Winning by a neck: tall giraffes avoid competing with shorter browsers

With their vertically elongated body form, giraffes generally feed above the level of other browsers within the savanna browsing guild, despite having access to foliage at lower levels. They ingest more leaf mass per bite when foraging high in the tree, perhaps because smaller, more selective browsers deplete shoots at lower levels or because trees differentially allocate resources to promote shoot growth in the upper canopy. We erected exclosures around individual Acacia nigrescens trees in the greater Kruger ecosystem, South Africa. After a complete growing season, we found no differences in leaf biomass per shoot across height zones in excluded trees but significant differences in control trees. We conclude that giraffes preferentially browse at high levels in the canopy to avoid competition with smaller browsers. Our findings are analogous with those from studies of grazing guilds and demonstrate that resource partitioning can be driven by competition when smaller foragers displace larger foragers from shared resources. This provides the first experimental support for the classic evolutionary hypothesis that vertical elongation of the giraffe body is an outcome of competition within the browsing ungulate guild.     

Generalities in grazing and browsing ecology: using across-guild comparisons to control contingencies

In community ecology, broad-scale spatial replication can accommodate contingencies in patterns within species groups, but contingencies in processes across species groups remain problematic. Here, based on a focused review of grazing and browsing by large mammals, we use one trophic guild as a “control” for the other to identify generalities that are not contingent upon specific consumer-resource interactions. An example of such a generality is the Jarman–Bell principle, which explains how allometries of metabolism and digestion influence dietary tolerance and thereby enable resource partitioning within both guilds at multiple scales. By comparing the grazing succession with browsing stratification we show how competition from smaller herbivores, rather than facilitation from larger ones, is the underlying process structuring ungulate assemblages when shared resources become limiting. Also, grazing lawns and browsing hedges are functionally similar. In each case, plants expressing tolerance traits can withstand chronic grazing or browsing in sites where the nutritive value of the local food resource is enhanced in positive feedback to the actions of its consumers. The debate over whether ungulates accelerate or decelerate nutrient cycling can be resolved by comparing grazing and browsing effects in the same ecosystem type. Evidence from African savannas points to the rate of nutrient cycling being controlled by the mix of tolerance and resistance traits in plants; not the relative dominance of grazing or browsing by local herbivores. We recommend this across-guild comparative approach as a novel solution with widespread utility for resolving contingencies in community processes.     

Behaviour of goats,sheep and cattle and their selection of browse species on natural pasture in a Sahelian area

A study was conducted in the Sahelian zone of Burkina Faso of the behaviour of cattle, sheep and goats on natural pasture and their preference for browse species. Concurrently, a survey was undertaken on a sample of farmers (herders, women and livestock owners) in the study area to estimate the indigenous knowledge of browse species and their utilisation by ruminants. A herd of cattle and flocks of sheep and goats belonging to farmers were followed on pasture, each species during three consecutive days each month, from May 2003 to April 2004, and their activities were recorded regularly every 15 min, as well as the browse species selected and the height reached while browsing. The farmers had good knowledge of the browse species present in the area and their preferential classification depended on the availability of the species, their nutritive value and also other ways of utilising the species concerned. However, some divergence existed in the knowledge of farmers compared to the result from the behaviour study, e.g. some species were mentioned by farmers but not found in the inventory. There was a decline in the feeding activities of all animal species from rainy to dry season, while resting and ruminating activities were increasing at the same time. This decline in time spent feeding was more important for cattle (from 72 to 39% of total time) as they relied on the herbaceous biomass for feeding, while sheep and goats made a shift in the feeding activities from grazing to browsing when the herbaceous biomass decreased. Cattle browsed (leaves and litter) during all the study period for around 4.5% of the time spent on pasture. Sheep and goats showed a peak in browsing activity in the dry season, 28 and 52% of the time, respectively. During the whole observation period, cattle browsed 10 species, with Guiera senegalensis most often selected, with 59, 54 and 84% of browsing time, respectively, in the rainy, post rainy and dry season. G. senegalensis, Combretum micranthum and Balanites aegyptiaca were the most important species browsed by sheep among the 20 browse species selected. Goats browsed more than 20 species daily but the most preferred species were Acacia senegal, B. aegyptiaca and Pterocarpus lucens. The mean height reached by goats when browsing was higher (1.65 m) than that of cattle (1.47 m) and sheep (0.87 m). The result from this study can be used to select species for regeneration and to advice farmers on the importance of herd composition.     

