Iain J. Gordon

The allometry of food intake in grazing ruminants

(1) A simulation model of grazing mechanics in ruminants shows that, due to the allometric relations of bite size and metabolic requirements to body size, small animals are able to subsist on shorter swards than large animals. (2) The density of nutrients in the grazed horizon of the modelled swards markedly affected the ability of animals of a given body size to satisfy their energy requirements. (3) By extension, the allometric relationships would be expected to apply in selective grazing and browsing species in their choice of food items of different size and nutrient content. (4) The results support the argument that sexual segregation and habitat choice of dimorphic species is an effect of scramble competition for limited resources, the males thus being excluded from mutually preferred swards. (5) The model provides an explanation for two interspecific phenomena amongst grazers: grazing succession and grazing facilitation.

Distribution and abundance of small insects and arachnids in relation to structural heterogeneity of grazed, indigenous grasslands

1. The species composition and spatial distribution of small insects (Hemiptera, Coleoptera, Lepidoptera) and arachnids (Araneae, Opiliones, and Pseudoscorpiones) were investigated in three indigenous, upland grasslands identified as the National Vegetation Classification Festuca–Agrostis–Galium typical subcommunity (code U4a), Festuca–Agrostis–Galium, Vaccinium–Deschampsia subcommunity (code U4e), and Nardus stricta species‐poor sub‐community (code U5a), on which grazing management was manipulated experimentally.

Incisor arcade structure and diet selection in ruminants

The structure of the incisor arcade in ruminant ungulates is likely to affect the degree to which animals can be selective in choosing food items. Species of ruminant which feed predominantly on grasses have broader more flattened incisor arcades when compared at the same size with species which browse on woody dicotyledonous plants. This is likely to reflect the differential dispersion of food items on the food plants of these two different groups of ruminants.

Prediction of intake and digestion in ruminants by a model of rumen kinetics integrating animal size and plant characteristics

Simulation modelling was used to investigate interactions between forage degradation characteristics, rumen processes and body weight, and to predict the voluntary food intake and digestion of a range of forages. Predicted voluntary intake and digestion agreed well with empirical data, explaining 61 and 70%, respectively, of variance in observed values. Since the data covered a wide range of animal weights and forage qualities, these results suggest that the model is a useful means of integrating the effects of animal and forage variables. Interactions were examined between animal weight and diet quality, as defined by the proportion of potentially digestible cell contents and cell walls and their rates of digestion. Retention time of food in the digestive tract was shown by regression to scale with W⁰·²⁷. The time taken to comminute large fibre particles also scaled with W⁰·²⁷. Longer retention of digesta by large ruminants increases digestive efficiency compared with small animals and would permit them to survive on lower-quality foods. The model showed that maximum intake of metabolizable energy scales with c. W⁰·⁸⁷, greater than the scaling of maintenance with W⁰·⁷³

Landscape features affect gene flow of Scottish Highland red deer (Cervus elaphus)

Landscape features have been shown to strongly influence dispersal and, consequently, the genetic population structure of organisms. Studies quantifying the effect of landscape features on gene flow of large mammals with high dispersal capabilities are rare and have mainly been focused at large geographical scales. In this study, we assessed the influence of several natural and human‐made landscape features on red deer gene flow in the Scottish Highlands by analysing 695 individuals for 21 microsatellite markers. Despite the relatively small scale of the study area (115 × 87 km), significant population structure was found using F‐statistics (FST = 0.019) and the program structure, with major differentiation found between populations sampled on either side of the main geographical barrier (the Great Glen). To assess the effect of landscape features on red deer population structure, the ArcMap GIS was used to create cost‐distance matrices for moving between populations, using a range of cost values for each of the landscape features under consideration. Landscape features were shown to significantly affect red deer gene flow as they explained a greater proportion of the genetic variation than the geographical distance between populations. Sea lochs were found to be the most important red deer gene flow barriers in our study area, followed by mountain slopes, roads and forests. Inland lochs and rivers were identified as landscape features that might facilitate gene flow of red deer. Additionally, we explored the effect of choosing arbitrary cell cost values to construct least cost‐distance matrices and described a method for improving the selection of cell cost values for a particular landscape feature.

