Sampling subterranean termite species diversity and activity in tropical savannas: an assessment of different bait choices

Abstract.  1. Cellulose baits are commonly used for semi-quantitative sampling of subterranean wood-feeding termites, with a single food choice sampling programme implemented most often. In most situations, however, the composition and feeding preferences of the subterranean termite assemblage remain unknown.
2. The diversity, frequency, foraging activity, and intensity of attack of termites were assessed regularly at 144 baits representing 12 different bait choices over 8.5 months, in two northern Australian tropical savanna sites that differed in vegetation structure (closed vs open). Baits differed in type (paper rolls, cardboard, wooden stakes), position (surface, buried), and moisture status at installation (wet, dry).
3. Sixteen species were recorded, including 11 wood-feeders. Average species diversity, foraging activity, and bait consumption were greater at buried baits than at surface baits. Wooden baits were most attractive early in the experiment, and paper baits more attractive later. Mean species diversity was greatest at wooden stakes in the closed site. Species frequency of occurrence varied across bait choices.
4. A composite bait sampling protocol of stakes and paper rolls installed above and below ground gave an accurate assessment of the activity, diversity, and structure of the termite guild sampled across all baits over 8.5 months.
5. The choice of bait, its presentation, and time of examination are critical to the success of a termite baiting programme. If the aim is to characterise the structure and foraging activity of the subterranean termite assemblages that are attracted to baits, composite baiting protocols should be implemented.     

Stability structures tropical woody plant diversity more than seasonality: Insights into the ecology of high legume-succulent-plant biodiversity

Phylogenies of legume taxa are ecologically structured along a tropical seasonality gradient, which suggests phylogenetic niche conservatism. This seasonality gradient spans Neotropical wet forests, savannas, and highly seasonal drought-prone woody vegetation known as the succulent biome. Ecological phylogenetic structure was investigated using a community phylogenetic approach. We further analyzed bioclimatic and other independent variables that potentially explained phylogenetic beta diversity among 466 floristic sites that spanned the savanna and succulent biomes in eastern South America. Explanatory variables were selected using variance inflation factors, information criteria, and the ability to explain both species and phylogenetic beta diversity. A model involving annual precipitation suggests that a threshold of < 1200 mm explains community phylogenetic structure along the savanna–succulent biome transition. Variables involving temperatures or measures of seasonality were notably lacking from top-ranked models. The abundance and diversity of legumes across the tropical seasonality gradient suggest that a high nitrogen metabolism confers an advantage in one of two ways, both of which are related to rapid growth rates. Legumes adapted to the succulent biome may be responding to regular post-dry-season leaf-flush opportunities. Legumes adapted to the savanna biome may be responding to intermittent post-disturbance growing opportunities. A seasonal predominance of leaf flushing by woody plants implicates the role of ecological stability in the succulent biome because of the need to recover the cost of regenerating short-lived leaves. Ecological stability may be the fundamental cause of ecological phylogenetic structure across the tropical seasonality gradient and required for maintaining high levels of both leaf-flushing legume and succulent plant biodiversity.     

The termite (Isoptera) fauna of a monsoonal rainforest near Darwin, northern Australia

Abstract  Termite species richness, frequency of occurrence and functional diversity at Holmes Jungle Nature Reserve, a monsoon rainforest patch in northern Australia, was investigated at the end of the wet season in 2003. A sampling protocol that employed direct search, soil pits and baiting techniques was used to sample litter, wood, mound, soil and arboreal nest microhabitats for termites. Five species from five genera and three families (Mastotermitidae, Rhinotermitidae and Termitidae) were recorded. This included the first record of Ephelotermes taylori (Hill) from monsoon forest in Australia. The family Termitidae was dominant and represented 70% of termite occurrences. Termites were most frequently encountered in carton runways on tree trunks, followed by lying dead wood and baits. Four nesting habits were represented: arboreal, epigeal, hypogeal and within wood. The arboreal nest-builder Nasutitermes graveolus (Hill) accounted for 61% of termite encounters. Epigeal mound-building species were rare. Wood-feeders were the only trophic group represented. Relatively high activity of Mastotermes darwiniensis Froggatt was recorded at baits within 2.5 months. The low species richness recorded at Holmes Jungle is consistent with the low diversity reported for Australian rainforests generally, but remains relatively depauperate compared with other monsoon forest and savanna habitats of the Northern Territory.     

