Herbivore Assemblages Research Articles

Negative effects of cattle on soil carbon and nutrient pools reversed by megaherbivores

Wild herbivore populations are declining in many African savannas, which is related to replacement by livestock (mainly cattle) and the loss of megaherbivores. Although some livestock management practices may be compatible with the conservation of native savanna biodiversity, the sustainability of these integrated wild herbivore/livestock management practices is unknown. For instance, how will these herbivore mixes influence key processes for the long-term functioning of savanna ecosystems, such as soil carbon, nitrogen and phosphorus pools and cycling? The Kenya Long-term Exclosure Experiment studies the ecosystem consequences of manipulating the presence and absence of wild herbivores and cattle at moderate densities in a ‘black cotton’ savanna. Here we show that after 20 years, cattle presence decreased total soil carbon and nitrogen pools, while the presence of megaherbivores (mainly elephants) increased these pools and even reversed the negative effects of cattle. Our results suggest that a mix of cattle at moderate densities and wild herbivores can be sustainable, provided that the assemblage of wild herbivores includes the largest species. Cattle are replacing wildlife in many African savannas. This field study finds that wild megaherbivores, such as elephants, increased soil carbon and nitrogen, and hence soil fertility, normally lost when only cattle are present.

Quantifying water requirements of African ungulates through a combination of functional traits

AbstractClimate and land use change modify surface water availability in African savannas. Surface water is a key resource for both wildlife and livestock and its spatial and temporal distribution is important for understanding the composition of large herbivore assemblages in savannas. Yet, the extent to which ungulate species differ in their water requirements remains poorly quantified. Here, we infer the water requirements of 48 African ungulates by combining six different functional traits related to physiological adaptations to reduce water loss, namely minimum dung moisture, relative dung pellet size, relative surface area of the distal colon, urine osmolality, relative medullary thickness, and evaporation rate. In addition, we investigated how these differences in water requirements relate to differences in dietary water intake. We observed strong correlations between traits related to water loss through dung, urine and evaporation, suggesting that ungulates minimize water loss through multiple pathways simultaneously, which suggests that each trait can thus be used independently to predict water requirements. Furthermore, we found that browsers and grazers had similar water requirements, but browsers are expected to be less dependent on surface water because they acquire more water through their diet. We conclude that these key functional traits are a useful way to determine differences in water requirements and an important tool for predicting changes in herbivore community assembly resulting from changes in surface water availability.

Savannah trees buffer herbaceous plant biomass against wild and domestic herbivores

AbstractQuestionsGiven the growing abundance and dominance of domestic herbivores in savannah ecosystems, can trees maintain plant herbaceous standing biomass under increasing herbivore pressure? Are there differences in the capacity of leguminous and non‐leguminous trees in sustaining understory herbaceous biomass? And finally, to what extent does plant community composition underneath trees modulate the effects of herbivore assemblage and abundance on herbaceous biomass?LocationPasturelands and protected areas along the borders of the Serengeti National Park, Tanzania, East Africa.MethodMonthly herbaceous biomass was monitored non‐destructively using a calibrated pasture disc. Measurements were taken within a network of paired exclosures and open plots, underneath large leguminous and non‐leguminous tree canopies and outside canopies. Herbivore community assemblage compositions and abundances were characterised using monthly dung counts, and herbaceous plant community composition was surveyed annually every wet season over two years.ResultsOverall, we found that trees promote herbaceous standing biomass, particularly in the presence of moderate herbivory rather than under herbivore exclusion. Greater herbivore abundance and livestock dominance reduced herbaceous plant biomass, but trees, particularly leguminous trees, limited these negative effects. This capacity for trees to limit the effect of herbivores was related to herbaceous plant species composition. Understory plant communities that were compositionally typical of protected areas sustained the highest plant biomass when found in pasturelands with high herbivore pressure.ConclusionOur findings give greater credence to the importance of preserving large trees in savannah landscapes increasingly dominated by high abundances of livestock. Moreover, our results highlight that park managers and pastoralists need to maintain the specialist herbaceous understory community beneath trees in order to benefit from facilitative tree–understory interactions.

