Phantoms of the forest: legacy risk effects of a regionally extinct large carnivore

The increased abundance of large carnivores in Europe is a conservation success, but the impact on the behavior and population dynamics of prey species is generally unknown. In Europe, the recolonization of large carnivores often occurs in areas where humans have greatly modified the landscape through forestry or agriculture. Currently, we poorly understand the effects of recolonizing large carnivores on extant prey species in anthropogenic landscapes. Here, we investigated if ungulate prey species showed innate responses to the scent of a regionally exterminated but native large carnivore, and whether the responses were affected by human-induced habitat openness. We experimentally introduced brown bear Ursus arctos scent to artificial feeding sites and used camera traps to document the responses of three sympatric ungulate species. In addition to controls without scent, reindeer scent Rangifer tarandus was used as a noncarnivore, novel control scent. Fallow deer Dama dama strongly avoided areas with bear scent. In the presence of bear scent, all ungulate species generally used open sites more than closed sites, whereas the opposite was observed at sites with reindeer scent or without scent. The opening of forest habitat by human practices, such as forestry and agriculture, creates a larger gradient in habitat openness than available in relatively unaffected closed forest systems, which may create opportunities for prey to alter their habitat selection and reduce predation risk in human-modified systems that do not exist in more natural forest systems. Increased knowledge about antipredator responses in areas subjected to anthropogenic change is important because these responses may affect prey population dynamics, lower trophic levels, and attitudes toward large carnivores. These aspects may be of particular relevance in the light of the increasing wildlife populations across much of Europe

Recreation and hunting differentially affect deer behaviour and sapling performance

Humans are increasingly acknowledged as apex predators that shape landscapes of fear to which herbivores adapt their behaviour. Here, we investigate how humans modify deer space-use and their effects on vegetation at two spatial scales; zones with different types of human use (largescale risk factor) and, nested within that, trails (fine-scale risk factor). In zones with three contrasting types of human activities: 1) no recreation, no hunting, 2) with recreation, no hunting and 3) with recreation and hunting, we linked deer space-use (dropping counts) to browsing intensity, relative growth and survival of planted saplings. Plots were located at two distances to trails (20 versus 100 m) to test how trails affect deer space-use and sapling performance. Additionally, plots were distributed over forest and heathland as risk effects are habitat-dependent. Deer space-use was highest in the zone without recreation or hunting, resulting in higher browsing levels and lower sapling growth and survival, but only in heathland. In contrast, deer space-use and sapling performance did not differ between zones with recreation only and zones with recreation and hunting. Deer dropping counts were lower near trails used for recreation, but this was not associated with browsing impact or sapling performance. Our results show that recreational use modifies deer space-use which is associated with browsing impact on woody vegetation, while seasonal hunting activities in zones with recreation did not have additive year-round effects. Yet, effects were only observed at the larger scale of recreation zones and not near trails. Furthermore, deer space-use was only associated with sapling performance in open heathland, where high visibility presumably increases avoidance behaviour because it increases detectability and decreases escape possibilities. This suggests that recreation creates behaviourally mediated cascading effects that influence vegetation development, yet these effects are context-dependent. We advocate incorporating human-induced fear effects in conservation, management and research

Hunting as land use: Understanding the spatial associations among hunting, agriculture, and forestry

Hunting is a widespread but often overlooked land-use activity, providing major benefits to society. Hunting takes place in most landscapes, yet it remains unclear which types of landscapes foster or dampen hunting-related services, and how hunting relates to other land uses. A better understanding of these relationships is key for sustainable land-use planning that integrates wildlife management. This is particularly urgent for Europe, where wildlife populations are increasing. Focusing on Sweden, we explored the spatial associations among hunting, agriculture, and forestry to identify archetypical combinations of these land uses. Specifically, we combined indicators on the extent and intensity of agriculture and forestry, with data on hunting bags for 63 game species using self-organizing maps, a non-parametric clustering approach. We identified 15 typical bundles of co-occurring land uses at the municipality level across Sweden. The harvest of forest grouse, bears, and moose co-occurred with forestry in northern Sweden, whereas the harvest of small game, different deer species, and wild boar co-occurred with agriculture across southern Sweden, reflecting species’ biology, environmental factors, and management. Our findings also highlight the strength of associations among hunting and other land uses. Importantly, we identified large areas in central Sweden where harvest of game was below average, possibly indicating that intensity of hunting is out of balance with that of agriculture or forestry, potentially fostering conflict between wildlife and land use. Collectively, our results suggest that (1) hunting should be considered a major land use that, in Sweden, is more widespread than agriculture and forestry; (2) land-use planning must therefore integrate wildlife management; and (3) such an integration should occur in a regionalized manner that considers social-ecological context. Our approach identifies a first spatial template within which such context-specific land-use planning, aiming at aligning wildlife and diverse land uses, can take place

