Optimal diet choice for large herbivores: an extended contingency model

1. A more general contingency model of optimal diet choice is developed, allowing for simultaneous searching and handling, which extends the theory to include grazing and browsing by large herbivores.</p><p>2. Foraging resolves into three modes: purely encounter-limited, purely handling-limited and mixed-process, in which either a handling-limited prey type is added to an encounter-limited diet, or the diet becomes handling-limited as it expands.</p><p>3. The purely encounter-limited diet is, in general, broader than that predicted by the conventional contingency model,</p><p>4. As the degree of simultaneity of searching and handling increases, the optimal diet expands to the point where it is handling-limited, at which point all inferior prey types are rejected,</p><p>5. Inclusion of a less profitable prey species is not necessarily independent of its encounter rate and the zero-one rule does not necessarily hold: some of the less profitable prey may be included in the optimal diet. This gives an optimal foraging explanation for herbivores' mixed diets.</p><p>6. Rules are shown for calculating the boundary between encounter-limited and handling-limited diets and for predicting the proportion of inferior prey to be included in a two-species diet,</p><p>7. The digestive rate model is modified to include simultaneous searching and handling, showing that the more they overlap, the more the predicted diet-breadth is likely to be reduced.</p>

Herbivores strongly influence algal recruitment in both coral- and algal-dominated coral reef habitats

Coral reefs can exist as coral- and macroalgae-dominated habitats often separated by only a few hundred metres. While herbivorous fish are known to depress the abundance of algae and help maintain the function of coral-dominated habitats, less is known about their influence in algae-dominated habitats. Here, we quantified herbivorous fish and benthic algal communities over a 6 mo period in coral-dominated (back-reef) and algal-dominated (lagoon) habitats in a relatively undisturbed fringing coral reef (Ningaloo, Western Australia). Simulta - neously, we tested the effects of herbivorous fish on algal recruitment in both habitats using recruitment tiles and fish exclusion cages. The composition of established algal communities differed consistently between habitats, with the back-reef hosting a more diverse community than the Sargassum-dominated lagoon. However, total algal biomass and cover only differed between habitats in autumn, coinciding with maximum Sargassum biomass. The back-reef hosted high coral cover and a diverse herbivorous fish community, with herbivore biomass an order of magnitude greater than the lagoon. Despite these differences in herbivore composition, exclusion of large herbivores had a similar positive effect to foliose macroalgae recruitment on experimental tiles in both back-reef and lagoon habitats. Additionally, territorial damselfish found in the backreef increased turf algae cover and decreased crustose coralline algae cover on recruitment tiles. Collectively, our results show that disparate herbivorous fish communities in coral- and algaedominated habitats are similarly able to limit the recruitment of foliose macroalgae, but suggest that when herbivorous fish biomass and diversity are relatively low, macroalgal communities are able to escape herbivore control through increased growth.

Defoliation and patchy nutrient return drive grazing effects on plant and soil properties in a dairy cow pasture

Large herbivores can influence plant and soil properties in grassland ecosystems, but especially for belowground biota and processes, the mechanisms that explain these effects are not fully understood. Here, we examine the capability of three grazing mechanisms-plant defoliation, dung and urine return, and physical presence of animals (causing trampling and excreta return in patches)-to explain grazing effects in Phleum pratense-Festuca pratensis dairy cow pasture in Finland. Comparison of control plots and plots grazed by cows showed that grazing maintained original plant-community structure, decreased shoot mass and root N and P concentrations, increased shoot N and P concentrations, and had an inconsistent effect on root mass. Among soil fauna, grazing increased the abundance of fungivorous nematodes and Aporrectodea earthworms and decreased the abundance of detritivorous enchytraeids and Lumbricus earthworms. Grazing also increased soil density and pH but did not affect average soil inorganic-N concentration. To reveal the mechanisms behind these effects, we analyzed results from mowed plots and plots that were both mowed and treated with a dung and urine mixture. This comparison revealed that grazing effects on plant attributes were almost entirely explained by defoliation, with only one partly explained by excreta return. Among belowground attributes, however, the mechanisms were more mixed, with effects explained by defoliation, patchy excreta return, and cow trampling. Average soil inorganic-N concentration was not affected by grazing because it was simultaneously decreased by defoliation and increased by cow presence. Presence of cows created great spatial heterogeneity in soil N availability and abundance of fungivorous nematodes. A greenhouse trial revealed a grazing-induced soil feedback on plant growth, which was explained by patchiness in N availability rather than changes in soil biota. Our results show that grazing effects on plant attributes can be satisfactorily predicted using the effects of defoliation, whereas those on soil fauna and soil N availability need understanding of other mechanisms as well. The results indicate that defoliation-induced changes in plant ecophysiology and the great spatial variation in N availability created by grazers are the two key mechanisms through which large herbivores can control grassland ecosystems.

