Corinna Riginos

Grass competition suppresses savanna tree growth across multiple demographic stages

Savanna ecosystems, defined by the codominance of trees and grasses, cover one-fifth of the worlds land surface and are of great socioeconomic and biological importance. Yet, the fundamental question of how trees and grasses coexist to maintain the savanna state remains poorly understood. Many models have been put forward to explain tree-grass coexistence, but nearly all have assumed that grasses do not limit tree growth and demography beyond the sapling stage. This assumption, however, has rarely been tested. Here I show that grass can strongly suppress the growth of trees. I removed grass around trees of three size classes in an Acacia drepanolobium savanna in Laikipia, Kenya. For even the largest trees, grass removal led to a doubling in growth and a doubling in the probability of transitioning to the next size class over two years. These results suggest that grass competition in productive (nutrient-rich) savannas may limit tree growth as much as herbivory and fire (the main factors thought to determine tree demography within a rainfall region) and should be incorporated into savanna models if tree-grass coexistence and savanna dynamics are to be understood.

Context-dependent interactions between adult shrubs and seedlings in a semi-arid shrubland

Abstract Question: In semi-arid systems, rainfall gradients can cause plant-plant interactions to shift from negative to positive or vice versa. However, the importance of a second major abiotic factor, soil nutrients, has rarely been considered. We consider different combinations of both factors and ask: do net adult-seedling interactions become less competitive and more facilitative with increasing overall abiotic harshness? Location: Succulent Karoo, Western Cape, South Africa. Methods: We examined the interactions between seedlings and adult shrubs at two sites. Sites differ in rainfall, and each contain two habitats: Nutrient-rich mounds associated with underground termitaria and a relatively nutrient-poor matrix. We carried out a spatial pattern analysis of community-wide seedling-adult associations. We then conducted field and greenhouse experiments to test the effects of soil and the presence of neighbouring shrubs on the growth and survival of six seedling species. Results: At the higher rainfall site, both competitive and facilitative effects of adults on seedlings were found but did not differ by habitat, despite the more benign conditions in the mound habitat. At the lower rainfall site, adult shrubs generally had neutral effects on seedlings in the matrix habitat. In the nutrient-rich mound habitat, however, adult shrubs had strong and consistently competitive effects on seedlings. Conclusion: Seedling-adult interactions could not be predicted by a simple overall gradient of abiotic harshness, demonstrating the need for more complex, mechanistic models to predict plant-plant interactions. We suggest that rainfall and soil nutrients affect seedling-adult relations through their interactive effects on the life-history attributes of the species involved. Nomenclature: Germishuizen & Meyer (2003).

Native and domestic browsers and grazers reduce fuels, fire temperatures, and acacia ant mortality in an African savanna

Despite the importance of fire and herbivory in structuring savanna systems, few replicated experiments have examined the interactive effects of herbivory and fire on plant dynamics. In addition, the effects of fire on associated ant-tree mutualisms have been largely unexplored. We carried out small controlled burns in each of 18 herbivore treatment plots of the Kenya Long-term Exclosure Experiment (KLEE), where experimentally excluding elephants has resulted in 42% greater tree densities. The KLEE design includes six different herbivore treatments that allowed us to examine how different combinations of megaherbivore wildlife, mesoherbivore wildlife, and cattle affect fire temperatures and subsequent loss of ant symbionts from Acacia trees. Before burning, we quantified herbaceous fuel loads and plant community composition. We tagged all trees, measured their height and basal diameter, and identified the resident ant species on each. We recorded weather conditions during the burns and used ceramic tiles painted with fire-sensitive paints to estimate fire temperatures at different heights and in different microsites (under vs. between trees). Across all treatments, fire temperatures were highest at 0-50 cm off the ground and hotter in the grass under trees than in the grassy areas between trees. Plots with more trees burned hotter than plots with fewer trees, perhaps because of greater fine woody debris. Plots grazed by wildlife and by cattle prior to burning had lower herbaceous fuel loads and experienced lower burn temperatures than ungrazed plots. Many trees lost their ant colonies during the burns. Ant survivorship differed by ant species and at the plot level was positively associated with previous herbivory (and lower fire temperatures). Across all treatments, ant colonies on taller trees were more likely to survive, but even some of the tallest trees lost their ant colonies. Our study marks a significant step in understanding the mechanisms that underlie the interactions between fire and herbivory in savanna ecosystems.