The relative role of intrinsic and extrinsic drivers in regulating population change and survival of African wild dogs (Lycaon pictus)

DATA AVAILABILITY : The datasets generated during and/or analysed during the current study are available in the figshare repository https://doi.org/10.6084/m9.figshare.14368043.v1.CODE AVAILABILITY : The code generated during and/or analysed during the current study are available in the figshare repository https://doi.org/10.6084/m9.figshare.14368043.v1.Evaluating how intrinsic (intraspecific density), extrinsic (interspecific density and prey density) and anthropogenic (management intervention) factors affect African wild dog (Lycaon pictus) population performance is key to implementing effective conservation strategies. Lions (Panthera leo) can affect wild dog populations, and in small and highly managed protected areas, populations of wild dogs and lions often co-occur at high densities. It is unknown what mediates these co-occurring high densities and how trophic interactions facilitate the persistence of wild dogs in areas with high lion densities. In this study, we modelled how intrinsic and extrinsic factors affected population change and survival of 590 individually identifiable wild dogs in Hluhluwe–iMfolozi Park (HiP), South Africa, an area with high lion density. The wild dog population in HiP grew significantly and comprised one of the highest population densities and the largest pack sizes in Africa. Pup survival rate was also the highest known in Africa, and the median individual survival was 29 months. During low lion density periods, the rate of wild dog population change increased despite low prey density, while at higher lion density, the rate of population change decreased only when prey density decreased. Survival for all age classes increased as population density increased, suggesting there is an important density-dependent effect on survival for all age classes that manifests at the population level. While increasing lion density had negative effects on adult and yearling survival, it did not affect pup survival. Our results suggest that both intrinsic and extrinsic drivers regulated the HiP wild dog population. Importantly, populations of wild dogs in small (~ 900 km2), fenced protected areas are highly adaptable and can co-exist at high levels of interspecific competition over many generations as long as there is an intact and abundant prey base.The National Research Foundation of South Africa Scarce Skills Development Fund; the Rufford Small Grants Foundation; the Endangered Wildlife Trust (South Africa); the Wildlife ACT Fund (South Africa) and the Claude Leon Foundation.https://link.springer.com/journal/429912023-09-06hj2023Mammal Research InstituteZoology and Entomolog

Top–down limits on prey populations may be more severe in larger prey species, despite having fewer predators

Variation in the vulnerability of herbivore prey to predation is linked to body size, yet whether this relationship is size‐nested or size‐partitioned remains debated. If size‐partitioned, predators would be focused on prey within their preferred prey size range. If size‐nested, smaller prey species should become increasingly more vulnerable because increasingly more predators are capable of catching them. Yet, whether either of these strategies manifests in top–down prey population limitation would depend both on the number of potential predator species as well as the total mortality imposed. Here we use a rare ecosystem scale ‘natural experiment’ comparing prey population dynamics between a period of intense predator persecution and hence low predator densities and a period of active predator protection and population recovery. We use three decades of data on herbivore abundance and distribution to test the role of predation as a mechanism of population limitation among prey species that vary widely in body size. Notably, we test this within one of the few remaining systems where a near‐full suite of megaherbivores occur in high density and are thus able to include a thirtyfold range in herbivore body size gradient. We test whether top–down limitation on prey species of particular body size leads to compositional shifts in the mammalian herbivore community. Our results support both size‐nested and size‐partitioning predation but suggest that the relative top–down limiting impact on prey populations may be more severe for intermediate sized species, despite having fewer predators than small species. In addition we show that the gradual recovery of predator populations shifted the herbivore community assemblage towards large‐bodied species and has led to a community that is strongly dominated by large herbivore biomass.The Earthwatch Institute, Ezemvelo KwaZulu-Natal Wildlife and a Marie Curie Grant held by JC (grant # PCIG10-GA-2011-304128). ElR was supported by the South African National Research Foundation, Nelson Mandela Univ. and the Claude Leon Foundation.http://www.ecography.org2020-06-01hj2019Centre for Wildlife Managemen