India's biodiversity hotspot under anthropogenic pressure: A case study of Nilgiri Biosphere Reserve

This paper presents data on the impact of biotic pressure in terms of grazing by livestock and wood cutting by humans on the plant community in the Nilgiri Biosphere Reserve of India. Grass, and herbaceous plant biomass, number of cattle dung piles, number of woody stems available and damaged by human activities and weed biomass were assessed at different proximity along transects radiating from village-forest boundary to forest interior to measure the ecological impact of livestock grazing and fire wood collection. The grass biomass was positively correlated to overgrazing indicating the adverse effect on natural vegetation by cattle. Woodcutting was intense along the forest boundary and significantly declined as distance increased. Similarly, weed biomass and number of thorny species declined positively with proximity from village-forest boundary and the weed biomass was significantly higher in the pastoral sites compared to residential sites. The results suggest that human impact adversely affects natural vegetation and promotes weed proliferation in forest areas adjoining human settlements in the ecologically important Nilgiri Biosphere Reserve. Continued anthropogenic pressure could cause reduction in fodder availability to large herbivores like elephants, which in turn leads to an increase in human-elephant conflict. (C) 2011 Elsevier GmbH. All rights reserved.

The control of indirect effects of biomanipulation

Results from long-term investigations on biomanipulation show that indirect effects are at least as important as direct effects are for the stability of biomanipulation. Three types of indirect effects can be distinguished: (1) a change in quantity or quality of the resource base, (2) behavioural change of the prey, and (3) development of anti-predator traits. Although indirect effects of type (2), (e.g. a change in the pattern of vertical migration of zooplankton), and type (3), (e.g. development of helmets and neck teeth in Daphnia), are important mechanisms, the most essential indirect effects regarding biomanipulation belong to type (1). An example of the latter will be demonstrated: the complex of indirect effects of enhanced grazing by large herbivores on the phosphorus metabolism of the lake. It is concluded that control of the indirect effects is absolutely necessary to stabilize biomanipulation measures, but this is much more difficult than the control of direct effects and needs deeper insights into the structuring mechanisms of food webs. Proper management of fish stocks, in combination with the control of phosphorus load and/or the physical conditions, seems to be the most promising way of controlling the indirect effects of biomanipulation.

Comunidade de mamíferos de médio e grande porte em um mosaico de fitofisionomias na Mata Atlântica no estado do Espírito Santo, Brasil