Generalists are more specialized in low-resource habitats, increasing stability of ecological network structure

Linking mechanistic processes to the stability of ecological networks is a key frontier in ecology. In trophic networks, "modules"-groups of species that interact more with each other than with other members of the community-confer stability, mitigating effects of species loss or perturbation. Modularity, in turn, is shaped by the interplay between species' diet breadth traits and environmental influences, which together dictate interaction structure. Despite the importance of network modularity, variation in this emergent property is poorly understood in complex natural systems. Using two years of field data, we quantified interactions between a rich community of lepidopteran herbivores and their host plants across a mosaic of low-resource serpentine and high-resource nonserpentine soils. We used literature and our own observations to categorize herbivore species as generalists (feeding on more than one plant family) or specialists (feeding on one plant family). In both years, the plant-herbivore network was more modular on serpentine than on nonserpentine soils-despite large differences in herbivore assemblage size across years. This structural outcome was primarily driven by reduction in the breadth of host plant use by generalist species, rather than by changes in the composition of species with different fundamental diet breadths. Greater modularity-and thus greater stability-reflects environmental conditions and plastic responses by generalist herbivores to low host plant quality. By considering the dual roles of species traits and ecological processes, we provide a deeper mechanistic understanding of network modularity, and suggest a role for resource availability in shaping network persistence.

The landscape-scale drivers of herbivore assemblage distribution on the central basalt plains of Kruger National Park

AbstractThe distribution and abundance of herbivores in African savannas are constrained by interactions between abiotic and biotic factors. At the species-level, herbivores face trade-offs among foraging requirements, vegetation structure and the availability of surface water that change over spatial and temporal scales. Characterizing herbivore requirements is necessary for the management of the environment in which they occur, as conservation management interventions such as fencing and artificial water provision consequently have effects on how herbivores address these trade-offs. We tested the effects of environmental attributes on the probability of presence of herbivore functional types at different distances to water in the Satara section of Kruger National Park over the period of a year. Hypotheses about species’ relative distribution and abundance were developed through a literature review of forage and water availability constraints on feeding preference and body size of herbivore. We expected strong seasonal relationships between vegetation biomass and quality, and biomass of water-dependent herbivores with increasing distance to water. Our analyses of herbivore distribution across the region confirmed broad-scale descriptions of interactions between forage requirements and water availability across a set of species which differ in functional traits.

Large herbivore assemblages in a changing climate: incorporating water dependence and thermoregulation.

The coexistence of different species of large herbivores (ungulates) in grasslands and savannas has fascinated ecologists for decades. However, changes in climate, land‐use and trophic structure of ecosystems increasingly jeopardise the persistence of such diverse assemblages. Body size has been used successfully to explain ungulate niche differentiation with regard to food requirements and predation sensitivity. But this single trait axis insufficiently captures interspecific differences in water requirements and thermoregulatory capacity and thus sensitivity to climate change. Here, we develop a two‐dimensional trait space of body size and minimum dung moisture content that characterises the combined food and water requirements of large herbivores. From this, we predict that increased spatial homogeneity in water availability in drylands reduces the number of ungulate species that will coexist. But we also predict that extreme droughts will cause the larger, water‐dependent grazers as wildebeest, zebra and buffalo–dominant species in savanna ecosystems – to be replaced by smaller, less water‐dependent species. Subsequently, we explore how other constraints such as predation risk and thermoregulation are connected to this two‐dimensional framework. Our novel framework integrates multiple simultaneous stressors for herbivores and yields an extensive set of testable hypotheses about the expected changes in large herbivore community composition following climate change.