Stocks and dynamics of carbon in trees across a rainfall gradient in a tropical savanna

In this study, systematic variation in tree morphology across a rainfall gradient in Australia's tropical savanna biome and its implications for carbon stocks and dynamics were quantified. The aim was to support efforts to manage fire regimes to increase vegetative carbon stocks as a greenhouse gas mitigation strategy. The height of trees for a given trunk diameter declines with decreasing rainfall from 2000 to 300 mm and increasing dry season length across the Australian savanna biome. It is likely that increasing dry season length is the main driver of this decline rather declining rainfall per se. By taking account of the response of total basal area to rainfall and soil type, stand structure, and tree height and diameter relationships, the carbon stocks in live trees were estimated to decline from about 34 t ha^?1 in the wetter savannas to 6 t ha^?1 in the drier savannas. These values are broadly consistent with field‐based estimates. Because of the declining ratio of height to trunk diameter, trees of a given diameter in drier regions will be more likely to be killed by fires of a given intensity than trees in wetter regions. Thus single fires of given intensity are likely to have a greater proportionate impact on live tree carbon stock in drier savannas, but a much greater absolute impact in wetter savannas due to the greater total carbon stock. Projected decreases in early wet season rainfall under climate change scenarios, despite projections of little change in total precipitation in northern Australia, may lead to decreased carbon stock in live trees through two mechanisms: a reduction in total basal area and decreases in tree height for given trunk diameters.     

When is a ‘forest’ a savanna,and why does it matter?

Savannas are defined based on vegetation structure, the central concept being a discontinuous tree cover in a continuous grass understorey. However, at the high‐rainfall end of the tropical savanna biome, where heavily wooded mesic savannas begin to structurally resemble forests, or where tropical forests are degraded such that they open out to structurally resemble savannas, vegetation structure alone may be inadequate to distinguish mesic savanna from forest. Additional knowledge of the functional differences between these ecosystems which contrast sharply in their evolutionary and ecological history is required. Specifically, we suggest that tropical mesic savannas are predominantly mixed tree–C₄ grass systems defined by fire tolerance and shade intolerance of their species, while forests, from which C₄ grasses are largely absent, have species that are mostly fire intolerant and shade tolerant. Using this framework, we identify a suite of morphological, physiological and life‐history traits that are likely to differ between tropical mesic savanna and forest species. We suggest that these traits can be used to distinguish between these ecosystems and thereby aid their appropriate management and conservation. We also suggest that many areas in South Asia classified as tropical dry forests, but characterized by fire‐resistant tree species in a C₄ grass‐dominated understorey, would be better classified as mesic savannas requiring fire and light to maintain the unique mix of species that characterize them.     

Drought-induced tree death in savanna

Increasing densities of woody plants in savannas has been attributed to both elevated atmospheric CO₂ and reduced burning with grazing management, such that the biome could represent a substantial carbon sink. However, we show that extreme droughts (less than two-thirds expected rainfall over 3 years) occur in the drier half of the savanna biome and can cause substantial tree death. An Australian case study reveals that a net increase in tree cover over five decades of above-average rainfall was offset by sudden tree death during drought. The relationship between woody cover change and rainfall is moderated by competition with growth being facilitated by low woody cover and drought-induced death more likely as the woody component of savanna increases. The results are not supportive of a sustained increase in the woody component of xeric savannas resulting from CO₂ fertilization or land management. Extensive tree death in savanna regions will become a stark consequence of climate change if predictions of increasing severity and frequency of drought are realized.     

