Herbivory and Body Size: Allometries of Diet Quality and Gastrointestinal Physiology,and Implications for Herbivore Ecology and Dinosaur Gigantism

Digestive physiology has played a prominent role in explanations for terrestrial herbivore body size evolution and size-driven diversification and niche differentiation. This is based on the association of increasing body mass (BM) with diets of lower quality, and with putative mechanisms by which a higher BM could translate into a higher digestive efficiency. Such concepts, however, often do not match empirical data. Here, we review concepts and data on terrestrial herbivore BM, diet quality, digestive physiology and metabolism, and in doing so give examples for problems in using allometric analyses and extrapolations. A digestive advantage of larger BM is not corroborated by conceptual or empirical approaches. We suggest that explanatory models should shift from physiological to ecological scenarios based on the association of forage quality and biomass availability, and the association between BM and feeding selectivity. These associations mostly (but not exclusively) allow large herbivores to use low quality forage only, whereas they allow small herbivores the use of any forage they can physically manage. Examples of small herbivores able to subsist on lower quality diets are rare but exist. We speculate that this could be explained by evolutionary adaptations to the ecological opportunity of selective feeding in smaller animals, rather than by a physiologic or metabolic necessity linked to BM. For gigantic herbivores such as sauropod dinosaurs, other factors than digestive physiology appear more promising candidates to explain evolutionary drives towards extreme BM.     

The importance of aquatic vegetation in beaver diets and the seasonal and habitat specificity of aquatic‐terrestrial ecosystem linkages in a subarctic environment

The extent and ecological significance of trophic linkages across ecosystem boundaries have been the subject of considerable recent research attention. North American beavers Castor canadensis engineer terrestrial influences in aquatic ecosystems by constructing terrestrial food caches near their lodges and aquatic influences in terrestrial ecosystems by building dams and flooding low lying areas. However, it is poorly resolved to what extent beavers rely on aquatic food sources and whether this reliance is greater during winter when ice cover physically confines beavers to aquatic habitats or during summer when warm, ice free water promotes the growth and accessibility of aquatic vegetation. Working in a subarctic region, we surveyed the abundance of aquatic and terrestrial food sources in and around lotic and lentic environments and estimated their contributions to beaver diets during open water and ice covered periods using carbon and nitrogen stable isotope analysis of hair samples. Ponds had four times more aquatic vegetation than streams, but terrestrial habitats around ponds had less than half as much shrub cover as habitats adjacent to streams. Beaver diets in this subarctic environment are estimated to be comprised of 60 to 80% aquatic vegetation, with beavers occupying ponds consuming more aquatic vegetation in winter than beavers occupying streams, which rely more on terrestrial shrubs cached near their lodge. Collectively, these results show how the influence of physical barriers on ecosystem linkages can be modified by habitat‐ and season‐specific abundances of preferred resources and the potential for animals to consume food in ecosystems and seasons different from where and when the food was harvested.     

Experimental evidence that the ecosystem effects of aquatic herbivory by moose and beaver may be contingent on water body type

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Influence of Summer and Autumn Nutrition on Body Condition and Reproduction in Lactating Mule Deer

ABSTRACT Recent work suggests that availability and quality of forage in late summer and early autumn, a time when female ungulates face multiple energetic demands, is critical to reproduction in wild ungulates. Therefore, we examined direct links between nutritional quality of diets, body condition, and reproduction of lactating mule deer. Using captive mule deer, we tested the hypothesis that females consuming diets with lower digestible energy (DE; kJ/g) would have lower DE intake rates (DEI; MJ/day), have less body fat and muscle, have later estrus cycles, and have lower pregnancy and twinning rates. Deer fed lower DE diets had lower DEI during summer and autumn. In turn, deer with lower DEI, regardless of diet DE, had lower body mass, body fat, and muscle thickness. When nutritional quality of diets began to decline earlier in the summer, relationships between food quality, DEI, and body condition were stronger. Although DEI did not influence estrus date for deer that became pregnant before 21 December, deer with lower DEI had a lower probability of becoming pregnant and had a lower probability of producing twins. Measures of body condition in October (i.e., body mass, body fat, and muscle depth) predicted pregnancy and twinning rates in mule deer. Serum concentration of hormones leptin and Insulin Growth Factor 1 were not good predictors of body condition or reproduction. These findings suggest that managers concerned with productivity of mule deer populations should consider focusing on assessing and improving quality of forage available in summer and autumn.     

