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University of Groningen Spatial and Temporal Hotspots of Termite-Driven University of Groningen Spatial and temporal hotspots of termite-driven decomposition in the Serengeti Freymann, Bernd P.; de Visser, Sara N.; Olff, Han; Spence, John R. Published in: Ecography DOI: 10.1111/j.1600-0587.2009.05960.x IMPORTANT NOTE: You are advised to consult the publisher's version (publisher's PDF) if you wish to cite from it. Please check the document version below. Document Version Publisher's PDF, also known as Version of record Publication date: 2010 Link to publication in University of Groningen/UMCG research database Citation for published version (APA): Freymann, B. P., de Visser, S. N., Olff, H., & Spence, J. R. (Ed.) (2010). Spatial and temporal hotspots of termite-driven decomposition in the Serengeti. Ecography, 33(3), 443-450. DOI: 10.1111/j.1600- 0587.2009.05960.x Copyright Other than for strictly personal use, it is not permitted to download or to forward/distribute the text or part of it without the consent of the author(s) and/or copyright holder(s), unless the work is under an open content license (like Creative Commons). Take-down policy If you believe that this document breaches copyright please contact us providing details, and we will remove access to the work immediately and investigate your claim. Downloaded from the University of Groningen/UMCG research database (Pure): http://www.rug.nl/research/portal. For technical reasons the number of authors shown on this cover page is limited to 10 maximum. Download date: 10-02-2018 Ecography 33: 443Á450, 2010 doi: 10.1111/j.1600-0587.2009.05960.x # 2010 The Authors. Journal compilation # 2010 Ecography Subject Editor: John R. Spence. Accepted 17 July 2009 Spatial and temporal hotspots of termite-driven decomposition in the Serengeti Bernd P. Freymann, Sara N. de Visser and Han Olff B. P. Freymann ([email protected]), S. N. de Visser and H. Olff, Centre for Ecological and Evolutionary Studies, Community and Conservation Ecology Group, Univ. of Groningen, PO Box 14, NL-9750 AA Haren, The Netherlands. Ecosystem engineers are organisms that directly or indirectly control the availability of resources to other organisms by causing physical state changes in biotic or abiotic materials. Termites (Insecta, Isoptera) are among the most important ecosystem engineers in tropical ecosystems. We used a field experiment in the tall grasslands of Serengeti National Park, Tanzania, to investigate 1) the consumption by termites of grass litter and dung baits along the landscape gradient of catena position, and 2) seasonal variation in litter and dung removal. Our maps of termitaria and patterns of bait removal revealed clear spatial and temporal hotspots of termite activity. In the dry season termites removed more baits at the top- catena positions than at the bottom positions, but there was no effect of catena position in the wet season. Spatial hotspots of termite activity overlapped with those of both mammalian herbivores and predators. Within the framework of ecosystem engineering, this study suggests that intraspecific aspects of spatial heterogeneity and temporal variability deserve much greater consideration. Every living organism interacts with its environment, 2006, Jime´nez and Decae¨ns 2006). Some termites act as including other individuals of its own species and other herbivores as well as decomposers, feeding on a wide range biotic elements through competition, mutualism, parasitism of living, dead or decaying plant materials and soil and predation (Krebs 2001). Recently, the concept of (Adamson 1943, Lee and Wood 1971, Wood 1976, interaction types was extended to include the conceptual 1978, Bignell and Eggleton 2000, Traniello and Leuthold framework of ecosystem engineering (Jones et al. 1994, 2000). Also, termites may play an important role in 1997, Wright and Jones 2006). Ecosystem engineers are recycling herbivore dung in tropical ecosystems (Freymann organisms that directly or indirectly control the availability et al. 2008). Thus, termites play a crucial role in the of resources to other organisms by causing physical state decomposition processes of tropical ecosystems, providing changes in biotic or abiotic materials (Jones et al. 1997). ecosystem services via nutrient cycling (Lavelle et al. 2006). Termites (Insecta, Isoptera) and earthworms are among Termite mounds are common but heterogeneously the most important ecosystem engineers in terrestrial distributed landscape elements in several regions of Africa ecosystems (Lavelle et al. 1997). As eusocial insects, termites (Pomeroy 1977, Kaib et al. 1997, Meyer et al. 1999), but build nests of great architectural diversity ranging from the implications of this spatial and temporal heterogeneity inconspicuous below-ground nests to the massive above- for decomposition activity has only rarely been studied ground mounds of Macrotermes spp. (Noirot and Darling- (Schuurman 2005). At the landscape scale, the concept of ton 2000). Adamson (1943) suggested that termites are soil