Biodiversity Loss in Latin American Coffee Landscapes: Review of the Evidence on Ants,Birds, and Trees

Abstract: Studies have documented biodiversity losses due to intensification of coffee management (reduction in canopy richness and complexity). Nevertheless, questions remain regarding relative sensitivity of different taxa, habitat specialists, and functional groups, and whether implications for biodiversity conservation vary across regions. We quantitatively reviewed data from ant, bird, and tree biodiversity studies in coffee agroecosystems to address the following questions: Does species richness decline with intensification or with individual vegetation characteristics? Are there significant losses of species richness in coffee‐management systems compared with forests? Is species loss greater for forest species or for particular functional groups? and Are ants or birds more strongly affected by intensification? Across studies, ant and bird richness declined with management intensification and with changes in vegetation. Species richness of all ants and birds and of forest ant and bird species was lower in most coffee agroecosystems than in forests, but rustic coffee (grown under native forest canopies) had equal or greater ant and bird richness than nearby forests. Sun coffee (grown without canopy trees) sustained the highest species losses, and species loss of forest ant, bird, and tree species increased with management intensity. Losses of ant and bird species were similar, although losses of forest ants were more drastic in rustic coffee. Richness of migratory birds and of birds that forage across vegetation strata was less affected by intensification than richness of resident, canopy, and understory bird species. Rustic farms protected more species than other coffee systems, and loss of species depended greatly on habitat specialization and functional traits. We recommend that forest be protected, rustic coffee be promoted, and intensive coffee farms be restored by augmenting native tree density and richness and allowing growth of epiphytes. We also recommend that future research focus on potential trade‐offs between biodiversity conservation and farmer livelihoods stemming from coffee production.     

Ecological and Economic Services Provided by Birds on Jamaican Blue Mountain Coffee Farms

Abstract: Coffee farms can support significant biodiversity, yet intensification of farming practices is degrading agricultural habitats and compromising ecosystem services such as biological pest control. The coffee berry borer (Hypothenemus hampei) is the world's primary coffee pest. Researchers have demonstrated that birds reduce insect abundance on coffee farms but have not documented avian control of the berry borer or quantified avian benefits to crop yield or farm income. We conducted a bird‐exclosure experiment on coffee farms in the Blue Mountains, Jamaica, to measure avian pest control of berry borers, identify potential predator species, associate predator abundance and borer reductions with vegetation complexity, and quantify resulting increases in coffee yield. Coffee plants excluded from foraging birds had significantly higher borer infestation, more borer broods, and greater berry damage than control plants. We identified 17 potential predator species (73% were wintering Neotropical migrants), and 3 primary species composed 67% of migrant detections. Average relative bird abundance and diversity and relative resident predator abundance increased with greater shade‐tree cover. Although migrant predators overall did not respond to vegetation complexity variables, the 3 primary species increased with proximity to noncoffee habitat patches. Lower infestation on control plants was correlated with higher total bird abundance, but not with predator abundance or vegetation complexity. Infestation of fruit was 1–14% lower on control plants, resulting in a greater quantity of saleable fruits that had a market value of US$44–$105/ha in 2005/2006. Landscape heterogeneity in this region may allow mobile predators to provide pest control broadly, despite localized farming intensities. These results provide the first evidence that birds control coffee berry borers and thus increase coffee yield and farm income, a potentially important conservation incentive for producers.     

A simple plant mutation abets a predator-diversity cascade

Resource consumption often increases with greater consumer biodiversity. This could result either from complementarity among consumers or the inclusion of particular key species, and it is often difficult to differentiate between these two mechanisms. We exploited a simple plant mutation (reduced production of surface waxes) to alter foraging within a community of aphid predators, and thus perhaps shift the nature of resulting predator diversity effects. We found that greater predator species richness dramatically increased prey suppression and plant biomass only on mutant, reduced-wax pea plants (Pisum sativum). On pea plants from a sister line with wild type, waxier plant surfaces, predator species richness did not influence predators' impacts on herbivores or plants. Thus, a change in plant surface structure acted to turn on, or off, the cascading effects of predator diversity. Greater predator richness encouraged higher densities of true predators but did not lead to greater reproduction by a parasitoid, Aphidius ervi; fecundity of each natural enemy species was similar for the two plant types. Behavioral observations indicated that although A. ervi was less likely to forage within species-rich predator communities, low-wax plants mitigated this interference by encouraging generally greater A. ervi foraging and thus high rates of aphid dislodgement (aphids dropped from plants to escape A. ervi, but not the other predators). Thus, only species-rich, low-wax plants simultaneously encouraged strong species-specific effects of A. ervi, and strong complementarity among the other predator species. In summary, our study provides evidence that diversity effects in predator assemblages are sensitive to habitat characteristics. Further, we show that a simple plant morphological trait, controlled by a single gene mutation, can dramatically alter the cascading effects of predator species richness on herbivores and plants.     