A Mata Atlântica é considerada como um dos cinco hotspots mais diversos, sendo um conjunto complexo de ecossistemas que abriga uma parcela significativa da diversidade biológica do Brasil. O Estado do Espírito Santo, localizado na região Sudeste, possuía quase 90% de sua superfície coberta por Mata Atlântica, entretanto com o processo de colonização portuguesa, ocupação do território e industrialização, restou apenas 8% da cobertura florestal original. Com toda a informação existente acerca das espécies neotropicais de mamíferos de médio e grande porte ainda há muitas lacunas no nosso conhecimento. Nesse sentido, esta dissertação foi desenvolvida em quatro capítulos com o objetivo de realizar um levantamento de dados de riqueza, composição, abundância, densidade populacional, período de atividade, distribuição e uso do hábitat por espécies de mamíferos de médio e grande porte, da Reserva Natural Vale (RNV), norte do Espírito Santo. Espera-se que as informações aqui geradas possam contribuir com o incremento no conhecimento da ecologia dos mamíferos neotropicais, bem como fornecer informações desse grupo para subsidiar as políticas e ações ambientais que visem à conservação da biodiversidade. Para este estudo foi realizado amostragens mensais no período de abril de 2013 até junho de 2014 na RNV, através de armadilhamento fotográfico (39 armadilhas fotográficas) e também transecções lineares Os registros obtidos nas39 armadilhas fotográficas resultaram em um total de 7.020 dias de monitoramento. Foram observadas 23245 fotos de 26 espécies de mamíferos de médio e grande porte. Uma maior riqueza e frequência de registros ocorreram nas armadilhas estabelecidas na região norte da RNV. A distância do recurso hídrico estava positivamente relacionada com a composição de mamíferos e a quantidade de registros de caça no presente estudo teve um efeito marginalmente negativo na frequência de registros nas regiões oeste e sul, onde apresentaram uma maior incidência de caça. Nossos resultados mostram que essas espécies não estão distribuídas uniformemente dentro da reserva, com a ocupação das mesmas tendo sido afetada por quatro principais covariáveis: (1) distância entre árvores; (2) distância da estrada; (3) distância do recurso hídrico mais próximo e (4) densidade de lianas por hectare. A detectabilidade das espécies que são consideradas cinegéticas foi afetada pelo histórico de registros de caça. A maioria das espécies registradas que tiveram uma probabilidade de ocupação alta são raras ou até mesmo localmente extintas em outras áreas de domínio da Mata Atlântica. Adicionalmente nossos dados confirmam que a RNV abriga uma fauna de mamíferos de médio e grande porte rica, incluindo grandes herbívoros, dispersores de sementes e alguns carnívoros. Embora o estudo tenha sido realizado em apenas uma área, vale lembrar que a RNV, juntamente com a ReBio Sooretama, formam o maior bloco de área protegida dentro do estado do Espírito Santo e também um dos maiores blocos de Mata de Tabuleiro. Assim, para a conservação das espécies que habitam a RNV, é preciso considerar não apenas a reserva isolada, mas todo o bloco Linhares-Sooretama, uma vez que estão conectados permitirão o incremento populacional e as trocas genéticas entre as populações. Somente desse modo será aumentada a probabilidade de sobrevivência e manutenção das espécies em longo prazo, das espécies de mamíferos de médio e grande porte

“Islands” in Holocene forests: implications for forest openness, landscape clearance and “culture-steppe” species

Human activity has undoubtedly had a major impact on Holocene forested ecosystems, with the concurrent expansion of plants and animals associated with cleared landscapes and pasture, also known as 'culture-steppe'. However, this anthropogenic perspective may have underestimated the contribution of autogenic disturbance (e.g. wind-throw, fire), or a mixture of autogenic and anthropogenic processes, within early Holocene forests. Entomologists have long argued that the north European primary forest was probably similar in structure to pasture woodland. This idea has received support from the conservation biologist Frans Vera, who has recently strongly argued that the role of large herbivores in maintaining open forests in the primeval landscapes of Europe has been seriously underestimated. This paper reviews this debate from a fossil invertebrate perspective and looks at several early Holocene insect assemblages. Although wood taxa are indeed important during this period, species typical of open areas and grassland and dung beetles, usually associated with the dung of grazing animals, are persistent presences in many early woodland faunas. We also suggest that fire and other natural disturbance agents appear to have played an important ecological role in some of these forests, maintaining open areas and creating open vegetation islands within these systems. More work, however, is required to ascertain the role of grazing animals, but we conclude that fossil insects have a significant contribution to make to this debate. This evidence has fundamental implications in terms of how the palaeoecological record is interpreted, particularly by environmental archaeologists and palaeoecologists who may be more interested in identifying human-environment interactions rather than the ecological processes which may be preserved within palaeoecological records.

La conservation de la biodiversité dans les aires protégées en zones périurbaines : dynamique des communautés végétales au parc national du Mont-Saint-Bruno entre 1977 et 2013

Ce mémoire visait à déterminer si un petit parc périurbain, en l’occurrence le Parc national du Mont-Saint-Bruno, est parvenu à maintenir l’intégrité de sa flore au cours des trente dernières années en dépit de pressions humaines croissantes et de perturbations naturelles passées, en plus d’identifier les principaux changements floristiques survenus au cours de cette période et les facteurs responsables. Pour répondre à ces objectifs, une étude historique a été réalisée, en comparant un inventaire floristique ancien (1977) à un inventaire récent (2013). Mes résultats montrent d’abord une forte croissance de la diversité alpha indigène au cours des 35 dernières années, accompagnée d’un déclin significatif de la diversité bêta (30%). Malgré cette homogénéisation taxonomique, la diversité fonctionnelle de la flore forestière s’est accrue, la rendant probablement plus résiliente aux événements perturbateurs. D’autre part, mes analyses ont révélé la progression de traits fonctionnels souvent associés à des habitats forestiers intensément broutés, révélant une certaine influence du cerf de Virginie sur la composition et la structure de la flore forestière. Enfin, mes résultats ont montré que les herbiers botaniques se révèlent être une alternative fiable aux méthodes traditionnelles pour documenter et évaluer l’impact des grands herbivores sur la morphologie des plantes broutées. Au final, cette étude a montré que les petites aires protégées périurbaines peuvent jouer un rôle majeur dans la préservation de la diversité floristique d’habitats forestiers d’intérêt, particulièrement lorsque leur statut de protection permet d’encadrer de manière stricte les activités humaines.