Lessons on the relationship between livestock husbandry and biodiversity from the Kenya Long-term Exclosure Experiment (KLEE)

Although livestock and wildlife share most of their ranges worldwide, little controlled experimental research has been done on their interactions. Since 1995 we have been manipulating the presence of cattle and large wild ungulates in a Kenyan savanna rangeland in order to better understand the nature of competition and coexistence between these two guilds of herbivores and how they affect biodiversity. In a replicated experiment in which different combinations of cattle and wild herbivores are allowed access to large-scale plots, we have been monitoring the impacts of these herbivores on vegetation, on the wild herbivores, and cattle themselves, and on a variety of other taxa. We have also been conducting experimental research to examine other ways in which livestock management in eastern Africa might affect biodiversity. These include studies on the impacts of fire, livestock corrals, and changes in tree density. This research has revealed the following patterns. (1) Cattle suppress many species of wild herbivores, presumably through competition for their shared resources. The nature of this competition, however, is contingent on rainfall and the presence of other herbivores. (2) Wild herbivores both compete with and facilitate cattle, depending on rainfall. (3) The pastoral practice of housing livestock nightly in protective corral enclosures (“bomas”) over time produces long-lived nutrient hotspots preferred by both livestock and wild herbivores. (4) Fire, frequently used by pastoralists in the past, is valuable for improving grass quality, with benefits for many species of wild herbivores. (5) Pastoral practices that reduce woody cover, including burning and boma construction, create local habitat patches that are preferred by wild herbivores, apparently for their greater anti-predator visibility. (6) Despite competition between livestock and wild herbivores, coexistence between these two guilds can be managed, and there are several positive (facilitative) pathways between livestock husbandry and wild herbivores and other biodiversity.

Revolutionary land use change in the 21st century: is (Rangeland) science relevant?

Abstract Rapidly increasing demand for food, fiber, and fuel together with new technologies and the mobility of global capital are driving revolutionary changes in land use throughout the world. Efforts to increase land productivity include conversion of millions of hectares of rangelands to crop production, including many marginal lands with low resistance and resilience to degradation. Sustaining the productivity of these lands requires careful land use planning and innovative management systems. Historically, this responsibility has been left to agronomists and others with expertise in crop production. In this article, we argue that the revolutionary land use changes necessary to support national and global food security potentially make rangeland science more relevant now than ever. Maintaining and increasing relevance will require a revolutionary change in range science from a discipline that focuses on a particular land use or land cover to one that addresses the challenge of managing all lands that, at one time, were considered to be marginal for crop production. We propose four strategies to increase the relevance of rangeland science to global land management: 1) expand our awareness and understanding of local to global economic, social, and technological trends in order to anticipate and identify drivers and patterns of conversion; 2) emphasize empirical studies and modeling that anticipate the biophysical (ecosystem services) and societal consequences of large-scale changes in land cover and use; 3) significantly increase communication and collaboration with the disciplines and sectors of society currently responsible for managing the new land uses; and 4) develop and adopt a dynamic and flexible resilience-based land classification system and data-supported conceptual models (e.g., state-and-transition models) that represent all lands, regardless of use and the consequences of land conversion to various uses instead of changes in state or condition that are focused on a single land use. Resumen La creciente demanda de alimentos, fibras y combustibles de manera simultánea con las nuevas tecnologías y la movilidad global del capital están ocasionando cambios revolucionados en el uso de la tierra en todo el mundo. Los esfuerzos para incrementar la productividad de la tierra incluyen la conversión de millones de hectáreas de pastizales a la producción de granos, incluyendo tierras marginales con bajo resistencia y resilencia a la degradación. Sostener la productividad de estas tierras requiere planeación cuidadosa del uso de la tierra y sistemas de manejo innovadores. Históricamente, esta responsabilidad se ha dejado a agrónomos y otros expertos en producción de granos. En este articulo, discutimos que los revolucionados cambios en uso de la tierra necesarios para sostener la seguridad alimentaria nacional y mundial potencialmente hacen a la ciencia del pastizal más relevante ahora que nunca. Mantener e incrementar esa relevancia requerirá de cambios revolucionarios en la ciencia del pastizal de una disciplina que se enfoca en un uso particular de la tierra o cubierta vegetal a una que considere el reto de manejar todas las tierras que en algún tiempo fueron consideradas marginales para la producción de granos. Proponemos cuatro estrategias para aumentar la relevancia de la ciencia del pastizal a un manejo global de la tierra: 1) extender nuestra conocimiento y concientización del ámbito local a tendencias globales económicas, sociales y tecnológicas con el fin de anticipar e identificar conductores y patrones de conversión, 2) enfatizar en estudios empíricos y modelaje que anticipe las consecuencias biofísicas (servicios de los ecosistemas) y sociales de cambios en la cobertura y uso de la tierra en gran escala, 3) aumentar significativamente la comunicación y colaboración con las disciplinas y sectores de la sociedad actualmente responsables en el manejo del nuevo uso de la tierra, y 4) desarrollar y adoptar un sistema de clasificación dinámica y flexible basado en la resilencia de la tierra y modelos conceptuales apoyados en datos (ejm. Modelos de Estado y Transición) que representan todas las tierras, independientemente del uso y las consecuencias en la conversión de tierras para varios usos el lugar de cambios en el estado y condición que se enfocan en un solo uso de la tierra.