Simulated predator extinctions: predator identity affects survival and recruitment of oysters

The rate of species loss is increasing at a global scale, and human-induced extinctions are biased toward predator species. We examined the effects of predator extinctions on a foundation species, the eastern oyster (Crassostrea virginica). We performed a factorial experiment manipulating the presence and abundance of three of the most common predatory crabs, the blue crab (Callinectes sapidus), stone crab (Menippe mercenaria), and mud crab (Panopeus herbstii) in estuaries in the eastern United States. We tested the effects of species richness and identity of predators on juvenile oyster survival, oyster recruitment, and organic matter content of sediment. We also manipulated the density of each of the predators and controlled for the loss of biomass of species by maintaining a constant mass of predators in one set of treatments and simultaneously using an additive design. This design allowed us to test the density dependence of our results and test for functional compensation by other species. The identity of predator species, but not richness, affected oyster populations. The loss of blue crabs, alone or in combination with either of the other species, affected the survival rate of juvenile oysters. Blue crabs and stone crabs both affected oyster recruitment and sediment organic matter negatively. Mud crabs at higher than ambient densities, however, could fulfill some of the functions of blue and stone crabs, suggesting a level of ecological redundancy. Importantly, the strong effects of blue crabs in all processes measured no longer occurred when individuals were present at higher-than-ambient densities. Their role as dominant predator is, therefore, dependent on their density within the system and the density of other species within their guild (e.g., mud crabs). Our findings support the hypothesis that the effects of species loss at higher trophic levels are determined by predator identity and are subject to complex intraguild interactions that are largely density dependent. Understanding the role of biodiversity in ecosystem functioning or addressing practical concerns, such as loss of predators owing to overharvesting, remains complicated because accurate predictions require detailed knowledge of the system and should be drawn from sound experimental evidence, not based on observations or generalized models.     

Herbivores on a dominant understory shrub increase local plant diversity in rain forest communities

Indirect effects of trophic interactions on biodiversity can be large and common, even in complex communities. Previous experiments with dominant understory Piper shrubs in a Costa Rican rain forest revealed that increases in herbivore densities on these shrubs caused widespread seedling mortality as a result of herbivores moving from Piper to seedlings of many different plant genera. We tested components of the Janzen-Connell hypothesis by conducting focused studies on the effects of specialist and generalist Piper herbivores on local seedling diversity. Whereas specialist herbivores are predicted to increase mortality to neighboring seedlings that are closely related to the source plant, true generalists moving from source plants may cause density-dependent mortality of many species, and possibly increase richness if new species replace abundant species that have been thinned by herbivores. Therefore, we hypothesized that seedling richness would be greater in understory control plots created in patches of Piper that had normal densities of generalist herbivores compared to plots from which we removed generalist herbivores manually from all Piper shrubs. After 15 months, generalist-herbivore-removal plots had > 40% fewer seedlings, > 40% fewer species, and 40% greater seedling evenness, on average, than control plots with generalist herbivores intact. Using a complementary approach in unmanipulated plots in four forests, we used path analysis to test for a positive association between seedling diversity and herbivore damage on Piper species. In unmanipulated plots, for both generalist and specialist herbivores, our data were significant fits to the causal model that Piper herbivores decrease evenness and increase plant species richness, corroborating the experimental results. Because herbivores changed how individuals were apportioned among the species and families present (lower evenness), one interpretation of these associations between herbivores on Piper shrubs and local seedling richness is that high seedling mortality in dominant families allowed the colonization or survival of less common species. If interspecific or apparent competition allowed for a relative increase in species richness, then the Janzen-Connell hypothesis may extend its predictions to generalist seedling predators. We speculate that apparent competition may explain some of the deviations from neutral model predictions, especially at small scales.     