The impact of herbivore exclusion on forb diversity: Comparing species and functional responses during a drought

AbstractRainfall and herbivory shape savannah herbaceous communities, but these disturbances are being altered globally. To assess potential consequences of such alterations, we evaluated herbivore effects on species and functional diversity during an episodic drought in a sodic savannah using data collected from long‐term herbivore exclosures in the Kruger National Park, South Africa. Herbaceous life forms are rarely acknowledged as distinct functional entities. Moreover, the functional ecology of forbs remains elusive. Here, we present disturbances–responses by forbs separately from grasses. We hypothesised that combinations of intense utilisation and drought would be associated with low diversity and high dominance at species and functional levels for both life forms. Contrary to our hypothesis, low forb and grass diversity was associated with long‐term herbivore exclusion, which exceeded expected undesirable effects of intense utilisation and drought. Grasses responded less sensitively, suggesting that forbs respond dynamically to changes in herbivore assemblage when these alterations are combined with drought. Consistent with patterns in savannah systems, forbs contributed significantly to species and functional trait diversity. High forb diversity is suggested to enhance resilience of this nutrient‐rich ecosystem against declines in its functioning when subjected to drought and alterations in herbivory.

Taxonomic and functional diversity of insect herbivore assemblages associated with the canopy-dominant trees of the Azorean native forest.

Oceanic islands have been providing important insights on the structuring of ecological communities and, under the context of the present biodiversity crisis, they are paramount to assess the effects of biological invasions on community assembly. In this study we compare the taxonomic and functional diversity of insect herbivore assemblages associated with the dominant tree species of Azorean native forests and investigate the ecological processes that may have originated current patterns of plant-herbivore associations. Five dominant trees—Erica azorica, Ilex perado subsp. azorica, Juniperus brevifolia, Laurus azorica and Vaccinium cylindraceum—were sampled in the remnants of the native forest of Terceira Island (Azores) using a standardised methodology. The taxonomic and functional diversity of insect herbivore assemblages was assessed using complementary metrics and beta diversity partitioning analysis (species replacement and richness differences) aiming to evaluate the variation in insect herbivore assemblages within and between the study plant species. Sixty two insect species, mostly bugs (Hemiptera) and caterpillars (Lepidoptera), were found in the five study plants with indigenous (endemic and native non-endemic) insects occurring with higher species richness and abundance than introduced ones. Species replacement was the most important component of insect herbivore taxonomic beta diversity while differences in trait richness played a major role on functional beta diversity. The endemic E. azorica stands out from the other study plants by having associated a very distinct insect herbivore assemblage with a particular set of functional attributes, mainly composed by large bodied and long shaped species that feed by chewing. Despite the progressive biotic homogenization witnessed in the Azores during the last few decades, several strong associations between the endemic trees and their indigenous insect herbivores remain.

Resource availability drives trait composition of butterfly assemblages.

How species respond to environmental change is a fundamental question in ecology and species traits can help to tackle this question. In this study, we analyze how the functional structure of species assemblages changes with selected environmental variables along an elevational gradient. In particular, we used species traits of local butterfly communities (body size, voltinism, overwintering stages, and host specificity) in a national nature reserve in China to assess the impacts of temperature, net primary productivity, and land use. Our results show that productivity, measured as NDVI, had a stronger influence on the functional community structure of butterflies than temperature. Within the butterfly assemblages, net primary productivity mainly affected body size and supported few but large species. Length of vegetation period demonstrated dominating effects on the functional structure of local butterfly assemblages. However, an observed increase in dietary generalists with longer vegetation periods contradicted expectations based on niche breadth hypothesis, that more stable conditions should favor specialists. Furthermore, the general positive impact of vegetation period on species abundances differed considerably among functional groups. Only the group containing species hibernating as egg decreased with the length of vegetation period. Our results suggest that trait associations are instructive to explain environment-herbivore relationships, that resource availability can predominantly influence the functional composition of herbivore assemblages, and that conservation priority should be given to specialist butterfly species overwintering as egg, especially in the face of global warming.