Hermetically sealed baits for subterranean termites (Isoptera: Rhinotermitidae)

Cellulose baits containing 0.5% hexaflumuron were hermetically sealed in a closed cell polyethylene sheet envelope and placed in soil to test their durability and efficacy against field colonies of the Formosan subterranean termite, Coptotermes formosanus Shiraki, or the eastern subterranean termite, Reticulitermes flavipes (Kollar). The closed cell polyethylene sheet was readily tunneled through by termites, yet it was impervious to water and protected the cellulose baits and hexaflumuron from the environment. Only in a few incidents did the polyethylene envelope become infiltrated by plant roots, resulting in water intrusion and apparent degradation of cellulose baits. After consuming one to three sealed baits, three colonies each of both termite species were eliminated. The sealed baits may be placed in soil for months or years without the need of monitoring, and they are readily penetrated and fed upon by termites when they are present. Application of baits hermetically sealed in a protective sheet may save labor costs by bypassing the monitoring phase, circumvent the station avoidance by some termite species, and enable the use of baiting technologies in large areas such as agricultural fields in which the manual monitoring is impractical.     

Are the eucalypt and non-eucalypt components of Australian tropical savannas independent?

Eucalypts (Eucalyptus and Corymbia spp.) dominate (>60%) the tree biomass of Australia's tropical savannas but account for only a fraction (28%) of the tree diversity. Because of their considerable biomass and adaptation to environmental stressors, such as fire, the eucalypts may drive tree dynamics in these savannas, possibly to the exclusion of non-eucalypts. We evaluated whether the eucalypt and non-eucalypt components in tropical savannas are dependent so that changes in one component are matched by opposite trends in the other. Using tree inventory data from 127 savanna sites across the rainfall and fire frequency gradients, we found that eucalypt and non-eucalypt basal area and species richness had a negative relationship. This relationship was maintained across the rainfall gradient, with rainfall having a positive effect on the basal area and species richness of both components, but with a greater effect in non-eucalypts. Fire frequency negatively affected basal area, but not species richness, although basal area and species richness of eucalypts and non-eucalypts did not differ in their response to fire. Rainfall appears to set the upper bounds to woody biomass in these mesic savannas, while fire maintains woody biomass below carrying capacity and facilitates coexistence of the components. The magnitude of the component responses, particularly for non-eucalypts, is determined by rainfall, but their dependence is likely due to their differential response to both rainfall and fire, but not to competition for resources. Thus, while eucalypts dominate biomass overall, at high rainfall sites non-eucalypt basal area and diversity are highest, especially where fire frequency is low.     

Neotropical savanna ants show a reversed latitudinal gradient of species richness,with climatic drivers reflecting the forest origin of the fauna

Aim

To evaluate the extent to which ant species richness in Neotropical savannas varies with macrogeographic variables, and to identify the potential climatic drivers of such variation.

Location

The Cerrado savanna biome of central Brazil, in a region spanning ca. 20° of latitude and 18°of longitude.

Methods

Standardized sampling of the arboreal and ground‐dwelling faunas was performed in 29 well‐preserved savanna sites using pitfall traps. Species were classified according to their habitat affinities: open‐savanna specialists, forest‐associated species or habitat generalists. We used generalized linear models to evaluate the importance of geographic (latitude, longitude and elevation) and climatic (mean temperature and three metrics of rainfall) variables as predictors of species richness.

Results

The total number of species recorded at each site varied more than twofold (from 59 to 144), and latitude was the best geographic correlate of overall species richness. However, contrary to the expected pattern, more species were found at higher than lower latitudes. This reversed latitudinal pattern of diversity occurred for both the arboreal and ground‐dwelling faunas, and for the habitat generalists and forest specialists. The savanna specialists showed a mid‐latitudinal peak in diversity. Overall, there was a significant positive association between rainfall and species richness, but the strength of this relationship varied with ant habitat affinity.

Main conclusions

The Cerrado ant fauna shows a reverse latitudinal gradient in species diversity, and this can be explained by increasing rainfall during the warmest months of the year (and therefore in plant productivity) with increasing latitude. The sensitivity of Cerrado ant diversity to declining rainfall contrasts with the high resilience to aridity of the Australian savanna ant fauna, and this reflects the contrasting evolutionary histories of these faunas. Our findings highlight the importance of historical processes as drivers of intercontinental contrasts in macroecological patterns.     