Defensive traits of savanna trees – the role of shoot exposure to browsers

To investigate patterns of plant defence, a study was conducted on 13 tree species in Botswana. We tested the hypotheses that (1) shoots on small, young trees are more defended than shoots at the same height on tall, mature individuals and (2) shoots within browsing height are more defended than shoots above the reach of browsers, on mature trees. Plant traits studied were total phenolics, tannin activity, nitrogen (N), acid-detergent fibre (ADF), and degree of spinescence (in three species). We found some support for the first hypothesis. Overall young trees had a significantly higher concentration of total phenolics than mature trees and on species level these differences were significant for three of the 13 species. However, we found no differences in tannin activity, spinescence or ADF concentration between young and mature trees, and four species had higher levels of N in young trees, suggesting higher overall palatability. Contrary to our second hypothesis, shoots within browsing height were less chemically defended (total phenolics and tannin activity) than shoots above browsing height in six of 13 species. Two species had less ADF in lower shoots, and no significant differences were found in N or spinescence for any species. This study included a broad range of tree species at two sites and the results indicate that general theories of plant defence, originating from boreal and northern temperate forests, cannot be simply applied to savanna ecosystems.     

Seasonal grazing effects by semi-domesticated reindeer on subarctic mountain birch forests

In northern Fennoscandia, the spatial and temporal grazing practices of semi-domesticated reindeer (Rangifer tarandus tarandus) vary, which implies different grazing effects dependent on natural conditions as well as management regime (i.e., timing and intensity of grazing). We compared density and biomass of main plant groups in semi-dry mountain birch forests exposed to either long-term summer or winter grazing in three reindeer herding districts in the northernmost Finland. Percent plant cover, height, and biomass of reindeer lichens (Cladonia spp.) and dwarf shrubs (Vaccinium uliginosum, Calluna vulgaris, and Betula nana) were lower on summer ranges compared with winter ranges. The biomass of other dwarf shrubs (Vaccinium myrtillus and V. vitis-idaea), and graminoids and herbs, and the % cover of non-vegetated bare soil and litter were, however, higher on summer ranges than on winter ranges. Young mountain birch shoots (Betula pubescens ssp. czerepanovii) were less frequent on summer ranges than on winter ranges. The total leaf biomass under the browsing height of reindeer (<1.5 m) was also lower on summer ranges compared with winter ranges. Especially in drier and nutrient poor mountain birch forests, intensive summer grazing reduces the quantity of lichens and total plant biomass which affects the ecological state and productivity of these forests and also reduces especially their winter grazing value for reindeer. Therefore, in addition to regulating the maximum sustained numbers of reindeer, pasture rotation systems that effectively protect dry and nutrient poor vegetation from summer grazing and trampling should be encouraged.     

African Elephants Loxodonta africana Amplify Browse Heterogeneity in African Savanna

There is a growing concern that the feeding habits of the African elephant, which include pushing over, uprooting and snapping trees, may have a negative impact on other herbivores. Browsed trees are known to respond by either increasing production (shoots and leaves) or defence (secondary compounds). It is not clear, however, what proportion of the browsed biomass can be made available at lower feeding heights after a tree is pushed over or snapped; thus, it is also unclear how the forage quality is affected. In a field survey in Kruger National Park, South Africa, 708 Mopane trees were measured over four elephant utilization categories: snapped trees, pushed‐over trees, uprooted trees and control trees. The elephants' impact on the leaf biomass distribution was quantified, and the forage quality (Ca, P, K and Mg, N, digestibility and condensed tannin [CT] concentrations) were analyzed. Pushed‐over and uprooted trees had the maximum leaf biomass at lower heights (<1 m), snapped trees at medium heights (1–2 m) and control trees at higher heights (>2 m). In all three utilization categories, the minimum leaf biomass was seven times higher than it was for control trees at a height of below 1 m. Leaf nitrogen content increased in all three categories and was significantly higher in snapped trees. CT concentrations increased slightly in all trees that were utilized by elephants, especially on granitic soils in the dry season. The results provide the insight that elephants facilitate the redistribution and availability of browse and improve the quality, which may positively affect small browsing herbivores.     