Savanna tree density, herbivores, and the herbaceous community: bottom-up vs. top-down effects

Herbivores choose their habitats both to maximize forage intake and to minimize their risk of predation. For African savanna herbivores, the available habitats range in woody cover from open areas with few trees to dense, almost-closed woodlands. This variation in woody cover or density can have a number of consequences for herbaceous species composition, cover, and productivity, as well as for ease of predator detection and avoidance. Here, we consider two alternative possibilities: first, that tree density affects the herbaceous vegetation, with concomitant "bottom-up" effects on herbivore habitat preferences; or, second, that tree density affects predator visibility, mediating "top-down" effects of predators on herbivore habitat preferences. We sampled sites spanning a 10-fold range of tree densities in an Acacia drepanolobium-dominated savanna in Laikipia, Kenya, for variation in (1) herbaceous cover, composition, and species richness; (2) wild and domestic herbivore use; and (3) degree of visibility obstruction by the tree layer. We then used structural equation modeling to consider the potential influences that tree density may have on herbivores and herbaceous community properties. Tree density was associated with substantial variation in herbaceous species composition and richness. Cattle exhibited a fairly uniform use of the landscape, whereas wild herbivores, with the exception of elephants, exhibited a strong preference for areas of low tree density. Model results suggest that this was not a response to variation in herbaceous-community characteristics, but rather a response to the greater visibility associated with more open places. Elephants, in contrast, preferred areas with higher densities of trees, apparently because of greater forage availability. These results suggest that, for all but the largest species, top-down behavioral effects of predator avoidance on herbivores are mediated by tree density. This, in turn, appears to have cascading effects on the herbaceous vegetation. These results shed light on one of the major features of the "landscape of fear" in which African savanna herbivores exist.     

Spatial pattern and ecological process in the coffee agroforestry system

The coffee agroforestry system provides an ideal platform for the study of spatial ecology. The uniform pattern of the coffee plants and shade trees allows for the study of pattern generation through intrinsic biological forces rather than extrinsic habitat patchiness. Detailed studies, focusing on a key mutualism between an ant (Azteca instabilis) and a scale insect (Coccus viridis), conducted in a 45-ha plot in a coffee agroforestry system have provided insights into (1) the quantitative evaluation of spatial pattern of the scale insect Coccus viridis on coffee bushes, (2) the mechanisms for the generation of patterns through the combination of local satellite ant nest formation and regional control from natural enemies, and (3) the consequences of the spatial pattern for the stability of predator-prey (host-parasitoid) systems, for a key coccinelid beetle preying on the scale insects and a phorid fly parasitoid parasitizing the ant.     

Reduced Diversity and Complexity in the Leaf-Litter Ant Assemblage of Colombian Coffee Plantations

Abstract:  Coffee agroecosystems have recently undergone a dramatic intensification in Colombia, a megadiverse country, especially in terms of the nature of shade cover. We tested for changes in the composition, ecological associations, and diversity of ants (Hymenoptera: Formicidae) along a gradient of intensification of coffee production in the Colombian Andes. We surveyed 16 farms in two regions, classified into four management types: (1) forest (no agriculture), (2) organic polygeneric shaded coffee, (3) monogeneric shaded coffee, and (4) sun coffee (unshaded). Forty sampling units (20 1-m² plots on the ground and 20 coffee bushes) were established at each farm between 2001 and 2002. We sampled with a mini-winkler litter extraction technique and through visual searching. Organic polygeneric shaded-coffee plantations contained significantly higher ant species richness, and their ant assemblages resembled the forest patches more than any other management type. The number of statistically significant associations among ant species dropped with production intensification, as did the number of ant species involved in such associations. The network of ant associations in shaded systems transformed into an extremely simplified network in sun coffee, with a few dominant ants extending almost entirely throughout the crop. Intensification of coffee agriculture not only caused loss of litter ant species (especially forest species) but also a reduction in the complexity of the ant assemblage in the leaf litter of this agroecosystem.     