Multi‐scale patterns and bush encroachment in an arid savanna with a shallow soil layer

Abstract. Question: Bush encroachment (i.e. an increase in density of woody plants often unpalatable to domestic livestock) is a serious problem in many savannas and threatens the livelihood of many pastoralists. Can we derive a better understanding of the factors causing bush encroachment by investigating the scale dependency of patterns and processes in savannas? Location: An arid savanna in the Khomas Hochland, Namibia. Methods: Patterns of bush, grass, and soil nutrient distribution were surveyed on several scales along a rainfall gradient, with emphasis on intraspecific interactions within the dominant woody species, Acacia reficiens. Results: Savannas can be interpreted as patch‐dynamic systems where landscapes are composed of many patches (a few ha in size) in different states of transition between grassy and woody dominance. Conclusions: In arid savannas, this patchiness is driven both by rainfall that is highly variable in space and time and by inter‐tree competition. Within the paradigm of patch‐dynamic savannas, bush encroachment is part of a cyclical succession between open savanna and woody dominance. The conversion from a patch of open savanna to a bush‐encroached area is initiated by the spatial and temporal overlap of several (localized) rainfall events sufficient for Acacia germination and establishment. With time, growth and self‐thinning will transform the bush‐encroached area into a mature Acacia stand and eventually into open savanna again. Patchiness is sustained due to the local rarity (and patchiness) of rainfall sufficient for germination of woody plants as well as by plant‐soil interactions.     

Decadal dynamics of tree cover in an Australian tropical savanna

Spatio‐temporal variation in tropical savanna tree cover remains poorly understood. We aimed to quantify the drivers of tree cover in tropical mesic savannas in Kakadu National Park by relating changes in tree cover over 40 years to: mean annual rainfall, fire activity, initial tree cover and prior changes in tree cover. Aerial photography, acquired in 1964, 1984 and 2004, was obtained for fifty sites in Kakadu that spanned a rainfall gradient from approximately 1200 to 1600 mm. The remotely sensed estimates of tree cover were validated via field survey. Linear mixed effects modelling and multi‐model inference were used to assess the strength and form of the relationships between tree cover and predictor variables. Over the 40 years, tree cover across these savannas increased on average by 4.94 ± 0.88%, but was spatio‐temporally variable. Tree cover showed a positive albeit weak trend across the rainfall gradient. The strength of this positive relationship varied over the three measurement times, and this suggests that other factors are important in controlling tree cover. Tree cover was positively related to prior tree cover, and negatively correlated with fire activity. Over 20 years tree cover was more likely to increase if (i) tree cover was initially low or (ii) had decreased in the previous 20‐year interval or (iii) there had been fewer fires. Across the examined rainfall gradient, the greater variability in fire activity and inherently higher average tree cover at the wetter latitudes resulted in greater dynamism of tree cover compared with the drier latitudes. This is consistent with savanna tree cover being determined by interactions between mean annual rainfall, tree competition and frequent fire in these tropical mesic savannas.     

Evolution of termite functional diversity: analysis and synthesis of local ecological and regional influences on local species richness