TRY plant trait database – enhanced coverage and open access

Plant traits—the morphological, anatomical, physiological, biochemical and phenological characteristics of plants—determine how plants respond to environmental factors, affect other trophic levels, and influence ecosystem properties and their benefits and detriments to people. Plant trait data thus represent the basis for a vast area of research spanning from evolutionary biology, community and functional ecology, to biodiversity conservation, ecosystem and landscape management, restoration, biogeography and earth system modelling. Since its foundation in 2007, the TRY database of plant traits has grown continuously. It now provides unprecedented data coverage under an open access data policy and is the main plant trait database used by the research community worldwide. Increasingly, the TRY database also supports new frontiers of trait-based plant research, including the identification of data gaps and the subsequent mobilization or measurement of new data. To support this development, in this article we evaluate the extent of the trait data compiled in TRY and analyse emerging patterns of data coverage and representativeness. Best species coverage is achieved for categorical traits—almost complete coverage for ‘plant growth form’. However, most traits relevant for ecology and vegetation modelling are characterized by continuous intraspecific variation and trait–environmental relationships. These traits have to be measured on individual plants in their respective environment. Despite unprecedented data coverage, we observe a humbling lack of completeness and representativeness of these continuous traits in many aspects. We, therefore, conclude that reducing data gaps and biases in the TRY database remains a key challenge and requires a coordinated approach to data mobilization and trait measurements. This can only be achieved in collaboration with other initiatives.Rest of authors: Decky Junaedi, Robert R. Junker, Eric Justes, Richard Kabzems, Jeffrey Kane, Zdenek Kaplan, Teja Kattenborn, Lyudmila Kavelenova, Elizabeth Kearsley, Anne Kempel, Tanaka Kenzo, Andrew Kerkhoff, Mohammed I. Khalil, Nicole L. Kinlock, Wilm Daniel Kissling, Kaoru Kitajima, Thomas Kitzberger, Rasmus Kjøller, Tamir Klein, Michael Kleyer, Jitka Klimešová, Joice Klipel, Brian Kloeppel, Stefan Klotz, Johannes M. H. Knops, Takashi Kohyama, Fumito Koike, Johannes Kollmann, Benjamin Komac, Kimberly Komatsu, Christian König, Nathan J. B. Kraft, Koen Kramer, Holger Kreft, Ingolf Kühn, Dushan Kumarathunge, Jonas Kuppler, Hiroko Kurokawa, Yoko Kurosawa, Shem Kuyah, Jean-Paul Laclau, Benoit Lafleur, Erik Lallai, Eric Lamb, Andrea Lamprecht, Daniel J. Larkin, Daniel Laughlin, Yoann Le Bagousse-Pinguet, Guerric le Maire, Peter C. le Roux, Elizabeth le Roux, Tali Lee, Frederic Lens, Simon L. Lewis, Barbara Lhotsky, Yuanzhi Li, Xine Li, Jeremy W. Lichstein, Mario Liebergesell, Jun Ying Lim, Yan-Shih Lin, Juan Carlos Linares, Chunjiang Liu, Daijun Liu, Udayangani Liu, Stuart Livingstone, Joan Llusià, Madelon Lohbeck, Álvaro López-García, Gabriela Lopez-Gonzalez, Zdeňka Lososová, Frédérique Louault, Balázs A. Lukács, Petr Lukeš, Yunjian