Diet composition of the African manatee: Spatial and temporal variation within the Sanaga River Watershed,Cameroon

The present study aimed to investigate the diet of African manatees in Cameroon to better inform conservation decisions within protected areas. A large knowledge gap on diet and seasonal changes in forage availability limits the ability to develop informed local management plans for the African manatee in much of its range. This research took place in the Sanaga River Watershed, which includes two protected areas in the Littoral Region of Cameroon: the Douala‐Edea National Park and the Lake Ossa Wildlife Reserve. We analyzed 113 manatee fecal samples and surveyed shoreline emergent and submerged vegetation within the Sanaga River Watershed. We used microhistological analyses to determine the relative contribution of each plant species to African manatee diets and compared across locations and across seasons (wet vs. dry season). We found that the shoreline vegetation is diverse with over 160 plant species, unevenly distributed across space and season, and dominated by emergent vegetation mostly represented by the antelope grass (Echinochloa pyramidalis). We recorded a total of 36 plant species from fecal samples with a spatial and temporal distribution mostly reflecting that of the corresponding shoreline vegetation. African manatees appear to be primarily opportunistically feeding on available vegetation across the seasons and habitat. This work documents the current, but changing, state of plant availability in the Sanaga River Watershed and reports the African manatee diet in Cameroon for the first time. This information can play a critical role in successfully managing the species and these protected areas. If we wish to protect the African manatee and the aquatic ecosystems within the Sanaga River Watershed, we must understand how forage availability changes over time, especially as its waters become nutrient enriched, eutrophic, and exposed to invasive species of plants in a changing world.     

Selective foraging on woody plant species by the Eurasian beaver (Castor fiber) in Telemark, Norway

Beavers Castor spp. are generalist herbivores, feeding on the bark, shoots and leaves of woody plants, terrestrial herbs and forbs, ferns and aquatic vegetation. As central-place foragers, beavers move out from water to select and cut trees and vegetation, and then transport it back to their refuge. These terrestrial forays are energetically costly; therefore, beavers should concentrate their foraging activity near the central place and increase the degree of selectivity for specific sizes or species of food with increasing distance from the water. The aim of this study was to test the predictions of the central place and the optimal foraging theories for the food selection of the Eurasian beaver Castor fiber , and show the foraging preferences of the focal species in the boreal conifer forest zone of Europe. Foraging intensity by beavers and the abundance of woody species were surveyed in transects positioned randomly at seven beaver territories. In compliance with the central-place foraging theory, the foraging intensity declined with increasing distance from the river. Beavers fed preferentially on willows ( Salix ), rowan ( Sorbus ) and birches ( Betula ), although alders ( Alnus ) dominated their diet. Size selectivity showed similar patterns to previous North American studies, which were also carried out in habitats with predominantly small saplings. The probability of selection of small saplings dropped as distance increased, which is consistent with the predictions of optimal foraging models that larger prey items are more likely to be favoured with increasing provisioning distance.     

Contrasting timing of parturition of chital Axis axis and gaur Bos gaurus in tropical South India – the role of body mass and seasonal forage quality

Seasonal variation in forage availability and quality is understood to affect the annual timing of parturition in large herbivores. In India–where seasonal monsoonal rains define variation in forage availability and quality–chital Axis axis exhibit stronger seasonality in parturition than the larger gaur Bos gaurus. We hypothesized that this difference can be explained by forage requirements determined by body mass. We developed a model to simulate changes in leaf biomass and nitrogen content based on plant available moisture and nutrients, and calibrated our model with field data. Our results show that the minimum forage nitrogen content required by lactating gaur was available throughout the year, but that required by lactating chital was available for less than 40% of the year, i.e. during the early wet season, which coincides with the annual peak period of chital births. The three to four month spread of chital births, which begins in the dry season, implies that the period of highest quality is also important for females to replenish maternal reserves for future reproduction and help maximize the growth rate of neonates. This spread also indicates low synchrony of chital births and suggests that predator swamping was less important in influencing their timing of parturition. As monsoonal rain exhibits annual temporal variation, we analyzed our model under different rainfall patterns while keeping the total annual rainfall constant. We found that the difference between the durations of how long forage quality is available to satisfy the minimum requirements of lactating gaur and lactating chital is similar for all simulated patterns. This insensitivity to variable rainfall patterns lends support to our hypothesis that forage requirements determined by body mass is one plausible explanation for the variation in parturition strategies among large herbivores species.     