Growth,chemical responses and herbivory after simulated leaf browsing in <Emphasis Type="Italic">Combretum apiculatum</Emphasis>

Vegetative and chemical responses to simulated leaf browsing during the growth season, and their subsequent effect on herbivory, were studied on Combretum apiculatum Sonder (Combretaceae) in Botswana. Treatments (50% and 100% leaf and shoot apex removal) were performed just before the shoot growth curve levelled out, and responses recorded 3 months later, just before leaf fall. Compared to controls, defoliation treatments, removing apical dominance, reduced growth in tree height and increased shoot mortality, although the production of lateral shoots increased. At the end of the trial, there was no difference in total length of annual shoots between treatment groups. Significant refoliation occurred only after 100% defoliation. Refoliated leaves were smaller and the 100% defoliated trees had a lower final leaf biomass. Total leaf biomass production was, however, equal for all treatment groups. Refoliated leaves contained higher levels of N, lower levels of acid-detergent fibre (ADF) and total phenolics, and showed a trend towards lower levels of condensed tannins, compared to leaves on control trees. Such chemical changes may be due to either carbon stress or to younger physiological age of new leaves. In spite of the observed potential increase in food quality, we found no evidence of increased levels of insect or ungulate herbivory on refoliated leaves, which, at least for insect herbivory, may be explained by the reduction in temporal availability of leaves. We conclude that the single severe defoliation was not detrimental to C. apiculatum in the short-term, although the resource loss and induced compensatory growth may produce negative effects during subsequent growth seasons.     

Cascading effects of predatory fish exclusion on the detritus-based food web of a lake littoral zone (Lake Vico, central Italy)

An exclosure experiment was carried out in the reed-dominated littoral zone of a volcanic lake (Lake Vico, central Italy) to test whether the impact of predatory fish on benthic invertebrates cascades on fungal colonisation and breakdown of leaf detritus. The abundance, biomass, and Shannon diversity index of the invertebrate assemblage colonising Phragmites australis leaf packs placed inside: (1) full-exclosure cages, (2) cages allowing access only to small-sized fish predators, and (3) cageless controls, were monitored over a 45-day period together with the mass loss and associated fungal biomass of leaf packs. The species composition of the fungal assemblage was further assessed at the end of the manipulation. In general, invertebrate predators did not show any significant response to fish exclusion, either on a trophic guild or on a single taxon level. In contrast, the exclusion of large predatory fish induced a diverse spectrum of changes in the abundance and population size-structure of dominant detritivore taxa, ultimately increasing the biomass and Shannon diversity index of the whole detritivorous guild. These changes corresponded with significant variations in leaf detritus decay rates as well as in the biomass and assemblage structure of associated fungal colonisers. Our experimental findings provide evidence that in Lake Vico effects of fish predators on invertebrate detritivores influence the fungal conditioning and breakdown of the detrital substrate. We conclude that in lacustrine littoral zones predator-driven constraints may structure lower trophic levels of detritus-based food webs and affect the decomposition of leaf detritus originated from the riparian vegetation.     