Luo, Michele Lussu, Siyan Ma, Camilla Maciel Rabelo Pereira, Michelle Mack, Vincent Maire, Annikki Mäkelä, Harri Mäkinen, Ana Claudia Mendes Malhado, Azim Mallik, Peter Manning, Stefano Manzoni, Zuleica Marchetti, Luca Marchino, Vinicius Marcilio-Silva, Eric Marcon, Michela Marignani, Lars Markesteijn, Adam Martin, Cristina Martínez-Garza, Jordi Martínez-Vilalta, Tereza Mašková, Kelly Mason, Norman Mason, Tara Joy Massad, Jacynthe Masse, Itay Mayrose, James McCarthy, M. Luke McCormack, Katherine McCulloh, Ian R. McFadden, Brian J. McGill, Mara Y. McPartland, Juliana S. Medeiros, Belinda Medlyn, Pierre Meerts, Zia Mehrabi, Patrick Meir, Felipe P. L. Melo, Maurizio Mencuccini, Céline Meredieu, Julie Messier, Ilona Mészáros, Juha Metsaranta, Sean T. Michaletz, Chrysanthi Michelaki, Svetlana Migalina, Ruben Milla, Jesse E. D. Miller, Vanessa Minden, Ray Ming, Karel Mokany, Angela T. Moles, Attila Molnár V, Jane Molofsky, Martin Molz, Rebecca A. Montgomery, Arnaud Monty, Lenka Moravcová, Alvaro Moreno-Martínez, Marco Moretti, Akira S. Mori, Shigeta Mori, Dave Morris, Jane Morrison, Ladislav Mucina, Sandra Mueller, Christopher D. Muir, Sandra Cristina Müller, François Munoz, Isla H. Myers-Smith, Randall W. Myster, Masahiro Nagano, Shawna Naidu, Ayyappan Narayanan, Balachandran Natesan, Luka Negoita, Andrew S. Nelson, Eike Lena Neuschulz, Jian Ni, Georg Niedrist, Jhon Nieto, Ülo Niinemets, Rachael Nolan, Henning Nottebrock, Yann Nouvellon, Alexander Novakovskiy, The Nutrient Network, Kristin Odden Nystuen, Anthony O'Grady, Kevin O'Hara, Andrew O'Reilly-Nugent, Simon Oakley, Walter Oberhuber, Toshiyuki Ohtsuka, Ricardo Oliveira, Kinga Öllerer, Mark E. Olson, Vladimir Onipchenko, Yusuke Onoda, Renske E. Onstein, Jenny C. Ordonez, Noriyuki Osada, Ivika Ostonen, Gianluigi Ottaviani, Sarah Otto, Gerhard E. Overbeck, Wim A. Ozinga, Anna T. Pahl, C. E. Timothy Paine, Robin J. Pakeman, Aristotelis C. Papageorgiou, Evgeniya Parfionova, Meelis Pärtel, Marco Patacca, Susana Paula, Juraj Paule, Harald Pauli, Juli G. Pausas, Begoña Peco, Josep Penuelas, Antonio Perea, Pablo Luis Peri, Ana Carolina Petisco-Souza, Alessandro Petraglia, Any Mary Petritan, Oliver L. Phillips, Simon Pierce, Valério D. Pillar, Jan Pisek, Alexandr Pomogaybin, Hendrik Poorter, Angelika Portsmuth, Peter Poschlod, Catherine Potvin, Devon Pounds, A. Shafer Powell, Sally A. Power, Andreas Prinzing, Giacomo Puglielli, Petr Pyšek, Valerie Raevel, Anja Rammig, Johannes Ransijn, Courtenay A. Ray, Peter B. Reich, Markus Reichstein, Douglas E. B. Reid, Maxime Réjou-Méchain, Victor Resco de Dios, Sabina Ribeiro, Sarah Richardson, Kersti Riibak, Matthias C. Rillig, Fiamma Riviera, Elisabeth M. R. Robert, Scott Roberts, Bjorn Robroek, Adam Roddy, Arthur Vinicius Rodrigues, Alistair Rogers, Emily Rollinson, Victor Rolo, Christine Römermann, Dina Ronzhina, Christiane Roscher, Julieta A. Rosell, Milena Fermina Rosenfield, Christian Rossi, David B. Roy, Samuel Royer-Tardif, Nadja Rüger, Ricardo Ruiz-Peinado, Sabine B. Rumpf, Graciela M. Rusch, Masahiro Ryo, Lawren Sack, Angela Saldaña, Beatriz Salgado-Negret, Roberto Salguero-Gomez, Ignacio Santa-Regina, Ana Carolina Santacruz-García, Joaquim Santos, Jordi Sardans, Brandon