catenae is particularly important in semi-arid ecosys- crucial for maintaining the fertility of tropical soils and the tems such as the African plains (Jarman and Sinclair 1979). productivity of the tropical ecosystems they inhabit because Milne (1935) introduced the pedological term catena they promote aeration and mixing of the soil, drainage and (Latin: chain) in senso stricto to describe a sequence of penetration of plant roots. In addition, termites accelerate repetitive soil patterns along the slope of a hill. Soils at the formation of humus and nutrient cycling by consuming a top position of a catena are characterised by a higher sand variety of cellulose containing material. content and a lower clay and water content in comparison Many studies have demonstrated the beneficial ecosys- to the soils at the bottom position; this pattern is common tem impacts of termites (Wood and Sands 1978, Anderson in the central region of Serengeti National Park (SNP) and Wood 1984, Abbadie et al. 1992, Holt and Lepage (Anderson et al. 2006). Despite the ecological importance 2000), some of which deal explicitly with their ecosystem of these landscape features, we know of no studies of termite engineering activities (Black and Okwakol 1997, Lavelle activity in relation to variation in soil texture and depth et al. 1997, Dangerfield et al. 1998, Jouquet et al. 2005, along catenae. 443 In this study we explored the spatial and temporal Field experiment heterogeneity of termite activity in the long-grass plains of Serengeti National Park, Tanzania. We hypothesised that Experimental sites were located randomly within the long spatial and temporal hotspots of termite activity and grass region of SNP, stratified according to top or bottom therefore decomposition would follow from the reported position of a catena. In total, eight experimental plots were spatio-temporal heterogeneity of termite mounds (Sinclair established on each ridge position. The minimum distance 1979) in this savanna ecosystem. We used a field experi- between plots located on the same hill was 500 m. The ment to investigate whether termite consumption of grass distance between top and bottom positions was about 1 km. litter and dung differed between the top and bottom We randomly chose a point in the field as the centre of a positions of catena ridges. We also compared termite 33 m plot. We marked the plots and subdivided them activity along the catena between the dry and wet seasons. into nine 11 m subplots in which we placed the baits. In addition, we quantified the spatial distribution of Each 33 m experimental plot formed the centre of a termitaria in this ecosystem to establish relationships wider 100100 m plot used for the later mapping of the between nest location and ecologically significant activities present termite mounds. related to decomposition. In a pilot experiment we identified a mesh-size of 1.5 mm to be suitable to allow the dominant termite taxa of SNP to access baits placed in metal mesh-bags (10 Material and methods 10 cm), whereas the majority of the remaining arthropod fauna, especially dung beetles, could be excluded. Study area Baits were collected and processed as follows: from the vicinity of each experimental plot we randomly harvested This study was conducted in the long grass region in the nine samples of the characteristic grass species of the southern part of Serengeti National Park (SNP), Tanzania particular site and air-dried them for three weeks. This (2828?S, 34853??E; 1570 m a.s.l.; Fig. 1A). The prevailing treatment was necessary since the dominant termite taxa in habitat type of the study area is a semi-arid grassland, this area (Odontotermes sp., Macrotermes sp.) consume dry dominated by long-grass species such as Themeda triandra grass litter. Of each grass sample a portion of 15Á20 g was and Pennisetum mezianum. For further details see Sinclair randomly chosen, filled in a mesh-bag and labelled. The (1979). The study site is characterised by an undulating exact filling weight of each of the 144 litter baits was topography with small ridges, catenae, showing an elevational determined using a digital scale accurate to 0.001g. We difference of 15Á25 m between hill tops and bases. In contrast collected 144 fresh wildebeest (Connochaetes taurinus) to these catenae located throughout the study area, the five droppings. Of each dropping a random sample of about hills of the so-called five-hills-track, also situated in the long 50 g was put into a mesh-bag and processed in the same grass region of SNP, show considerably larger elevational way as the litter baits. Placement of the baits took place differences (75Á100 m) between top and bottom. The dry within one day (day 0 of the experiment). In each 11m season component of the experiment was conducted in late subplot we placed one litter and one dung mesh-bag in July 2005, and the wet season component in early May 2006. close contact with the soil. Prior to the re-collection of the (A) (B) KENYA SNP TANZANIA LGP 05025 Kilometers Figure 1.
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