Evaluating Effects of Habitat Loss and Land-Use Continuity on Ant Species Richness in Seminatural Grassland Remnants

Abstract:  Seminatural grasslands in Europe are susceptible to habitat destruction and fragmentation that result in negative effects on biodiversity because of increased isolation and area effects on extinction rate. However, even small habitat patches of seminatural grasslands might be of value for conservation and restoration of species richness in a landscape with a long history of management, which has been argued to lead to high species richness. We tested whether ant communities have been negatively affected by habitat loss and increased isolation of seminatural grasslands during the twentieth century. We examined species richness and community composition in seminatural grasslands of different size in a mosaic landscape in Central Sweden. Grasslands managed continuously over centuries harbored species-rich and ecologically diverse ant communities. Grassland remnant size had no effect on ant species richness. Small grassland remnants did not harbor a nested subset of the ant species of larger habitats. Community composition of ants was mainly affected by habitat conditions. Our results suggest that the abandonment of traditional land use and the encroachment of trees, rather than the effects of fragmentation, are important for species composition in seminatural grasslands. Our results highlight the importance of considering land-use continuity and dispersal ability of the focal organisms when examining the effects of habitat loss and fragmentation on biodiversity. Landscape history should be considered in conservation programs focusing on effects of land-use change.     

Predator diversity and identity drive interaction strength and trophic cascades in a food web

Declining predator diversity may drastically affect the biomass and productivity of herbivores and plants. Understanding how changes in predator diversity can propagate through food webs to alter ecosystem function is one of the most challenging ecological research topics today. We studied the effects of predator removal in a simple natural food web in the Sierra Nevada mountains of California (USA). By excluding the predators of the third trophic level of a food web in a full-factorial design, we monitored cascading effects of varying predator diversity and composition on the herbivorous beetle Chrysomela aeneicollis and the willow Salix orestera, which compose the first and second trophic levels of the food web. Decreasing predator diversity increased herbivore biomass and survivorship, and consequently increased the amount of plant biomass consumed via a trophic cascade. Despite this simple linear mean effect of diversity on the strength of the trophic cascade, we found additivity, compensation, and interference in the effects of multiple predators on herbivores and plants. Herbivore survivorship and predator-prey interaction strengths varied with predator diversity, predator identity, and the identity of coexisting predators. Additive effects of predators on herbivores and plants may have been driven by temporal niche separation, whereas compensatory effects and interference occurred among predators with a similar phenology. Together, these results suggest that while the general trends of diversity effects may appear linear and additive, other information about species identity was required to predict the effects of removing individual predators. In a community that is not temporally well-mixed, predator traits such as phenology may help predict impacts of species loss on other species. Information about predator natural history and food web structure may help explain variation in predator diversity effects on trophic cascades and ecosystem function.     

Species identity dominates the relationship between predator biodiversity and herbivore suppression

Agricultural pest suppression is an important ecosystem service that may be threatened by the loss of predator diversity. This has stimulated interest in the relationship between predator biodiversity and biological control. Multiple-predator studies have shown that predators may complement or interfere with one another, but few experiments have determined if the resulting effects on prey are caused by changes in predator abundance, identity, species richness, or some combination of these factors. We experimentally isolated the effect of predator species richness on the biological control of an important agricultural pest, the green peach aphid. We found no evidence that increasing predator species richness affects aphid biological control; overall there was no strong complementarity or interference among predator species that altered the strength of aphid suppression. Instead, our experiments revealed strong effects of predator species identity, because predators varied dramatically in their per capita consumption rates. Our results are consistent with other multiple-predator studies finding strong species-identity effects and suggest that, for the biological control of aphids, conservation strategies that directly target key species will be more effective than those targeting predator biodiversity more broadly.     

Functional diversity loss and change in nocturnal behavior of mammals under anthropogenic disturbance