Herbivory and growth in terrestrial and aquatic populations of amphibious stream plants

1. Many amphibious plant species grow in the transition between terrestrial and submerged vegetation in small lowland streams. We determined biomass development, leaf turnover rate and invertebrate herbivory during summer in terrestrial and aquatic populations of three amphibious species to evaluate advantages and disadvantages of aerial and submerged life.
2. Terrestrial populations had higher area shoot density, biomass and leaf production than aquatic populations, while leaf turnover rate and longevity were the same. Terrestrial populations experienced lower percentage grazing loss of leaf production (average 1.2–5.1%) than aquatic populations (2.9–17.3%), while the same plant dry mass was consumed per unit ground area.
3. Grazing loss increased linearly with leaf age apart from the youngest leaf stages. Grazing loss during the lifetime of leaves was therefore 2.4–3.1 times higher than mean apparent loss to standing leaves of all ages. The results imply that variation in density of grazers relative to plant production can account for differences in grazing impact between terrestrial and aquatic populations, and that fast leaf turnover keeps apparent grazing damage down.
4. We conclude that the ability of amphibious plants to grow submerged permits them to expand their niche and escape intense competition on land, but the stream does not provide a refugium against grazing and constrains plant production compared with the terrestrial habitat.     

Foraging behaviour of coypus Myocastor coypus: why do coypus consume aquatic plants?

Foraging behaviour of wild coypu was studied to examine two hypotheses that had been previously proposed to explain the species’ preference for aquatic plants. First, the nutritional benefit hypothesis which states that aquatic plants are more nutritional than terrestrial plants. Second, the behavioural trade-off hypothesis which states that coypus avoid foraging far from the water because of the costs associated with other types of behaviour. In order to test the nutritional benefit hypothesis, we studied the diet composition of coypus in relation to the protein content of the diet and of the plants available in the environment. Fieldwork was conducted seasonally from November 1999 to August 2000 at one study site located in the Province of Buenos Aires, east central Argentina. Behavioural observations showed that coypus remained foraging in the water and microhistological analysis of faeces indicated that their diet was principally composed of hygrophilic monocotyledons (Lemna spp. and Eleocharis spp.) throughout the year. We did not find support for the nutritional benefit hypothesis: nutritional quality (based on nitrogen content) of hygrophilic plants was not higher than that of terrestrial plants, and seasonal changes in diet quality did not match either fluctuations in vegetation quality or proportion of hygrophilic plants in the diet. Although not directly tested, the behavioural trade-off hypothesis may explain why coypus prefer to forage in or near the water as a mechanism for reducing predation risk.     

Herbivores Enforce Sharp Boundaries Between Terrestrial and Aquatic Ecosystems

The transitions between ecosystems (ecotones) are often biodiversity hotspots, but we know little about the forces that shape them. Today, often sharp boundaries with low diversity are found between terrestrial and aquatic ecosystems. This has been attributed to environmental factors that hamper succession. However, ecosystem properties are often controlled by both bottom-up and top-down forces, but their relative importance in shaping riparian boundaries is not known. We hypothesize that (1) herbivores may enforce sharp transitions between terrestrial and aquatic ecosystems by inhibiting emergent vegetation expansion and reducing the width of the transition zone and (2) the vegetation expansion, diversity, and species turnover are related to abiotic factors in the absence of herbivores, but not in their presence. We tested these hypotheses in 50 paired grazed and ungrazed plots spread over ten wetlands, during two years. Excluding grazers increased vegetation expansion, cover, biomass, and species richness. In ungrazed plots, vegetation cover was negatively related to water depth, whereas plant species richness was negatively related to the vegetation N:P ratio. The presence of (mainly aquatic) herbivores overruled the effect of water depth on vegetation cover increase but did not interact with vegetation N:P ratio. Increased local extinction in the presence of herbivores explained the negative effect of herbivores on species richness, as local colonization rates were unaffected by grazing. We conclude that (aquatic) herbivores can strongly inhibit expansion of the riparian vegetation and reduce vegetation diversity over a range of environmental conditions. Consequently, herbivores enforce sharp boundaries between terrestrial and aquatic ecosystems.     