Trophic interactions and abiotic factors drive functional and phylogenetic structure of vertebrate herbivore communities across the Arctic tundra biome

Communities are assembled from species that evolve or colonise a given geographic region, and persist in the face of abiotic conditions and interactions with other species. The evolutionary and colonisation histories of communities are characterised by phylogenetic diversity, while functional diversity is indicative of abiotic and biotic conditions. The relationship between functional and phylogenetic diversity infers whether species functional traits are divergent (differing between related species) or convergent (similar among distantly related species). Biotic interactions and abiotic conditions are known to influence macroecological patterns in species richness, but how functional and phylogenetic diversity of guilds vary with biotic factors, and the relative importance of biotic drivers in relation to geographic and abiotic drivers is unknown. In this study, we test whether geographic, abiotic or biotic factors drive biome‐scale spatial patterns of functional and phylogenetic diversity and functional convergence in vertebrate herbivores across the Arctic tundra biome. We found that functional and phylogenetic diversity both peaked in the western North American Arctic, and that spatial patterns in both were best predicted by trophic interactions, namely vegetation productivity and predator diversity, as well as climatic severity. Our results show that both bottom–up and top–down trophic interactions, as well as winter temperatures, drive the functional and phylogenetic structure of Arctic vertebrate herbivore assemblages. This has implications for changing Arctic ecosystems; under future warming and northward movement of predators potential increases in phylogenetic and functional diversity in vertebrate herbivores may occur. Our study thus demonstrates that trophic interactions can determine large‐scale functional and phylogenetic diversity just as strongly as abiotic conditions.

Vertebrate herbivory on shrub seedlings in California sage scrub: important but understudied interactions

Herbivory is poorly characterized in the endangered California sage scrub ecosystem, because previous studies were conducted only in winter and spring. To better understand vertebrate herbivore impacts, we first examined if and when vertebrate herbivores influence Artemisia californica seedling survivorship by examining differences in mortality between caged and uncaged seedlings. Then, we examined: (1) vertebrate herbivore preference of common sage scrub shrub species, and (2) which vertebrate species constitute important herbivores in the ecosystem. We found that mortality for uncaged A. californica seedlings was twice that of caged seedlings, and occurred in the summer and early fall, when previous studies in the system were not conducted. Short leaved shrubs, Artemisia californica and Eriogonum fasciculatum, were more susceptible to herbivory than broader leaved Salvia apiana and Eriodictyon trichocalyx. However, susceptibility likely depends on the herbivore assemblage, as the three main herbivore species [Sylvilagus audubonii (desert cottontail), Zonotrichia leucophrys (white-crowned sparrow; a bird, not a mammal), and Otospermophilus beecheyi (California gray squirrel)] all had different shrub preferences. Our results contrast with previous studies that suggests that native vertebrate herbivores facilitate shrub seedling survivorship by highlighting that late-season vertebrate herbivory may negatively affect shrub seedling survivorship. We demonstrate the complex but understudied role herbivores play in the sage scrub ecosystem, and highlight that further study of these key ecological interactions is warranted.

War-induced collapse and asymmetric recovery of large-mammal populations in Gorongosa National Park, Mozambique.

How do large-mammal communities reassemble after being pushed to the brink of extinction? Few data are available to answer this question, as it is rarely possible to document both the decline and recovery of wildlife populations. Here we present the first in-depth quantitative account of war-induced collapse and postwar recovery in a diverse assemblage of large herbivores. In Mozambique’s Gorongosa National Park, we assembled data from 15 aerial wildlife counts conducted before (1968–1972) and after (1994–2018) the Mozambican Civil War (1977–1992). Pre-war total biomass density exceeded 9,000 kg km-2, but populations declined by >90% during the war. Since 1994, total biomass has substantially recovered, but species composition has shifted dramatically. Formerly dominant large herbivores—including elephant (Loxodonta africana), hippo (Hippopotamus amphibius), buffalo (Syncerus caffer), zebra (Equus quagga), and wildebeest (Connochaetes taurinus)—are now outnumbered by waterbuck (Kobus ellipsiprymnus) and other small to mid-sized antelopes. Waterbuck abundance has increased by an order of magnitude, with >55,000 individuals accounting for >74% of large-herbivore biomass in 2018. By contrast, elephant, hippo, and buffalo, which totaled 89% of pre-war biomass, now comprise just 23%. These trends mostly reflect natural population growth following the resumption of protection under the Gorongosa Restoration Project; reintroductions (465 animals of 7 species) accounted for a comparatively small fraction of the total numerical increase. Waterbuck are growing logistically, apparently as-yet unchecked by interspecific competition or predation (apex-carnivore abundance has been low throughout the post-war interval), suggesting a community still in flux. Most other herbivore populations have increased post-war, albeit at differing rates. Armed conflict remains a poorly understood driver of ecological change; our results demonstrate the potential for rapid post-war recovery of large-herbivore biomass, given sound protected-area management, but also suggest that restoration of community structure takes longer and may require active intervention.