Composição química e valor nutricional para grandes herbívoros das folhas e frutos de aninga (Montrichardia linifera, Araceae)

Montrichardia linifera (Araceae), conhecida popularmente como 'aninga', faz parte dos ecossistemas de várzea da Amazônia e da dieta natural de animais como peixe-boi, tartarugas, peixes, búfalo e gado. Com o objetivo de contribuir para o conhecimento químico e valor nutricional da mesma, folhas e frutos de M. linifera foram coletados às margens dos rios Guamá e Maratauíra, no Estado do Pará, Brasil. Em folhas e frutos foram realizadas análises de umidade, resíduo mineral fixo (cinzas), lipídios, proteínas, fibra bruta, concentração de carboidratos e valor calórico. A composição mineral (Ca, Mg, Cu, Fe, Zn e Mn) foi obtida por espectrometria de absorção atômica de chama. Observou-se que tanto as folhas quanto os frutos da aninga, apesar de calóricos (289,75 kcal e 355,12 kcal, respectivamente), possuem baixo valor protéico (0,44% e 0,24 %, respectivamente). As concentrações de manganês obtidas (folha = 3279,46 mg kg^-1e fruto = 18151,53 mg kg^-1) foram consideradas tóxicas, extrapolando o limite máximo tolerável para ruminantes (1000 mg kg^-1). A M. linifera, tem capacidade de absorver e bioacumular grandes quantidades de Ca, Mg e Mn presentes no solo, o que torna inadequada a sua utilização exclusiva na alimentação de quelônios, bovinos e bubalinos, havendo necessidade de mais estudos para sua aplicação como parte da ração.

Caracterização física, química e nutricional dos frutos da Aninga

Montrichardia linifera, conhecida popularmente como aninga, pertence à família Araceae e forma grandes populações às margens dos rios e igarapés da Amazônia, principalmente de águas brancas (barrentas). Seus frutos fazem parte da dieta alimentar de peixes, tartarugas e de grandes herbívoros, como peixe-boi e búfalo, havendo, portanto, a hipótese de seu aproveitamento na nutrição animal. Entretanto, apesar de sua vasta distribuição na paisagem amazônica, não existem dados na literatura sobre a composição nutricional desses frutos. Com os objetivos de avaliar o seu potencial nutricional e contribuir para o conhecimento dessa espécie, foi realizada, neste trabalho, a caracterização física, química e nutricional dos frutos. A infrutescência pesa em média 500 g, com um conjunto de aproximadamente 80 frutos. O seu valor nutritivo se resume, basicamente, ao seu valor energético (≅ 350 kcal), devido principalmente ao teor de carboidratos (≅ 80%), possuindo baixo valor proteico (< 0,5%). As concentrações de manganês (≅ 1.800 mg kg^-1) foram consideradas tóxicas, extrapolando o limite máximo tolerável em nutrição de bubalinos (1.000 mg kg^-1).

Seedling fate across different habitats: the effects of herbivory and soil fertility

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Antler Size of Alaskan Moose Alces alces gigas: Effects of Population Density, Hunter Harvest and Use of Guides

Moose Alces alces gigas in Alaska, USA, exhibit extreme sexual dimorphism, with adult males possessing large, elaborate antlers. Antler size and conformation are influenced by age, nutrition and genetics, and these bony structures serve to establish social rank and affect mating success. Population density, combined with anthropogenic effects such as harvest, is thought to influence antler size. Antler size increased as densities of moose decreased, ostensibly a density-dependent response related to enhanced nutrition at low densities. The vegetation type where moose were harvested also affected antler size, with the largest-antlered males occupying more open habitats. Hunts with guides occurred in areas with low moose density, minimized hunter interference and increased rates of success. Such hunts harvested moose with larger antler spreads than did non-guided hunts. Knowledge and abilities allowed guides to satisfy demands of trophy hunters, who are an integral part of the Alaskan economy. Heavy harvest by humans was also associated with decreased antler size of moose, probably via a downward shift in the age structure of the population resulting in younger males with smaller antlers. Nevertheless, density-dependence was more influential than effects of harvest on age structure in determining antler size of male moose. Indeed, antlers are likely under strong sexual selection, but we demonstrate that resource availability influenced the distribution of these sexually selected characters across the landscape. We argue that understanding population density in relation to carrying capacity (K) and the age structure of males is necessary to interpret potential consequences of harvest on the genetics of moose and other large herbivores. Our results provide researchers and managers with a better understanding of variables that affect the physical condition, antler size, and perhaps the genetic composition of populations, which may be useful in managing and modeling moose populations.