The canary in the coal mine: Sprouts as a rapid indicator of browse impact in managed forests

Forest managers are frequently confronted with sustaining vegetation diversity and structure in landscapes experiencing high ungulate browsing pressure. Often, managers monitor browse damage and risk to plant communities using vegetation as indicators (i.e., phytoindicators). Although useful, the efficacy of traditional phytoindicators is sometimes hampered by limited distribution and abundance, variable browse susceptibility, and lagged responses. In contrast, sprouts possess traits which make them readily available and attractive to browsers, yet fairly resilient to tissue loss. Here, we experimentally evaluate whether hardwood tree stump sprouts are effective and sensitive phytoindicators of deer browse pressure. We measured sprout abundance and height in fenced and unfenced plots at 17 shelterwood harvested sites scattered across a 6500 km² region where deer densities varied by nearly an order of magnitude. We found browsing did not alter the proportion of stumps sprouting and sprout density; however, browse pressure varied among the four most abundant species. Acer rubrum and Acer saccharum were heavily browsed, although browse pressure on A. saccharum decreased in areas with greater canopy openness. Fagus grandifolia and Prunus serotina were less preferred. Differences in palatability altered size hierarchies. Averaged across all species, browsing reduced sprout height by 39%, relative to protected sprouts. Under ambient browsing, P. serotina was 60–100% taller than other species and significantly taller than A. saccharum and F. grandifolia. However, within fences A. saccharum and A. rubrum doubled in size, relative to browsed individuals, and were as tall as P. serotina. Deer impact on sprout height within unfenced forest stands was negatively correlated with estimated deer densities (R² = 0.46). Thus, we suggest sprout surveys can provide a measure of impact across much larger areas. Our results demonstrate that sprouts, particularly those of Acer species, offer an abundant, easily measured, and reliable indicator of browse pressure. Moreover, browse impacts on sprouts emerged before impacts were detected on seedling abundance, height, or biomass. We argue sprouts can warn of imminent browse risk to seedlings (and perhaps non-woody vegetation) and thereby allow managers to take actions to mitigate or avert losses to the regenerating seedling cohort.     

Correlation of relative muzzle width and relative incisor width with dietary preference in ungulates

Qualitative observations suggest that grazing ungulates have relatively broader muzzles than browsing ones, and that grazers have incisors that are all of a similar size, as opposed to the large central and smaller lateral incisors seen in browsers. These differences may be correlated respectively with the need for grazing ungulates to maintain a large daily intake, or for browsing ungulates to forage selectively in a stand of vegetation. Quantitative examination of relative muzzle width and incisor width ratio in 95 species of living ungulates, correlated with seven different types of dietary preferences, substantiated these observations, although phylogenetic history may exhort a strong influence on morphological proportions. For example, equids have relatively narrower muzzles than grazing ruminants despite their less selective mode of feeding. The narrowest relative muzzle widths are not found in regular browsers, but in high level browsers and in mixed feeders in open habitats. Incisor width ratio can distinguish grazers from browsers, but can not be used to discriminate mixed feeders from other feeding types, and grazers appear to have incisors that are relatively broader overall than those of other dietary types, in correlation with their relatively broader muzzles.     

Physical and chemical response of juvenile Acacia tortilis trees to browsing. Experimental evidence

1. Changes in nutritional value and accessibility of leaves following browsing are important in the dynamics of plant–herbivore interactions because they influence the fitness of the plant attacked and the future utilization of it by the herbivore.
2. Hand pruning of Acacia tortilis , a spinescent tree common in savanna ecosystems of eastern Africa, resulted in higher biomass of spines and new shoots in pruned trees than in unpruned controls.
3. Pruned trees allocated a higher proportion of shoot biomass to spines than unpruned ones, whereas the proportion of leaf biomass in new shoots was slightly reduced. Because increases in spine biomass and density following pruning are coupled with an increase in shoot production, it is concluded that higher production of spines is an inducible response of Acacia tortilis to pruning.
4. No significant changes in the concentration of total phenolics, condensed tannins or leaf nitrogen were induced by pruning.
5. Irrespective of treatment, high foliar concentrations of nitrogen were correlated with an increase in twig production for a given leaf biomass and a reduction in the concentration of secondary substances in leaves. This relation may lead to a conflict between foraging efficiency and nutrition for browsers of A. tortilis.     