Glade cascades: indirect legacy effects of pastoralism enhance the abundance and spatial structuring of arboreal fauna

Studies in community ecology are typically conducted over the span of a few years, and results are often interpreted as the product of contemporary processes and interactions. All landscapes have histories, however, and observed patterns of distribution and abundance frequently reflect enduring legacies of past ecological events, the existence and influence of which may not be obvious to investigators. In East Africa, most wildlife occurs outside national reserves and often coexists there with livestock, which are traditionally corralled at night in temporary thorn fence enclosures, or bomas. After being abandoned, bomas develop into nutrient-rich, treeless glades that can persist for more than a century. These hotspots of primary productivity attract both native and domestic large herbivores, but the extent to which their effects cascade to other consumers is unknown. Here, we document positive edge effects of glades on the mean size and growth rates of Acacia trees and show that the density and biomass of arboreal geckos (Lygodactylus keniensis) are elevated near glades and decrease with distance from glades. The edge response of geckos is an indirect effect arising from the positive influence of glades on arboreal arthropod biomass (a trophic effect) and average tree size (a non-trophic effect). By clearing plots of trees to simulate glades, we experimentally demonstrate that these legacy effects arise from the elevated nutrient content of glades as opposed to their distinctive structural features. Finally, we investigated interactions among glade edges, showing that legacy effects are dampened (rather than enhanced) by the presence of other glades nearby. Collectively, our results show that legacy effects of traditional pastoral practices cascade into the treetops, imparting spatial structure across multiple trophic levels in an otherwise homogeneous Kenyan savanna ecosystem.

Management and Analysis of Camera Trap Data: Alternative Approaches (Response to Harris et al. 2010)

A recent article in the ESA Bulletin introduces an approach to cataloging and analyzing the large numbers of photos typically generated by camera traps (Harris et al. 2010). The authors highlight the need for a simple, systematic way to archive and extract data from camera trap photos. The authors then present a system they have developed in an effort to meet this need. Here, we introduce two alternative approaches that each have distinct advantages over the approach outlined by Harris et al.

Herbivore Effects on Productivity Vary by Guild: Cattle Increase Mean Productivity While Wildlife Reduce Variability