Schamp, Michael Scherer-Lorenzen, Matthias Schleuning, Bernhard Schmid, Marco Schmidt, Sylvain Schmitt, Julio V. Schneider, Simon D. Schowanek, Julian Schrader, Franziska Schrodt, Bernhard Schuldt, Frank Schurr, Galia Selaya Garvizu, Marina Semchenko, Colleen Seymour, Julia C. Sfair, Joanne M. Sharpe, Christine S. Sheppard, Serge Sheremetiev, Satomi Shiodera, Bill Shipley, Tanvir Ahmed Shovon, Alrun Siebenkäs, Carlos Sierra, Vasco Silva, Mateus Silva, Tommaso Sitzia, Henrik Sjöman, Martijn Slot, Nicholas G. Smith, Darwin Sodhi, Pamela Soltis, Douglas Soltis, Ben Somers, Grégory Sonnier, Mia Vedel Sørensen, Enio Egon Sosinski Jr, Nadejda A. Soudzilovskaia, Alexandre F. Souza, Marko Spasojevic, Marta Gaia Sperandii, Amanda B. Stan, James Stegen, Klaus Steinbauer, Jörg G. Stephan, Frank Sterck, Dejan B. Stojanovic, Tanya Strydom, Maria Laura Suarez, Jens-Christian Svenning, Ivana Svitková, Marek Svitok, Miroslav Svoboda, Emily Swaine, Nathan Swenson, Marcelo Tabarelli, Kentaro Takagi, Ulrike Tappeiner, Rubén Tarifa, Simon Tauugourdeau, Cagatay Tavsanoglu, Mariska te Beest, Leho Tedersoo, Nelson Thiffault, Dominik Thom, Evert Thomas, Ken Thompson, Peter E. Thornton, Wilfried Thuiller, Lubomír Tichý, David Tissue, Mark G. Tjoelker, David Yue Phin Tng, Joseph Tobias, Péter Török, Tonantzin Tarin, José M. Torres-Ruiz, Béla Tóthmérész, Martina Treurnicht, Valeria Trivellone, Franck Trolliet, Volodymyr Trotsiuk, James L. Tsakalos, Ioannis Tsiripidis, Niklas Tysklind, Toru Umehara, Vladimir Usoltsev, Matthew Vadeboncoeur, Jamil Vaezi, Fernando Valladares, Jana Vamosi, Peter M. van Bodegom, Michiel van Breugel, Elisa Van Cleemput, Martine van de Weg, Stephni van der Merwe, Fons van der Plas, Masha T. van der Sande, Mark van Kleunen, Koenraad Van Meerbeek, Mark Vanderwel, Kim André Vanselow, Angelica Vårhammar, Laura Varone, Maribel Yesenia Vasquez Valderrama, Kiril Vassilev, Mark Vellend, Erik J. Veneklaas, Hans Verbeeck, Kris Verheyen, Alexander Vibrans, Ima Vieira, Jaime Villacís, Cyrille Violle, Pandi Vivek, Katrin Wagner, Matthew Waldram, Anthony Waldron, Anthony P. Walker, Martyn Waller, Gabriel Walther, Han Wang, Feng Wang, Weiqi Wang, Harry Watkins, James Watkins, Ulrich Weber, James T. Weedon, Liping Wei, Patrick Weigelt, Evan Weiher, Aidan W. Wells, Camilla Wellstein, Elizabeth Wenk, Mark Westoby, Alana Westwood, Philip John White, Mark Whitten, Mathew Williams, Daniel E. Winkler, Klaus Winter, Chevonne Womack, Ian J. Wright, S. Joseph Wright, Justin Wright, Bruno X. Pinho, Fabiano Ximenes, Toshihiro Yamada, Keiko Yamaji, Ruth Yanai, Nikolay Yankov, Benjamin Yguel, Kátia Janaina Zanini, Amy E. Zanne, David Zelený, Yun-Peng Zhao, Jingming Zheng, Ji Zheng, Kasia Ziemińska, Chad R. Zirbel, Georg Zizka, Irié Casimir Zo-Bi, Gerhard Zotz, Christian Wirth.Max Planck Institute for Biogeochemistry; Max Planck Society; German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig; International Programme of Biodiversity Science (DIVERSITAS); International Geosphere-Biosphere Programme (IGBP); Future Earth; French Foundation for Biodiversity Research (FRB); GIS ‘Climat, Environnement et Société'.http://wileyonlinelibrary.com/journal/gcbhj2021Plant Production and Soil Scienc