Is escaping deer browse just a matter of time in Picea sitchensis? A chemical and dendroecological approach

We combined chemical and dendroecological analyses to understand the mechanisms that are involved in escaping deer browse by young Sitka spruce (Picea sitchensis) exposed to browsing by Sitka black-tailed deer (Odocoileus hemionus sitchensis) on Haida Gwaii (British Columbia, Canada). We compared chemical defences (terpenes), nutritive compounds (nitrogen, non-structural constituents, cellulose, and lignin), as well as age and radial growth of two young spruce categories growing side by side: (1) stunted spruces that were heavily browsed, shorter than the browse line, and (2) escaped spruces that were taller than the browse line but still browsed below the browse line. Escaped and stunted spruces did not differ in terpene concentrations, or in nutritive compound contents, suggesting that they had similar palatability. Escaped spruces were older that stunted spruces. Stunted and escaped trees had similar slow growth when young, suggesting no difference in initial browsing between the two spruce categories. For escaped spruce, there was a dramatic increase in radial growth at about 12-13 years old, suggesting that the apex of the trees had escaped deer browse. Because the two categories of spruces were equally accessible and did not differ in chemical defences or in nutritive compounds, and because escaped spruces were older than stunted trees and had a similar slow radial growth in their first 12-13 years, we conclude that morphological differences between stunted and escaped browsed trees are due to age and that it is only a matter of time before spruce escape deer on Haida Gwaii.     

Physiological acclimation and growth response to partial shading in <Emphasis Type="Italic">Salix matsudana</Emphasis> in the Mu Us Sandland in China

In a controlled experiment, Salix matsudana plants were subjected to uniform nonshading (F-S), partial shading (P-S) and uniform shading (U-S). The shoots of the plants in the F-S and U-S treatments were referred to as H-H and L-L, respectively. The plants in the P-S treatment had two kinds of shoots: (1) shoots under the nonshading treatment that were connected to others under the shading treatment (H-L).(2) Shoots under the shading treatment that were connected to others under the nonshading treatment (L-H). The physiological acclimation and growth response of the species to the partial shading were examined. The partial shading had significant effects on photosynthetic dynamics, transpiration and stomatal conductance, but no effect on instantaneous water use efficiency and maximum quantum yield. Water saturation deficit and coefficient of water loss were significantly smaller in the H-L shoots than in the H-H shoots. Leaf natality, leaf mortality and leaf turnover were greater in the H-L shoots than in the H-H shoots. In contrast, these three parameters were smaller in the L-H shoots than in the L-L shoots. The H-L shoots had significantly larger branching ratio, total branch length and shoot biomass than the H-H shoots. The L-H shoots had smaller branching ratio, total branch length and shoot biomass than the L-L shoots. Total plant biomass in the treatments increased as follows: F-S相似文献   

Shift in Black Rhinoceros Diet in the Presence of Elephant: Evidence for Competition?

In African large herbivore assemblages, megaherbivores dominate the biomass and utilise the greatest share of available resources. Consequently, they are considered a separate trophic guild that structures the food niches of coexisting large herbivores. However, there exists little empirical evidence on how food resources are shared within this guild, and none for direct competition for food between megaherbivores. Using the histological analysis of faeces, we explore this phenomenon for African elephant Loxodonta africana and black rhinoceros Diceros bicornis in the Addo Elephant National Park, South Africa, where the accumulated impacts of elephant have reduced browse availability. Despite being unable to generalise beyond our study sites, our observations support the predictions of competition theory (as opposed to optimality theory) by showing (1) a clear seasonal separation in resource use between these megaherbivores that increased as resource availability declined, and (2) rhinoceros changed their selectivity in the absence of elephant (using an adjacent site) by expanding and shifting their diet along the grass-browse continuum, and in relation to availability. Although black rhinoceros are generally considered strict browsers, the most significant shift in diet occurred as rhinoceros increased their preferences for grasses in the presence of elephant. We speculate that the lack of specialised grazing adaptations may increase foraging costs in rhinoceros, through reduced harvest- and handling-efficiencies of grasses. In the short-term, this may be off-set by an enhanced tolerance for low quality food and by seasonally mobilising fat reserves; however, the long-term fitness consequences require further study. Our data suggest that managing elephant at high densities may compromise the foraging opportunities of coexisting browsers. This may be particularly important in small, fenced areas and overlapping preferred habitats where impacts intensify.     