In the Anthropocene, understanding the impacts of anthropogenic influence on biodiversity and behavior of vulnerable wildlife communities is increasingly relevant to effective conservation. However, comparative studies aimed at disentangling the concurrent effect of different types of human disturbance on multifaceted biodiversity and on activity patterns of mammals are surprisingly rare. We applied a multiregion community model to separately estimate the effects of cumulative human modification (e.g., settlement, agriculture, and transportation) and human presence (aggregated presence of dogs, people, and livestock) on species richness and functional composition of medium- and large-bodied mammals based on camera trap data collected across 45 subtropical montane forests. We divided the detected mammal species into three trophic guilds–carnivores, herbivores, and omnivores–and assessed the nocturnal shifts of each guild in response to anthropogenic activities. Overall, species richness tended to increase (β coefficient = 0.954) as human modification increased but richness decreased as human presence increased (β = –1.054). Human modification was associated with significantly lower functional diversity (mean nearest taxon distance [MNTD], β = –0.134; standardized effect sizes of MNTD, β = –0.397), community average body mass (β = –0.240), and proportion of carnivores (β = –0.580). Human presence was associated with a strongly reduced proportion of herbivores (β = –0.522), whereas proportion of omnivores significantly increased as human presence (β = 0.378) and habitat modification (β = 0.419) increased. In terms of activity patterns, omnivores (β = 12.103) and carnivores (β = 9.368) became more nocturnal in response to human modification. Our results suggest that human modification and human presence have differing effects on mammals and demonstrate that anthropogenic disturbances can lead to drastic loss of functional diversity and result in a shift to nocturnal behavior of mammals. Conservation planning should consider concurrent effects of different types of human disturbance on species richness, functional diversity, and behavior of wildlife communities.     

Multiple functions increase the importance of biodiversity for overall ecosystem functioning

Biodiversity is proposed to be important for the rate of ecosystem functions. Most biodiversity-ecosystem function studies, however, consider only one response variable at a time, and even when multiple variables are examined they are analyzed separately. This means that a very important aspect of biodiversity is overlooked: the possibility for different species to carry out different functions at any one time. We propose a conceptual model to explore the effects of species loss on overall ecosystem functioning, where overall functioning is defined as the joint effect of many ecosystem functions. We show that, due to multifunctional complementarity among species, overall functioning is more susceptible to species loss than are single functions. Modeled relationships between species richness and overall ecosystem functioning using five empirical data sets on monocultures reflected the range of effects of species loss on multiple functions predicted by the model. Furthermore, an exploration of the correlations across functions and the degree of redundancy within functions revealed that multifunctional redundancy was generally lower than single-function redundancy in these empirical data sets. We suggest that by shifting the focus to the variety of functions maintained by a diversity of species, the full importance of biodiversity for the functioning of ecosystems can be uncovered. Our results are thus important for conservation and management of biota and ecosystem services.     

Compositional and functional stability of arthropod communities in the face of ant invasions

There is a general consensus that the diversity of a biotic community can have an influence on its stability, but the strength, ubiquity, and relative importance of this effect is less clear. In the context of biological invasions, diversity has usually been studied in terms of its effect on a community's invasibility, but diversity may also influence stability by affecting the magnitude of compositional or functional changes experienced by a community upon invasion. We examined the impacts of invasive ants on arthropod communities at five natural area sites in the Hawaiian Islands, and assessed whether differences among sites in community diversity and density variables were related to measures of stability. Ant invasion was usually associated with significant changes in overall community composition, as measured by Bray-Curtis distances, particularly among endemic subsets of the communities. Changes in mean species richness were also strong at three of the five sites. Among sites, diversity was negatively related to stability as measured by resistance to overall compositional change, but this effect could not be separated from the strong negative effect of invasive ant density on compositional stability. When compositional stability was measured as proportional change in richness, the best predictor of stability among endemic community subsets was endemic richness, with richer communities losing proportionately more species than species-poor communities. This effect was highly significant even after controlling for differences in invasive ant density and suggested that communities that had already lost many endemic species were resistant to further species loss upon ant invasion, while more intact communities remained vulnerable to species loss. Communities underwent strong but idiosyncratic functional shifts in association with ant invasion, both in terms of trophic structure and total arthropod biomass. There were no apparent relationships, however, between functional stability and community diversity or density measures. Instead, invasive ant density was the best among-site predictor of the magnitude of functional change. Overall, diversity appeared to be a poor predictor of stability in the face of ant invasion in these communities, possibly because any actual diversity effects were overshadowed by community-specific factors and variation in the magnitude of the ant-mediated perturbation.     