Habitat heterogeneity as a driver of ungulate diversity and distribution patterns: interaction of body mass and digestive strategy

Aim: Classic island biogeographical theory predicts that reserves have to be large to conserve high biodiversity. Recent literature, however, suggests that habitat heterogeneity can counterbalance the effect of small reserve size. For savanna ungulates, body mass is said to drive habitat selection and facilitate species coexistence, where large species use a higher proportion of the landscape than smaller species, because a wider food quality tolerance allows them to use a higher diversity of habitat types. In this case, high habitat heterogeneity would facilitate diverse assemblages of different-sized ungulates. Digestive physiology should further modify this relationship, because non-ruminants have a wider diet tolerance than ruminants. We tested this hypothesis with an empirical dataset on distribution and habitat preference of different-sized African grazers. Location: Hluhluwe-iMfolozi Park, Republic of South Africa. Methods: We recorded herbivore dung and habitat type on 24 line transects varying between 4 and 11 km with a total length of 190 km to determine habitat selection and landscape distribution of six grazer species, three ruminants and three non-ruminants. Results: Larger ruminant grazers were more evenly distributed than smaller ruminants, had a more diverse use of habitats and used more low quality habitat. In contrast, non-ruminant grazers were more evenly distributed than similar-sized ruminants and body mass did not clearly influence diversity of habitat use and use of low quality habitat. Main conclusions: We confirm that body mass influences diversity of habitat use of large herbivores but digestive strategy potentially modifies this relationship. Hence, habitat heterogeneity might facilitate herbivore diversity in savanna ecosystems and high heterogeneity might counterbalance the effects of fragmentation and declining reserve size. Concluding, processes that homogenize the landscape, such as fire (mis)management and artificial waterholes, might be as threatening to biodiversity as landscape fragmentation, especially for smaller ruminant herbivores.

Elephant effects on treefall and logfall highlight the absence of megaherbivores in coarse woody debris conceptual frameworks

Despite the importance of coarse woody debris (CWD) in woody ecosystems, conceptual frameworks of its dynamics currently exclude the role of the megaherbivores, focusing instead on the role of insects, disease, fire, wind and droughts. However, recognizing the ecological roles of the megaherbivores is one of the most urgent contemporary issues, particularly as their decline will likely have unanticipated outcomes at the ecosystem-level. Here we used sites with and without elephants in a semi-arid woodland ecosystem in South Africa to test whether elephants changed the quantitative and qualitative features of the CWD profile in relation to other wood producers. We show that elephants increased the abundance of CWD, influenced its quality by toppling mostly maturing trees, and changed the distribution of large woody items along gentle hillslopes. Surprisingly, and despite the recognized importance of both CWD and elephants for biodiversity, there is almost no published data on the role of elephants in shaping wood-living communities by changing fallen woody debris dynamics. Our study contributes towards developing a broader conceptual framework of CWD dynamics that includes the role of megaherbivores and provides a novel view of the ecological consequences of the loss of the Pleistocene megaherbivores. Our findings have important implications for CWD restoration efforts in transformed systems that previously supported megaherbivores

Managing invasions at the cost of native habitat? An experimental test of the impact of fire on the invasion of Chromolaena odorata in a South African savanna

Successfully managing invasive plants in natural systems is extremely difficult. Recently however, progress has been made with an approach focused on changing ecosystem processes through the disturbance regime. We performed a large-scale (3 ha) full-factorial field experiment in densely invaded woodland in Hluhluwe-iMfolozi Park, a savanna reserve in South Africa, to study the effect of fire on the control of the pan-tropical invasive exotic shrub Chromolaena odorata in combination with the conventional method, i.e. manual clearing and herbicide application. We show how fire interacted with the conventional clearing of C. odorata and induced an intense canopy fire that caused a shift from woodland to grassland. After 2.5 years of monitoring, grasses were still dominant and re-invasion minimal. It is important to note that fire without prior clearing did not have the same effect and was not successful in reducing densities of C. odorata. An integrated control practice targeting the species with mechanical and chemical methods, while simultaneously targeting its habitat through fire, effectively controlled dense C. odorata thickets during the course of the experiment. However, this approach transformed regular surface fires into high-intensity canopy fires that are rare in savannas. We discuss how this altered fire regime may threaten native habitats, including fire-sensitive forest patches and riverine woodlands within the savanna mozaic. This is an important dilemma for managers that should not be overlooked and asks for long-term data on the impact of control programs on the native vegetation.

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.