Physiological basis of spacing effects on tree growth and form in Eucalyptus globulus

We examine the effects of spacing and layout on the growth and form of 3- to 4-year-old Eucalyptus globulus in a farm forestry context. Four planting layouts were chosen. These represented the range commonly in use in farm forestry: block plantings (2Ǹ m), triple rows (2Ǹ m) at 10-m intervals, single rows (2᎒ m) and isolated trees (10᎒ m). The physiological significance of key results is interpreted in terms of changes in the parameters of a simple plantation growth model. Under conditions where levels of direct light are high, for example during summer, block-planted trees intercepted only 38% of the light intercepted by isolated trees. On a stand basis, however, the combination of incident radiation and ground coverage declined with lower stand densities. While stand leaf area index declined from around 6 to 1 with increased spacing, individual tree leaf areas rose from around 50 m² in block plantings to 150 m² in isolated trees. The proportion of above-ground biomass found in stems declined with increasing spacing as the mass in foliage and branches increased. Stems accounted for 65% of above-ground biomass in block-planted trees but only 35% in isolated trees. The contributions of leaves and branches correspondingly rose from 19% to 35% and from 16% to 29%, respectively. Changes in biomass distribution were accompanied by increasing branch number, branch thickness, flatter branch angles and the longer retention of lower branches with greater spacing. These changes have implications for the merchantability of the timber. The efficiency of above-ground radiation conversion was constant at 0.67 g MJ^-1 irrespective of spacing. We estimated that foliar maintenance respiration (R_m) accounted for about 90% of above-ground R_m. On a stand basis R_m costs block plantings 23.90 t DM ha^-1 year^-1 (50% annual above-ground photosynthetic production) compared with 6.22 t DM ha^-1 year^-1 (40% annual above-ground photosynthetic production) in stands of isolated trees.     

Plant structural defences against browsing birds: a legacy of New Zealand's extinct moas

Browsing by large vertebrates has been a major force in the evolution of terrestrial plants but Holocene extinctions of the browsers have left a legacy of broken biotic partnerships. Ratite birds were the largest herbivores in several regions, such as the moas of New Zealand. Many woody plants there have a distinct form of branching, described as "divaricate", with thin, wide angled, branches intertwining to form a tangled canopy. Divaricate branching has been interpreted as a form of protection against climate extremes or as an anachronistic defense against the extinct moas. Here we report the first experimental evidence that many of these plants are defended against extant ratite browsers. In feeding experiments on two tree species with different (heteroblastic) juvenile and adult branch morphology, emus and ostriches obtained adequate feeding rates from adult shoots but sub-maintenance feeding rates from juvenile shoots with the ratite-resistant traits. Divaricate juvenile shoots suffered 30–70% less biomass removal to the birds than adult shoots. Ratites browse by a distinctive clamping and tugging action. Structural defence traits that exploit the limitations of this feeding mode include narrow, strong, elastic branches that resist being torn off, wide branching angle ("divaricate") that makes shoots difficult to swallow, and small, widely spaced leaves. This novel plant architectural defence has developed in at least 20% of the native woody flora of New Zealand, including 10 heteroblastic tree species that exhibit the ratite-resistant strategy until they reach ca 3 metres height. It is also a major axis of variation amongst homoblastic woody shrub species. The defences are useless against mammalian browsers that shear shoots, contributing to marked decreases in the abundances of ratite-resistant species in New Zealand after the introduction of mammals.     

Browse characteristics and impact of browsing on trees and shrubs in African savannas

Abstract. Studies on browse and browsing impact on woody plants in African savannas show that browse can be a significant part of the total food for large herbivores. Utilization of browse is little studied and patterns are therefore difficult to examine. The quality of browse has been described mainly in terms of nutrients, fibre and digestibility. Studies on chemical and physical defences of plants against herbivory can add considerably to the understanding of the browse-browser system. The various ways woody plants respond to browsing are little known. Interactions between browsers and woody plants are dynamic, since responses of woody plants to browsing result in changes in feeding behaviour of the browsing animal, both in savannas and in temperate ecosystems.     