Marine protected areas enhance coral reef functioning by promoting fish biodiversity

AbstractPreserving biodiversity and ecosystem function in the Anthropocene is one of humanity's greatest challenges. Ecosystem‐based management and area closures are considered an effective way to maintain ecological processes, especially in marine systems. Although there is strong evidence that such measures positively affect community structure, their impact on the rate of key ecological processes remains unclear. Here, we provide evidence that marine protected areas enhance herbivory rates on coral reefs via direct and indirect pathways. Using meta‐analysis and a path‐analytical framework, we demonstrate that, on average, protected areas increase the species richness of herbivorous fishes, which, in turn, enhances browsing rates on macroalgae. However, in all three regions studied (the Atlantic, Indian, and Pacific Ocean), a small subset of the herbivore assemblage accounted for the majority of browsing. Our results therefore indicate that ecosystem functioning on coral reefs may respond positively to both area closures and the protection of key species.

Depauperation and divergence of plant‐specialist herbivore assemblages in a fragmented tropical landscape

1. Our understanding of the structure and spatial organisation of biological assemblages in human‐modified tropical landscapes has critical importance to improve conservation actions. Investigations on this topic have focused on local (α) diversity patterns, overlooking the changes in species turnover (β diversity) between sites, and its consequences on total (γ) diversity.2. This study assessed the differences in α, β and γ diversities of galling insects and their host plants (saplings) in a fragmented Atlantic forest landscape in northeast Brazil. Both assemblages were recorded in 30 plots (total of 0.1 ha for each forest type) located in the interior and on the edges of a large fragment and small forest fragments (10 plots per forest type).3. α diversity of host plants and galling insect assemblages was significantly higher in interior (reference) plots than in edge and fragment plots. Yet, both assemblages showed higher β diversity in fragment and edge plots than in reference plots – a finding potentially associated with the hyperdynamism of fragmented forests and consistent with the landscape divergence hypothesis.4. However, biotic differentiation of host plant and galling insects was not great enough to compensate the loss of α diversity, and thus γ diversity, because most host plant and galling insect species in forest fragments were also registered in reference plots. Our findings indicate that, despite each small forest fragment being very dissimilar from each other, they have low importance for the conservation of plant assemblages and their specialized herbivores at landscape scale.