Aim To (1) describe termite functional diversity patterns across five tropical regions using local species richness sampling of standardized areas of habitat; (2) assess the relative importance of environmental factors operating at different spatial and temporal scales in influencing variation in species representation within feeding groups and functional taxonomic groups across the tropics; (3) achieve a synthesis to explain the observed patterns of convergence and divergence in termite functional diversity that draws on termite ecological and biogeographical evidence to‐date, as well as the latest evidence for the evolutionary and distributional history of tropical rain forests. Location Pantropical. Methods A pantropical termite species richness data set was obtained through sampling of eighty‐seven standardized local termite diversity transects from twenty‐nine locations across five tropical regions. Local‐scale, intermediate‐scale and large‐scale environmental data were collected for each transect. Standardized termite assemblage and environmental data were analysed at the levels of whole assemblages and feeding groups (using components of variance analysis) and at the level of functional taxonomic groups (using correspondence analysis and canonical correspondence analysis). Results Overall species richness of local assemblages showed a greater component of variation attributable to local habitat disturbance level than to region. However, an analysis accounting for species richness across termite feeding groups indicated a much larger component of variation attributable to region. Mean local assemblage body size also showed the greater overall significance of region compared with habitat type in influencing variation. Ordination of functional taxonomic group data revealed a primary gradient of variation corresponding to rank order of species richness within sites and to mean local species richness within regions. The latter was in the order: Africa > south America > south‐east Asia > Madagascar > Australia. This primary gradient of species richness decrease can be explained by a decrease in species richness of less dispersive functional taxonomic groups feeding on more humified food substrates such as soil. Hence, the transects from more depauperate sites/regions were dominated by more dispersive functional taxonomic groups feeding on less humified food substrates such as dead wood. Direct gradient analysis indicated that ‘region’ and other large‐scale factors were the most important in explaining patterns of local termite functional diversity followed by intermediate‐scale geographical and site variables and, finally, local‐scale ecological variables. Synthesis and main conclusions Within regions, centres of termite functional diversity lie in lowland equatorial closed canopy tropical forests. Soil feeding termite evolution further down food substrate humification gradients is therefore more likely to have depended on the long‐term presence of this habitat. Known ecological and energetic constraints upon contemporary soil feeders lend support for this hypothesis. We propose further that the anomalous distribution of termite soil feeder species richness is partly explained by their generally very poor dispersal abilities across oceans. Evolution, radiation and dispersal of soil feeder diversity appears to have been largely restricted to what are now the African and south American regions. The inter‐regional differences in contemporary local patterns of termite species richness revealed by the global data set point to the possibility of large differences in consequent ecosystem processes in apparently similar habitats on different continents.     

Contrasting demographics of tropical savanna and temperate forest eucalypts provide insight into how savannas and forests function. A case study using Corymbia clarksoniana from north‐eastern Australia

Eucalypts (Eucalyptus spp. and Corymbia spp.) dominate many communities across Australia, including frequently burnt tropical savannas and temperate forests, which receive less frequent but more intense fires. Understanding the demographic characteristics that allow related trees to persist in tropical savannas and temperate forest ecosystems can provide insight into how savannas and forests function, including grass–tree coexistence. This study reviews differences in critical stages in the life cycle of savanna and temperate forest eucalypts, especially in relation to fire. It adds to the limited data on tropical eucalypts, by evaluating the effect of fire regimes on the population biology of Corymbia clarksoniana, a tree that dominates some tropical savannas of north‐eastern Australia. Corymbia clarksoniana displays similar demographic characteristics to other tropical savanna species, except that seedling emergence is enhanced when seed falls onto recently burnt ground during a high rainfall period. In contrast to many temperate forest eucalypts, tropical savanna eucalypts lack canopy‐stored seed banks; time annual seed fall to coincide with the onset of predictable wet season rain; have very rare seedling emergence events, including a lack of mass germination after each fire; possess an abundant sapling bank; and every tropical eucalypt species has the ability to maintain canopy structure by epicormically resprouting after all but the most intense fires. The combination of poor seedling recruitment strategies, coupled with characteristics allowing long‐term persistence of established plants, indicate tropical savanna eucalypts function through the persistence niche rather than the regeneration niche. The high rainfall‐promoted seedling emergence of C. clarksoniana and the reduction of seedling survival and sapling growth by fire, support the predictions that grass–tree coexistence in savannas is governed by rainfall limiting tree seedling recruitment and regular fires limiting the growth of juvenile trees to the canopy.     

Elimination of subterranean termite (Isoptera: Rhinotermitidae) colonies using a refined cellulose bait matrix containing noviflumuron when monitored and replenished quarterly

Using a quarterly (3-mo) monitoring and bait-replenishment interval, 122 subterranean termite colonies throughout the United States were baited with a refined cellulose bait matrix containing 0.5% noviflumuron. All colonies were eliminated in less than 1 yr after initiation of baiting as determined by long-term monitoring and genetic markers. Sixty-three percent of the colonies were eliminated during the first quarter after the initiation of baiting and 77% of colonies were eliminated after consuming two bait tubes or less. This suggests that a single baiting cycle and bait installed in response to a single active monitoring device were sufficient to eliminate the majority of colonies. Although termites temporarily abandoned stations after depleting bait, workers resumed feeding when baits were replenished. Colonies that consumed large amounts of bait before elimination foraged into multiple stations, thus allowing adequate amounts of bait to sustain feeding. The time to eliminate termite colonies with bait replenished quarterly was similar to that previously reported for laminated cellulose bait replenished monthly. Our data support the conclusion that extending the bait replenishment interval from monthly to quarterly for bait tubes with refined cellulose containing 0.5% noviflumuron did not adversely impact colony elimination.     