Short-Term Effects of Cattle Browsing on Tree Sapling Growth in Mountain Wooded Pastures

Browsing by livestock has been identified as an important factor preventing tree regeneration in wooded pastures. Two field experiments were performed to investigate the effects of cattle browsing on tree sapling growth in a mountain-wooded pasture. Two size classes (ca. 12–17 cm and 41–59 cm) of each of 4 species (Picea abies, Abies alba, Acer pseudoplatanus and Fagus sylvatica) were exposed to zero, low and high grazing intensities. We measured the proportion of saplings browsed and the effect of browsing on growth. A mowing treatment within the zero grazing intensity treatment showed no significant effect on sapling growth. One percent of the large saplings but 25% of the small saplings escaped browsing. Saplings were better protected when surrounded by taller vegetation. The proportion of saplings browsed was not significantly different among species although evergreen tree saplings lost a larger proportion of biomass than deciduous species when browsed. Under grazing, average current-year shoot production and total aboveground biomass of all species were significantly reduced. Browsing effects tended to be smaller at the lower grazing intensity. Because the deciduous species were least reduced in aboveground growth, especially under the low grazing intensity, we conclude that at least in short-term, deciduous species are less affected by cattle browsing.     

Effects of grass and browse consumption on the winter mass dynamics of elk

Much of the research into herbivore ecology and evolution has focused on patterns and mechanisms of niche partitioning in the diets of specialized grazers and browsers, but the significance of the balance of grazing and browsing within intermediate feeders present unique questions that have received less attention. We explored the nutritional effects of varying the balance of grass and browse in the winter diets of elk (Cervus elaphus). We compiled published data from three similar studies that monitored the mass dynamics of captive elk fed diets of pure grass, pure browse or 1 of 14 mixed diets in winter feeding trials. Elk lost mass (up to 22% of initial body weight) in 29 of 33 feeding trials, similar to wild elk in winter. We used regression models of mass dynamics, considering the linear, quadratic or logarithmic effects of the proportion of the diet that was grass (≈1 − proportion of the diet that was browse) and the additive and interactive effects of nitrogen intake. Diet composition had strong effects on mass dynamics, and all models explained ≥73% of the variation (adjusted r ²) in mass dynamics. Nitrogen intake had uniformly positive effects on mass balance, and increasing grass intake caused improved maintenance of body mass up to a point, but further increases in the proportion of grass in the diet had neutral or negative effects on body mass. Overall, the data suggest that elk are adapted to consuming mixed diets. Nonetheless, data on the foraging behavior and diet selection of wild elk in many populations show that elk often consume grass-dominated diets and sometimes consume browse-dominated diets, but rarely consume mixed diets. Physiological adaptations to mixed diets may place unique spatio-temporal constraints on diet selection in intermediate feeders and impose large penalties for a sub-optimal balance of grazing and browsing.     

Balanced dispersal or source–sink–do both models describe wood mice in farmed landscapes?

Plant responses to browsing can affect root and shoot morphology, which is important to subsequent herbivory, nutrient acquisition and competition. This paper examines the above- and below-ground responses of three browse species, with different growth strategies, to simulated browsing damage at different times of year. Saplings were grown in pots in sand culture to enable whole sapling analysis. At winter dormancy or budburst, 50% of previous year's shoots (and associated leaves/buds) were clipped. Subsequent sapling growth and morphology was compared with that of unclipped control saplings. Treatment differences in growth parameters of each species were observed, including changes in branching patterns, shoot lengths, diameters and ratios, leaf sizes and end-of-season bud numbers. Some responses were damage-induced per se; others differed according to timing of damage. Compensatory growth by the two deciduous species ( Betula pendula , Sorbus aucuparia ) resulted in few biomass differences by the end of the year of damage as compared to controls, but both above- and below-ground growth of clipped Pinus sylvestris was poor, particularly those damaged at budburst, giving strong differences between control, dormant, and budburst clipped saplings.