西太湖水生植物时空变化

水生植物在浅水湖泊生态系统中具有十分重要的作用。根据中国科学院太湖湖泊生态系统研究站1989年以来的常规监测资料,将西太湖（除东太湖以外的湖区）划分为9个区,采用点截法（point intercept method）,于2002～2005年对各区水生植物的种类、生物量和空间分布情况进行了6次调查。结果表明：西太湖现有水生植物16种,分属于11科12属;水生植物总面积约10220hm^2,其中沉水植物分布面积约占64．58％;挺水植物约占0．29％;漂浮植物约占38．16％。各个种之间生物量差异显著,马来眼子菜、荇菜、芦苇的生物量在所有水生植物中居前3位。多样性分析表明,水生植物种类4a来未发生明显变化,但种类和生物量季节性差异较大。水生植物呈环状分布在距湖岸5km以内的水域和部分岛屿周围,东岸和南岸为水生植物的主要集中分布区域,分布区连续性好,且水草种类齐全。挺水植物种类单一,仅有芦苇（Phragmites communis）一种,分布区域多限于水深小于1．6m的湖岸;沉水植物共有8种,为水生植物的主要组成部分,马来眼子菜（Potamogeton malaianus）的分布频度最高,在西山岛周围水域逐年扩张,成为该区域的先锋种;漂浮植物3种,主要以荇菜（Nymphoides peltata）为主,在七都水域有逐渐扩张的趋势。马来眼子菜、芦苇、荇菜表现出对水环境较强的适应能力,目前为西太湖的3个优势种。20世纪50年代以来,西太湖水生植物种类减少了50种,其中水质下降是导致水生植物种类不断减少甚至消失的一个重要原因。围网养殖和不合理的捕捞方式也对局部水域的植物造成极大的破坏。水生植物生存环境日益严峻,种群单一化趋势日益明显。     

洱海漂浮草垫的物种组成及分布

研究通过调查洱海2017年夏季和冬季漂浮草垫的物种组成及分布模式, 探究洱海在水位上升过程中, 哪些挺水植物更易形成漂浮草垫而存活下来。夏季共调查到漂浮草垫18个, 植物分属15科, 18属, 24种, 冬季共调查到漂浮草垫22个, 植物分属13科, 17属, 22种, 形成洱海漂浮草垫的植物共15科, 19属, 26个物种。草垫下水体中总氮(TN)、总磷(TP)、溶解性总磷(TDP)都明显高于开阔水域的(P<0.05), 而开阔水域的溶氧(DO)明显高于草垫下水体的(P<0.05)。结果表明在夏季和冬季漂浮草垫面积大小与其组成草垫的物种数和漂浮草垫生物量都呈显著正相关(P<0.01), 而在夏季和冬季漂浮草垫面积大小与漂浮草垫单位面积平均生物量相关性都不显著(P>0.05), 漂浮草垫面积大小与漂浮草垫下最长根在夏季相关性显著(P<0.01), 而在冬季相关性不显著(P>0.05)。漂浮草垫主要分布在离岸距离60 m以内以及水深不超过2 m的水域, 漂浮草垫面积大部分都小于600 m2(夏季占87%, 冬季占95%), 每个漂浮草垫的物种数都小于10个, 漂浮草垫下最长根主要分布在40—120 cm。组成漂浮草垫的物种中菰(Zizania latifolia)的频度和相对生物量在夏季和冬季(频度: 夏季-73.33%, 冬季-66.67%; 生物量: 夏季-43.38%, 冬季-41.91%)都是最高的, 说明菰相比其他挺水植物更容易形成漂浮草垫从而避免洱海水位上升对其生长和繁殖的影响, 这可能是菰成为洱海单优挺水植物的主要原因。而菰比其他挺水植物更容易形成漂浮草垫的机制则需要更进一步的研究。     

Beaver herbivory on aquatic plants

Herbivores have strong impacts on marine and terrestrial plant communities, but their impact is less well studied in benthic freshwater systems. For example, North American beavers (Castor canadensis) eat both woody and non-woody plants and focus almost exclusively on the latter in summer months, yet their impacts on non-woody plants are generally attributed to ecosystem engineering rather than herbivory. Here, we excluded beavers from areas of two beaver wetlands for over 2 years and demonstrated that beaver herbivory reduced aquatic plant biomass by 60%, plant litter by 75%, and dramatically shifted plant species composition. The perennial forb lizard’s tail (Saururus cernuus) comprised less than 5% of plant biomass in areas open to beaver grazing but greater than 50% of plant biomass in beaver exclusions. This shift was likely due to direct herbivory, as beavers preferentially consumed lizard’s tail over other plants in a field feeding assay. Beaver herbivory also reduced the abundance of the invasive aquatic plant Myriophyllum aquaticum by nearly 90%, consistent with recent evidence that native generalist herbivores provide biotic resistance against exotic plant invasions. Beaver herbivory also had indirect effects on plant interactions in this community. The palatable plant lizard’s tail was 3 times more frequent and 10 times more abundant inside woolgrass (Scirpus cyperinus) tussocks than in spatially paired locations lacking tussocks. When the protective foliage of the woolgrass was removed without exclusion cages, beavers consumed nearly half of the lizard’s tail leaves within 2 weeks. In contrast, leaf abundance increased by 73–93% in the treatments retaining woolgrass or protected by a cage. Thus, woolgrass tussocks were as effective as cages at excluding beaver foraging and provided lizard’s tail plants an associational refuge from beaver herbivory. These results suggest that beaver herbivory has strong direct and indirect impacts on populations and communities of herbaceous aquatic plants and extends the consequences of beaver activities beyond ecosystem engineering.     