Productivity and carbon fluxes of tropical savannas

Aim (1) To estimate the local and global magnitude of carbon fluxes between savanna and the atmosphere, and to suggest the significance of savannas in the global carbon cycle. (2) To suggest the extent to which protection of savannas could contribute to a global carbon sequestration initiative. Location Tropical savanna ecosystems in Africa, Australia, India and South America. Methods A literature search was carried out using the ISI Web of Knowledge, and a compilation of extra data was obtained from other literature, including national reports accessed through the personal collections of the authors. Savanna is here defined as any tropical ecosystem containing grasses, including woodland and grassland types. From these data it was possible to estimate the fluxes of carbon dioxide between the entire savanna biome on a global scale. Results Tropical savannas can be remarkably productive, with a net primary productivity that ranges from 1 to 12 t C ha⁻¹ year⁻¹. The lower values are found in the arid and semi‐arid savannas occurring in extensive regions of Africa, Australia and South America. The global average of the cases reviewed here was 7.2 t C ha⁻¹ year⁻¹. The carbon sequestration rate (net ecosystem productivity) may average 0.14 t C ha⁻¹ year⁻¹ or 0.39 Gt C year⁻¹. If savannas were to be protected from fire and grazing, most of them would accumulate substantial carbon and the sink would be larger. Savannas are under anthropogenic pressure, but this has been much less publicized than deforestation in the rain forest biome. The rate of loss is not well established, but may exceed 1% per year, approximately twice as fast as that of rain forests. Globally, this is likely to constitute a flux to the atmosphere that is at least as large as that arising from deforestation of the rain forest. Main conclusions The current rate of loss impacts appreciably on the global carbon balance. There is considerable scope for using many of the savannas as sites for carbon sequestration, by simply protecting them from burning and grazing, and permitting them to increase in stature and carbon content over periods of several decades.     

Patterns of Ant (Hymenoptera: Formicidae) Richness and Relative Abundance along an Aridity Gradient in Western Venezuela

In xeric ecosystems, ant diversity response to aridity varies with rainfall magnitude and gradient extension. At a local scale and with low precipitation regimes, increased aridity leads to a reduction of species richness and an increased relative abundance for some ant species. In order to test this pattern in tropical environments, ant richness and relative abundance variation were evaluated along 35 km of an aridity gradient in the Araya Peninsula, state of Sucre, Venezuela. Three sampling stations comprising five transects each were set up. Pitfall traps and direct collecting from vegetation were assessed per transect. Overall, 52 species, 23 genera, and 7 subfamilies of ants were recorded in the peninsula. The total number of species and genera recorded by both sampling stations and transects decreased linearly with increasing aridity. Total relative abundance was highest in the most arid portion of the peninsula, with Crematogaster rochai (Forel) and Camponotus conspicuus zonatus (Emery) (Hymenoptera: Formicidae) being the numerically dominant species. Spatial and multivariate analyses revealed significant changes in ant composition every 11 km of distance, and showed a decrease of ant diversity with the increase of harsh conditions in the gradient. Here, we discuss how local geographic and topographic features of Araya originate the aridity gradient and so affect the microhabitat conditions for the ant fauna.     

A patch-dynamics approach to savanna dynamics and woody plant encroachment – Insights from an arid savanna

The coexistence of woody and grassy plants in savannas has often been attributed to a rooting-niche separation (two-layer hypothesis). Water was assumed to be the limiting resource for both growth forms and grasses were assumed to extract water from the upper soil layer and trees and bushes from the lower layers. Woody plant encroachment (i.e. an increase in density of woody plants often unpalatable to domestic livestock) is a serious problem in many savannas and is believed to be the result of overgrazing in ‘two-layer systems’. Recent research has questioned the universality of both the two-layer hypothesis and the hypothesis that overgrazing is the cause of woody plant encroachment.

