the microtopography of meadows. Ant mounds were found mainly in the transition zone between terrestrial and aquatic habitats, with wetland type, including Calamagrostis augustifolia wet meadow, C.augustifolia marsh meadow, shrubs marsh meadow and Carexmeyeriana–Carexappendiculata wetland, being a significant factor. Most of the mounds detected were inhabited by Lasius flavus Fabricius, Lasius niger Linnaeus and Formica sanguinea Latreille, which occupied 52.9%, 26.5% and 20.6% of the mounds surveyed, respectively. The density, height, diameter and mound composition were significantly different among the mounds of F. sanguinea Latreille, L. flavus Fabricius and L. niger Linnaeus. The average density and diameter of L. flavus mounds was significantly higher than those of other ant mounds. The average height of F. sanguinea mounds was highest among the mounds detected. Mound building activities changed soil particle size distribution, with the silt and clay content of mounds higher than for non-mound soil. Compared with adjacent, non-mound soil, the bulk density (0–30 cm) and water content (0–25 cm) of mound soil were significantly lower, but there were no significant differences between the mound soil of F. sanguinea Latreille and L. flavus Fabricius. The spatial distribution of ant mounds with different height and diameter also changed the micro-geomorphology of the soil surface, increasing the degree of fluctuation of the microtopography. The ant distribution characteristics and their ecological roles respond to a wide range of environmental alterations. The biogenic structures of ant and the specific environment associated with them have been defined as the “functional domain”, a sphere of influence that may significantly affect soil processes at certain spatial and temporal scales. Our results suggest that the distribution and structure of ant mounds can indicate wetland environmental changes, with mounds influencing ecosystem functions and enhancing wetland degradation.