Tintinnids of the western Arabian Sea

The results of an investigation of tintinnids from the western Arabian Sea are described. A total of 134 closing-net samples was obtained from 22 stations of the German "Meteor" expedition 1964/1965. Distribution charts of the dominant species of tintinnids from the study area are presented as well as a list of the world-wide distribution of these species as derived from the literature. Tintinnids were most abundant in the surface waters. The layer from 0 - 25 m yielded a maximum 94.3% and a minimum of 61.3% of the tintinnids present from 0 - 175 m; the mean was 80%. There was no significant difference in the vertical distribution between day and night stations nor was there any indication of the influence of the thermocline upon vertical distribution of tintinnids. TS-diagrams show different water types in the western Arabian Sea. Temperatur-salinity-tintinnid -diagrams indicate regional patterns in the distribution of various species of tintinnids. Some tintinnids can be used as indicator species: Climacocylis scalaria, Parundella lohmanni and Amphorella amphora were typical for the Somali Current whereas Rhabdonella apophysata and Branditella palliata indicated the presence of East African Coastal Current water. The concentration of tintinnids in the upper 25 m raged between 4,800 and 39,300 individuals/m**3 (mean 19,000/m**3). Plasma volume of tintinnids was calculated to permit comparison of different links in the food chain. There was a mean of 51 mm**3/m**2 in the upper layer, equivalent to a concentration of 2 mm**3/m**3. Carbon values were computed from the plasma volume of tintinnids, phytoplankton and larger zooplankton. The ratio of phytoplankton plus microzooplankton carbon to large zooplankton carbon was 1 : 0.8 in the Somali Current, 1 : 0.4 in the East African Coastal Current and 1 : 1.2 in the mixing zone of these current systems. Tintinnids are one of the first links in the food chain. It is very likely that a part of the organic detritus and of the nanoplankton is transfered to large herbivores or omnivores via tintinnids and other protozoans. This mechanism might be especially effective during seasons when large phytoplankters are not available in the ocean.

(Table A1) Annual biomass of plants and animals recorded on Bylot Island between 1993-2009

Determining the manner in which food webs will respond to environmental changes is difficult because the relative importance of top-down vs. bottom-up forces in controlling ecosystems is still debated. This is especially true in the Arctic tundra where, despite relatively simple food webs, it is still unclear which forces dominate in this ecosystem. Our primary goal was to assess the extent to which a tundra food web was dominated by plant-herbivore or predator--rey interactions. Based on a 17-year (1993-2009) study of terrestrial wildlife on Bylot Island, Nunavut, Canada, we developed trophic mass balance models to address this question. Snow Geese were the dominant herbivores in this ecosystem, followed by two sympatric lemming species (brown and collared lemmings). Arctic foxes, weasels, and several species of birds of prey were the dominant predators. Results of our trophic models encompassing 19 functional groups showed that <10% of the annual primary production was consumed by herbivores in most years despite the presence of a large Snow Goose colony, but that 20-100% of the annual herbivore production was consumed by predators. The impact of herbivores on vegetation has also weakened over time, probably due to an increase in primary production. The impact of predators was highest on lemmings, intermediate on passerines, and lowest on geese and shorebirds, but it varied with lemming abundance. Predation of collared lemmings exceeded production in most years and may explain why this species remained at low density. In contrast, the predation rate on brown lemmings varied with prey density and may have contributed to the high-amplitude, periodic fluctuations in the abundance of this species. Our analysis provided little evidence that herbivores are limited by primary production on Bylot Island. In contrast, we measured strong predator-prey interactions, which supports the hypothesis that this food web is primarily controlled by top-down forces. The presence of allochthonous resources subsidizing top predators and the absence of large herbivores may partly explain the predominant role of predation in this low-productivity ecosystem.