Seascapes as drivers of herbivore assemblages in coral reef ecosystems

AbstractNotionally herbivorous fishes maintains a critical ecosystem function on coral reefs by grazing algae and maintaining highly productive algal turf assemblages. Current paradigms implicate habitat complexity, predation, and primary productivity as major drivers of the distribution and abundance of herbivorous fish, yet little is known about the relative contribution of these factors. Here, we compare bottom‐up and top‐down drivers of notional herbivore assemblages across an environmental gradient of wave exposure in the Palau archipelago. We surveyed herbivore assemblages at reef slopes (6–9 m) across 18 sites, and quantified proxies of top‐down control (predator biomass, habitat complexity) and bottom‐up drivers (net primary production, nutrients) at each site. Despite substantial variability in herbivore biomass throughout the archipelago (6–65 g/m2), general additive models indicate that neither top‐down nor bottom‐up drivers significantly predicted biomass or density of herbivores among sites. In contrast to expectations, herbivore biomass was highest at sites with high predator biomass, low structural complexity, and low benthic productivity. Rather, the highest biomass of herbivores was associated with shallow, tidally emergent, productive reef flats located adjacent to steep vertical walls (“drop‐offs”). The emergent nature of this neighboring habitat precluded occupation by territorial fishes and multiple species of herbivores were observed to make foraging runs into this habitat once tidally inundated. We hypothesize that this habitat configuration provides an important cross‐habitat resource subsidy. Multivariate ordination and permutation of herbivore communities revealed strong evidence for biogeographic partitioning throughout the archipelago (western, southwestern, inner eastern, and outer eastern clusters), contributing to an emerging picture that the habitat heterogeneity of seascapes can overwhelm the effects of conventional top‐down and bottom‐up structuring of herbivory on coral reefs.

High specialization and limited structural change in plant‐herbivore networks along a successional chronosequence in tropical montane forest

Secondary succession is well‐understood, to the point of being predictable for plant communities, but the successional changes in plant‐herbivore interactions remains poorly explored. This is particularly true for tropical forests despite the increasing importance of early successional stages in tropical landscapes. Deriving expectations from successional theory, we examine properties of plant‐herbivore interaction networks while accounting for host phylogenetic structure along a succession chronosequence in montane rainforest in Papua New Guinea. We present one of the most comprehensive successional investigations of interaction networks, equating to > 40 person years of field sampling, and one of the few focused on montane tropical forests. We use a series of nine 0.2 ha forest plots across young secondary, mature secondary and primary montane forest, sampled almost completely for woody plants and larval leaf chewers (Lepidoptera) using forest felling. These networks comprised of 12 357 plant‐herbivore interactions and were analysed using quantitative network metrics, a phylogenetically controlled host‐use index and a qualitative network beta diversity measure. Network structural changes were low and specialisation metrics surprisingly similar throughout succession, despite high network beta diversity. Herbivore abundance was greatest in the earliest stages, and hosts here had more species‐rich herbivore assemblages, presumably reflecting higher palatability due to lower defensive investment. All herbivore communities were highly specialised, using a phylogenetically narrow set of hosts, while host phylogenetic diversity itself decreased throughout the chronosequence. Relatively high phylogenetic diversity, and thus high diversity of plant defenses, in early succession forest may result in herbivores feeding on fewer hosts than expected. Successional theory, derived primarily from temperate systems, is limited in predicting tropical host‐herbivore interactions. All succession stages harbour diverse and unique interaction networks, which together with largely similar network structures and consistent host use patterns, suggests general rules of assembly may apply to these systems.

Biodiversity of Kelp Forests and Coralline Algae Habitats in Southwestern Greenland

All marine communities in Greenland are experiencing rapid environmental change, and to understand the effects on those structured by seaweeds, baseline records are vital. The kelp and coralline algae habitats along Greenland’s coastlines are rarely studied, and we fill this knowledge gap for the area around Nuuk, west Greenland. Using subtidal swath surveys, photo-quadrats, and grab samples, we characterised the diversity of floral and faunal assemblages in kelp forests and coralline algae beds. The most abundant herbivore assemblages and the most diverse communities occur in the interstitial habitats of rhodolith beds. In kelp forests, species diversity is higher in epi-benthic (photo-quadrat) and mid-water (swath) surveys. These habitats are not mutually exclusive; Agarum clathratum is prominent in coralline algal habitats, while crustose coralline algae cover the bedrock under kelp holdfasts. Overall, the suite of surveys used capture the diverse communities within kelp forests and coralline algae in Greenland and their differing role in the life history of the inhabitants. Furthermore, coralline algae beds are an important carbonate store, with CaCO3 concentrations ranging from 28.06 to 103.73 g·m−3. Our research sets the baseline for continued investigations and monitoring of these important habitats and their supported fisheries.

Effects of large herbivores on tundra vegetation in a changing climate, and implications for rewilding.