Wild herbivores and livestock share the majority of rangelands worldwide, yet few controlled experiments have addressed their individual, additive, and interactive impacts on ecosystem function. While ungulate herbivores generally reduce standing biomass, their effects on aboveground net primary production (ANPP) can vary by spatial and temporal context, intensity of herbivory, and herbivore identity and species richness. Some evidence indicates that moderate levels of herbivory can stimulate aboveground productivity, but few studies have explicitly tested the relationships among herbivore identity, grazing intensity, and ANPP. We used a long-term exclosure experiment to examine the effects of three groups of wild and domestic ungulate herbivores (megaherbivores, mesoherbivore wildlife, and cattle) on herbaceous productivity in an African savanna. Using both field measurements (productivity cages) and satellite imagery, we measured the effects of different herbivore guilds, separately and in different combinations, on herbaceous productivity across both space and time. Results from both productivity cage measurements and satellite normalized difference vegetation index (NDVI) demonstrated a positive relationship between mean productivity and total ungulate herbivore pressure, driven in particular by the presence of cattle. In contrast, we found that variation in herbaceous productivity across space and time was driven by the presence of wild herbivores (primarily mesoherbivore wildlife), which significantly reduced heterogeneity in ANPP and NDVI across both space and time. Our results indicate that replacing wildlife with cattle (at moderate densities) could lead to similarly productive but more heterogeneous herbaceous plant communities in rangelands.

Influence of cattle on browsing and grazing wildlife varies with rainfall and presence of megaherbivores.

In many savanna ecosystems worldwide, livestock share the landscape and its resources with wildlife. The nature of interactions between livestock and wildlife is a subject of considerable interest and speculation, yet little controlled experimental research has been carried out. Since 1995, we have been manipulating the presence and absence of cattle and large mammalian herbivore wildlife in a Kenyan savanna in order to better understand how different herbivore guilds influence habitat use by specific wildlife species. Using dung counts as a relative assay of herbivore use of the different experimental plots, we found that cattle had a range of effects, mostly negative, on common mesoherbivore species, including both grazers and mixed feeders, but did not have significant effects on megaherbivores. The effect of cattle on most of the mesoherbivore species was contingent on both the presence of megaherbivores and rainfall. In the absence of megaherbivores, wild mesoherbivore dung density was 36% lower in plots that they shared with cattle than in plots they used exclusively, whereas in the presence of megaherbivores, wild mesoherbivore dung density was only 9% lower in plots shared with cattle than plots used exclusively. Cattle appeared to have a positive effect on habitat use by zebra (a grazer) and steinbuck (a browser) during wetter periods of the year but a negative effect during drier periods. Plots to which cattle had access had lower grass and forb cover than plots from which they were excluded, while plots to which megaherbivores had access had more grass cover but less forb cover. Grass cover was positively correlated with zebra and oryx dung density while forb cover was positively correlated with eland dung density. Overall these results suggest that interactions between livestock and wildlife are contingent on rainfall and herbivore assemblage and represent a more richly nuanced set of interactions than the longstanding assertion that cattle simply compete with (grazing) wildlife. Specifically, rainfall and megaherbivores seemed to moderate the negative effects of cattle on some mesoherbivore species. Even if cattle tend to reduce wildlife use of the landscape, managing simultaneously for livestock production (at moderate levels) and biodiversity conservation is possible.

Disruption of a protective ant-plant mutualism by an invasive ant increases elephant damage to savanna trees.

Invasive species can indirectly affect ecosystem processes via the disruption of mutualisms. The mutualism between the whistling thorn acacia (Acacia drepanolobium) and four species of symbiotic ants is an ecologically important one; ants strongly defend trees against elephants, which can otherwise have dramatic impacts on tree cover. In Laikipia, Kenya, the invasive big-headed ant (Pheidole megacephala) has established itself at numerous locations within the last 10-15 years. In invaded areas on five properties, we found that three species of symbiotic Crematogaster ants were virtually extirpated, whereas Tetraponera penzigi co-occurred with P. megacephala. T. penzigi appears to persist because of its nonaggressive behavior; in a whole-tree translocation experiment, Crematogaster defended host trees against P. megacephala, but were extirpated from trees within hours. In contrast, T. penzigi retreated into domatia and withstood invading ants for >30 days. In the field, the loss of defensive Crematogaster ants in invaded areas led to a five- to sevenfold increase in the number of trees catastrophically damaged by elephants compared to uninvaded areas. In savannas, tree cover drives many ecosystem processes and provides essential forage for many large mammal species; thus, the invasion of big-headed ants may strongly alter the dynamics and diversity of East Africas whistling thorn savannas by disrupting this systems keystone acaciaant mutualism.