We present an alternative hypothesis explaining both tree–grass coexistence and woody plant encroachment in arid savannas. We propose that woody plant encroachment is part of a cyclical succession between open savanna and woody dominance and is driven by two factors: rainfall that is highly variable in space and time, and inter-tree competition. In this case, savanna landscapes are composed of many patches (a few hectares in size) in different states of transition between grassy and woody dominance, i.e. we hypothesize that arid savannas are patch-dynamic systems. We summarize patterns of tree distribution observed in an arid savanna in Namibia and show that these patterns are in agreement with the patch-dynamic savanna hypothesis. We discuss the applicability of this hypothesis to fire-dominated savannas, in which rainfall variability is low and fire drives spatial heterogeneity.

We conclude that field studies are more likely to contribute to a general understanding of tree–grass coexistence and woody plant encroachment if they consider both primary (rain and nutrients) and secondary (fire and grazing) determinants of patch properties across different savannas.     

Rainfall Influences Ant Sampling in Dry Forests

The standardized 'Ants of the Leaf Litter' protocol aims to facilitate the use of ground-foraging and litter-dwelling ants in biodiversity assessment and monitoring programs. It was initially developed to characterize assemblages from tropical rain forests and is based on two main techniques: Winkler extractions and pitfall traps. Here, we tested to what extent this protocol was adapted to tropical dry forests and affected by the rainfall regime. Our 10 study sites were located along an aridity gradient (average annual rainfall: 350–1300mm) in the Gran Chaco. The number of species collected per sampling effort increased with aridity for pitfalls but followed an opposite trend for Winkler samples. This trend could be explained by the low daytime foraging activity in the leaf litter during drought periods. In arid and semiarid regions the good performance of pitfalls was probably related to their 24-h operation and to the attractiveness of the water they contained. Our results stress that the Winkler method used in the Ants of the Leaf Litter protocol may not be cost-effective during periods of drought and may lead to severe underestimations of litter ant diversity in tropical dry forests.     

Root foraging strategies and soil patchiness in a humid savanna

In Lamto (Côte d'Ivoire), the savanna is a patchy environment as far as soil is concerned: tree clumps and termite mounds lead to higher nutrient contents than in the surrounding savanna. Mature Borassus aethiopum (Mart.) specimens are tall palm trees dominating the community, with aerial parts located out of these nutrient-rich patches.Palm root densities were compared under tree clumps and in the surrounding savanna, and were also sampled along transects between palm trees and nutrient-rich patches (two clumps and one mound). Palm root densities were far higher (up to 10 times) in the nitrogen-rich soil of both clumps and termite mounds than in the surrounding savanna. Evidence is given that palm trees are able to extend their root system as far as 20 m towards these nutrient-rich patches where they proliferate. These results point out a particular root foraging strategy, which is one of the first known for a woody perennial. They also provide new insights for understanding nitrogen cycling and savannas high rate of primary production.     

Decomposition of tissue baits and termite density along a gradient of human land‐use intensification in Western Kenya

Termites are important decomposers and ‘ecosystem engineers’ in tropical ecosystems. Furthermore, termite assemblages are sensitive to human land‐use intensification and often termite density and the importance of soil‐feeding termites decrease with land‐use intensification. These changes in termite assemblages may also lead to a decrease in termite‐mediated ecosystem processes (e.g. soil formation, cellulose decomposition). We compared density and functional composition of termites with cellulose removal from undisturbed primary forests to farmlands (Kakamega Forest, Western Kenya). In contrast to the expectation, we found no response of termite abundance along the gradient of land‐use intensification. However, as expected, the relative abundance of soil‐feeders decreased from primary forests to farmlands. In contrast, frequency of attack on tissue paper baits and removal of tissue showed a clear hump‐shaped relationship to land‐use intensification with high values in secondary forests. These nonconcordant patterns of density and functional composition of termite assemblages with cellulose removal by termites suggest that it may be misleading to infer changes in a process by the characteristics of the assemblage of organisms that mediate that process.