In contrast to that of the Pleistocene epoch, between approximately 2.6 million and 10 000 years before present, the extant community of large herbivores in Arctic tundra is species-poor predominantly due to human extinctions. We here discuss how this species-poor herbivore guild influences tundra ecosystems, especially in relation to the rapidly changing climate. We show that present herbivore assemblages have large effects on tundra ecosystem composition and function and suggest that the effect on thermophilic species expected to invade the tundra in a warmer climate is especially strong, and that herbivores slow ecosystem responses to climate change. We focus on the ability of herbivores to drive transitions between different vegetation states. One such transition is between tundra and forest. A second vegetation transition discussed is between grasslands and moss- and shrub-dominated tundra. Contemporary studies show that herbivores can drive such state shifts and that a more diverse herbivore assemblage would have even higher potential to do so. We conclude that even though many large herbivores, and especially the megaherbivores, are extinct, there is a potential to reintroduce large herbivores in many arctic locations, and that doing so would potentially reduce some of the unwanted effects of a warmer climate.This article is part of the theme issue 'Trophic rewilding: consequences for ecosystems under global change'.

Predicting rates of consumer-mediated nutrient cycling by a diverse herbivore assemblage

Herbivores mediate the abundances of primary producers both from the top-down, by consuming them, and from the bottom-up, by recycling nutrients. Whereas the top-down effects of herbivores on algae in marine ecosystems are well-documented, less is known about their roles as mediators of local-scale nutrient availability. We conducted a series of surveys and measurements of tide pools and the grazers in those pools between October of 2016 and June of 2017 at an intertidal site on the coast of Southern California, USA (33°35′16.3″N, 117°52′1.5″W). We surveyed grazer abundances in the field, measured biomass of representatives from four different grazer groups (littorine snails, limpets, chitons, and turban snails), measured ammonium excretion rates, and quantified ammonium accumulation rates in tide pools at our study site. We found that different grazer groups were characterized by different per-biomass ammonium excretion rates. Some grazer groups—turban snails and chitons—contributed more ammonium than predicted by their biomass, whereas other grazer groups—littorine snails and limpets—contributed less ammonium than predicted by biomass. Because of these differences between grazer groups, ammonium accumulation rates in tide pools at our study site were effectively predicted based on the ammonium excretion rates of the different grazer groups. However, ammonium accumulation rates were not related to total herbivore biomass. Our results highlight the importance of grazer identity—and particularly the role of species such as turban snails that contribute disproportionately to nutrient recycling—in understanding the contributions of grazers as mediators of bottom-up processes in marine systems.

Tracking of Host Defenses and Phylogeny During the Radiation of Neotropical Inga-Feeding Sawflies (Hymenoptera; Argidae).

Coevolutionary theory has long predicted that the arms race between plants and herbivores is a major driver of host selection and diversification. At a local scale, plant defenses contribute significantly to the structure of herbivore assemblages and the high alpha diversity of plants in tropical rain forests. However, the general importance of plant defenses in host associations and divergence at regional scales remains unclear. Here, we examine the role of plant defensive traits and phylogeny in the evolution of host range and species divergence in leaf-feeding sawflies of the family Argidae associated with Neotropical trees in the genus Inga throughout the Amazon, the Guiana Shield and Panama. Our analyses show that the phylogenies of both the sawfly herbivores and their Inga hosts are congruent, and that sawflies radiated at approximately the same time, or more recently than their Inga hosts. Analyses controlling for phylogenetic effects show that the evolution of host use in the sawflies associated with Inga is better correlated with Inga chemistry than with Inga phylogeny, suggesting a pattern of delayed host tracking closely tied to host chemistry. Finally, phylogenetic analyses show that sister species of Inga-sawflies are dispersed across the Neotropics, suggesting a role for allopatric divergence and vicariance in Inga diversification. These results are consistent with the idea that host defensive traits play a key role not only in structuring the herbivore assemblages at a single site, but also in the processes shaping host association and species divergence at a regional scale.