Seaweed richness and herbivory increase rate of community recovery from disturbance

The importance of herbivores and of plant diversity for community succession and recovery from disturbance is well documented. However, few studies have assessed the relative magnitude of, or potential interactions between, these factors. To determine the combined effect of herbivory and surrounding algal species richness on the recovery of a rocky intertidal community, we conducted a 27-month field experiment assessing algal recruitment and succession in cleared patches that mimic naturally forming gaps in the ambient community. We crossed two herbivore treatments, ambient and reduced abundance, with monocultures and polycultures of the four most common algal species in a mid-high rocky intertidal zone of northern California. We found that both the presence of herbivores and high surrounding algal richness increased recovery rates, and the effect of algal richness was twice the magnitude of that of herbivores. The increased recovery rate of patches containing herbivores was due to the consumption of fast-growing, early colonist species that preempt space from perennial, late-successional species. Mechanisms linking algal richness and recovery are more numerous. In polycultures, herbivore abundance and species composition is altered, desiccation rates are lower, and propagule recruitment, survival, and growth are higher compared to monocultures, all of which could contribute the observed effect of surrounding species richness. Herbivory and species richness should jointly accelerate recovery wherever palatable species inhibit late-successional, herbivore-resistant species and recruitment and survival of new colonists is promoted by local species richness. These appear to be common features of rocky-shore seaweed, and perhaps other, communities.     

Predator diversity and ecosystem functioning: density modifies the effect of resource partitioning

The link between biodiversity and ecosystem functioning is now well established, but the challenge remains to develop a mechanistic understanding of observed effects. Predator-prey interactions provide an opportunity to examine the role of resource partitioning, thought to be a principal mediator of biodiversity-function relationships. To date, interactions between multiple predators and their prey have typically been investigated in simplified agricultural systems with limited scope for resource partitioning. Thus there remains a dearth of studies examining the functional consequences of predator richness in diverse food webs. Here, we manipulated a species-rich intertidal food web, crossing predator diversity with total predator density, to simultaneously examine the independent and interactive effects of diversity and density on the efficiency of secondary resource capture. The effect of predator diversity was only detectable at high predator densities where competitive interactions between individual predators were magnified; the rate of resource capture within the species mixture more than doubled that of the best-performing single species. Direct observation of species-specific resource use in monoculture, as quantified by patterns of prey consumption, provided clear evidence that species occupied distinct functional niches, suggesting a mechanistic explanation of the observed diversity effect.     

Geographic variation in resource dominance-discovery in Brazilian ant communities

A predictive framework for the ecology of species invasions requires that we learn what limits successful invaders in their native range. The red imported fire ant (Solenopsis invicta) is invasive in the United States, Puerto Rico, Australia, New Zealand, and China. Solenopsis invicta appears to be a superior competitor in its introduced range, where it can cause the local extirpation of native species, but little is known about its competitive ability in its native range in South America. Here we examine the competitive ability of S. invicta for food resources in three widely separated Brazilian ant communities. Each of these communities contains 20-40 ant species, 8-10 of which were common and frequently interacted with S. invicta. S. invicta at all three sites was attacked by several species-specific phorid parasitoids, and at one site, two other species were attacked by their own specialized parasitoids. We examined interactions in these local communities for evidence that trade-offs among ant species between resource dominance and resource discovery, and between resource dominance and parasitoid vulnerability facilitate local coexistence. The trade-off between resource dominance and resource discovery was strong and significant only at Santa Genebra, where parasitoids had no effect on the outcome of confrontations at resources. At Bonito, parasitoids significantly reduced the ability of S. invicta, which was the top-ranked behavioral dominant, from defending and usurping food resources from subordinate species. In the Pantanal, S. invicta ranked behind three other ant species in a linear hierarchy of behavioral dominance, and lost the majority of its interactions with a fourth more subordinate species, Paratrechina fulva, another invasive species. Parasitoids of S. invicta were uncommon in the Pantanal, and did not affect its low position in the hierarchy relative to the other two sites. Parasitoids, however, did affect the ability of Linepithema angulatum, the top-ranked behavioral dominant in this community, from defending and usurping resources from behavioral subordinates. These results indicate that both interspecific competition and trait-mediated indirect effects of phorid parasitoids affect the ecological success of the red imported fire ant in its native range, but that the relative importance of these factors varies geographically.     