Dietary phosphorus affects the growth of larval Manduca sexta

Although phosphorus has long been considered an important factor in the growth of diverse biota such as bacteria, algae, and zooplankton, insect nutrition has classically focused on dietary protein and energy content. However, research in elemental stoichiometry has suggested that primary producer biomass has similar N:P ratios in aquatic and terrestrial systems, and phosphorus-rich herbivores in freshwater systems frequently face phosphorus-limited nutritional conditions. Therefore, herbivorous insects should also be prone to phosphorus limitation. We tested this prediction by rearing Manduca sexta larvae on artificial and natural (Datura wrightii leaves) diets containing varying levels of phosphorus (approximately 0.20, 0.55, or 1.2% phosphorus by dry weight). For both artificial and natural diets, increased dietary phosphorus significantly increased growth rates and body phosphorus contents, and shortened the time to the final instar molt. Caterpillars did not consistently exhibit compensatory feeding for phosphorus on either type of diet. The growth and body phosphorus responses were not explicable by changes in amounts of potassium or calcium, which co-varied with phosphorus in the diets. Concentrations of phosphorus in D. wrightii leaves collected in the field varied over a range in which leaf phosphorus is predicted to affect M. sexta's growth rates. These results suggest that natural variation in dietary phosphorus is likely to affect the growth rate and population dynamics of M. sexta, and perhaps larval insects more generally.     

Beaver activity increases aquatic subsidies to terrestrial consumers

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Climate change modifies the size structure of assemblages of emerging aquatic insects

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Climate change and the future distributions of aquatic macrophytes across boreal catchments

Aim Aquatic–terrestrial ecotones are vulnerable to climate change, and degradation of the emergent aquatic macrophyte zone would have severe ecological consequences for freshwater, wetland and terrestrial ecosystems. Our aim was to uncover future changes in boreal emergent aquatic macrophyte zones by modelling the occurrence and percentage cover of emergent aquatic vegetation under different climate scenarios in Finland by the 2050s. Location Finland, northern Europe. Methods Data derived from different GIS sources were used to estimate future emergent aquatic macrophyte distributions in all catchments in Finland (848 in total). We used generalized additive models (GAM) with a full stepwise selection algorithm and Akaike information criterion to explore the main environmental determinates (climate and geomorphology) of emergent aquatic macrophyte distributions, which were derived from the national subclass of CORINE land‐cover classification. The accuracy of the distribution models (GAMs) was cross‐validated, using percentage of explained deviance and the area under the curve derived from the receiver‐operating characteristic plots. Results Our results indicated that emergent aquatic macrophytes will expand their distributions northwards from the current catchments and percentage cover will increase in all of the catchments in all climate scenarios. Growing degree‐days was the primary determinant affecting distributions of emergent aquatic macrophytes. Inclusion of geomorphological variables clearly improved model performance in both model exercises compared with pure climate variables. Main conclusions Emergent aquatic macrophyte distributions will expand due to climate change. Many emergent aquatic plant species have already expanded their distributions during the past decades, and this process will continue in the years 2051–80. Emergent aquatic macrophytes pose an increasing overgrowth risk for sensitive macrophyte species in boreal freshwater ecosystems, which should be acknowledged in management and conservation actions. We conclude that predictions based on GIS data can provide useful ‘first‐filter’ estimates of changes in aquatic–terrestrial ecotones.     

Importance of terrestrial subsidies for native brook trout in Appalachian intermittent streams

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Local Control of Aquatic Resources: Community and Ecology in Lake Titicaca, Peru

All 151 fishing communities in Lake Titicaca, Peru, maintain and defend communally controlled fishing territories. Environmental factors, particularly the slope of the lake bottom and the presence and abundance of aquatic vegetation, influence the distribution of the three types of such territories, which differ in the area they cover and in the maximum depth of water they contain. A cost-benefit model L employed to explain this spatial patterning. This study emphasizes the interactions between aquatic and terrestrial resources. It discusses the conflicts between the formal legal codes of the Peruvian state and the informal regulations of peasant communities. It argues for a refining of the terminology used to describe and analyze common-property resources.