The influence of intraguild predation on prey suppression and prey release: a meta-analysis

Intraguild predation (IGP) occurs when one predator species consumes another predator species with whom it also competes for shared prey. One question of interest to ecologists is whether multiple predator species suppress prey populations more than a single predator species, and whether this result varies with the presence of IGP. We conducted a meta-analysis to examine this question, and others, regarding the effects of IGP on prey suppression. When predators can potentially consume one another (mutual IGP), prey suppression is greater in the presence of one predator species than in the presence of multiple predator species; however, this result was not found for assemblages with unidirectional or no IGP. With unidirectional IGP, intermediate predators were generally more effective than the top predator at suppressing the shared prey, in agreement with IGP theory. Adding a top predator to an assemblage generally caused prey to be released from predation, while adding an intermediate predator caused prey populations to be suppressed. However, the effects of adding a top or intermediate predator depended on the effectiveness of these predators when they were alone. Effects of IGP varied across different ecosystems (e.g., lentic, lotic, marine, terrestrial invertebrate, and terrestrial vertebrate), with the strongest patterns being driven by terrestrial invertebrates. Finally, although IGP theory is based on equilibrium conditions, data from short-term experiments can inform us about systems that are dominated by transient dynamics. Moreover, short-term experiments may be connected in some way to equilibrium models if the predator and prey densities used in experiments approximate the equilibrium densities in nature.     

Birds as predators in tropical agroforestry systems

Insectivorous birds reduce arthropod abundances and their damage to plants in some, but not all, studies where predation by birds has been assessed. The variation in bird effects may be due to characteristics such as plant productivity or quality, habitat complexity, and/or species diversity of predator and prey assemblages. Since agroforestry systems vary in such characteristics, these systems provide a good starting point for understanding when and where we can expect predation by birds to be important. We analyze data from bird exclosure studies in forests and agroforestry systems to ask whether birds consistently reduce their arthropod prey base and whether bird predation differs between forests and agroforestry systems. Further, we focus on agroforestry systems to ask whether the magnitude of bird predation (1) differs between canopy trees and understory plants, (2) differs when migratory birds are present or absent, and (3) correlates with bird abundance and diversity. We found that, across all studies, birds reduce all arthropods, herbivores, carnivores, and plant damage. We observed no difference in the magnitude of bird effects between agroforestry systems and forests despite simplified habitat structure and plant diversity in agroforests. Within agroforestry systems, bird reduction of arthropods was greater in the canopy than the crop layer. Top-down effects of bird predation were especially strong during censuses when migratory birds were present in agroforestry systems. Importantly, the diversity of the predator assemblage correlated with the magnitude of predator effects; where the diversity of birds, especially migratory birds, was greater, birds reduced arthropod densities to a greater extent. We outline potential mechanisms for relationships between bird predator, insect prey, and habitat characteristics, and we suggest future studies using tropical agroforests as a model system to further test these areas of ecological theory.     

Rainfall and soils modify plant community response to grazing in Serengeti National Park

Terrestrial plant community responses to herbivory depend on resource availability, but the separate influences of different resources are difficult to study because they often correlate across natural environmental gradients. We studied the effects of excluding ungulate herbivores on plant species richness and composition, as well as available soil nitrogen (N) and phosphorus (P), across eight grassland sites in Serengeti National Park (SNP), Tanzania. These sites varied independently in rainfall and available soil N and P. Excluding herbivores decreased plant species richness at all sites and by an average of 5.4 species across all plots. Although plant species richness was a unimodal function of rainfall in both grazed and ungrazed plots, fences caused a greater decrease in plant species richness at sites of intermediate rainfall compared to sites of high or low rainfall. In terms of the relative or proportional decreases in plant species richness, excluding herbivores caused the strongest relative decreases at lower rainfall and where exclusion of herbivores increased available soil P. Herbivore exclusion increased among-plot heterogeneity in species composition but decreased coexistence of congeneric grasses. Compositional similarity between grazed and ungrazed treatments decreased with increasing rainfall due to greater forb richness in exclosures and greater sedge richness outside exclosures and was not related to effects of excluding herbivores on soil nutrients. Our results show that plant resources, especially water and P, appear to modulate the effects of herbivores on tropical grassland plant diversity and composition. We show that herbivore effects on soil P may be an important and previously unappreciated mechanism by which herbivores influence plant diversity, at least in tropical grasslands.