Comparative nutrient extraction from forages by grazing bovids and equids: a test of the nutritional model of equid/bovid competition and coexistence

SummaryRuminants are unevenly distributed across the range of body sizes observed in herbivorous mammals; among extant East African species they predominate, in numbers and species richness, in the medium body sizes (10–600 kg). The small and the large species are all hind-gut fermenters. Some medium-sized hind-gut fermenters, equid perissodactyls, coexist with the grazing ruminants, principally bovid artiodactyls, in grassland ecosystems. These patterns have been explained by two complementary models based on differences between the digestive physiology of ruminants and hind-gut fermenters. The Demment and Van Soest (1985) model accounts for the absence of ruminants among the small and large species, while the Bell/Janis/Foose model accounts both for the predominance of ruminants, and their co-existence with equids among the medium-sized species (Bell 1971; Janis 1976; Foose 1982). The latter model assumes that the rumen is competitively superior to the hind-gut system on medium quality forages, and that hind-gut fermenters persist because of their ability to eat more, and thus to extract more nutrients per day from high fibre, low quality forages. Data presented here demonstrate that compared to similarly sized grazing ruminants (bovids), hind-gut fermenters (equids) have higher rates of food intake which more than compensate for their lesser ability to digest plant material. As a consequence equids extract more nutrients per day than bovids not only from low quality foods, but from the whole range of forages eaten by animals of this size. Neither of the current nutritional models, nor refinements of them satisfactorily explain the preponderance of the bovids among medium-sized ungulates; alternative hypotheses are presented.

Megaherbivores influence trophic guilds structure in African ungulate communities

We used a data set of ungulate censuses from 31 natural ecosystems from East and Southern Africa to test two hypotheses: (1) megaherbivores should dominate ungulate communities in ecosystems with high rainfall and low soil nutrient status because of their ability to survive on poor quality food resources, and (2) the abundance of megaherbivores affects the abundance of the mesoherbivores, distinguishing the different feeding guilds: mesograzers, mesobrowsers and mesomixed feeders. Two axes of a multivariate analysis (77% of the variance) discriminated the sites well, the first separating sites dominated by megaherbivores from those dominated by mesoherbivores, and the second representing a gradient between mesograzers and mesobrowsers. Our analysis shows (1) that megaherbivores can be considered to be a separate trophic guild and (2) that mesograzers and mesobrowsers respond differently to variation in their trophic environments. The metabolic biomass density of megaherbivores increased with annual rainfall, but was not related to soil nutrient status, and as predicted, megaherbivores comprised a larger proportion of the biomass of ungulate communities in ecosystems with high rainfall and low nutrient soils. The metabolic biomass density of mesoherbivores increased with rainfall and soil nutrient status. Within the mesoherbivores, the metabolic biomass density of mesograzers showed the same trend, and seemed unaffected by megaherbivores. Conversely, mesobrowsers and mesomixed feeders appeared to be unaffected by rainfall or soil nutrient status, but mesomixed feeders declined when megaherbivores were abundant. This suggests that megaherbivores may compete with the mesomixed-feeder species for food or they may alter the vegetation communities unfavourably. A similar analysis using elephants alone instead of megaherbivores as a group showed that both mesobrowsers and mesomixed feeders were affected significantly by elephant, which is consistent with the fact that most of the effect of megaherbivores on browse resources or woodland habitat is due to elephants. This study shows that the different trophic guilds within African ungulate communities react differently to environmental factors (rain and soil), and that megaherbivores, and particularly elephants, appear to compete with mesomixed feeders and mesobrowsers. These results are relevant for the understanding of the functioning of African ungulate communities and call for further testing with longitudinal data.

The functional significance of the browser-grazer dichotomy in African ruminants

The allometric relationships for the fermentation rate of dry matter, the total energy concentration of volatile fatty acids (VFAs), the energy supplied from VFA production and the mass of the digesta contents within the rumen or caecum and proximal colon (hindgut) were used to test whether the digestive strategies of grazing and browsing African ruminants differ. The wet and dry mass of the contents of the rumen and hindgut were allometrically related to body mass (BM). These relationships did not differ between browsing and grazing ruminants. The fermentation rates in the rumen were strongly allometric and the intercepts of the relationships did not differ between browsers and grazers. The fermentation rates in the hindgut were not allometrically related to BM and did not differ between ruminants with different feeding habits. Likewise, the total energy concentration of the VFAs in the rumen and hindgut showed no allometric scaling and did not differ between browsing and grazing ruminants. The energy supplied by VFA production in both the rumen and hindgut of African ruminants scaled at around 0.8 with BM. Only in the case of the energy supplied by VFAs in the rumen were there significantly different intercepts for browsing and grazing ruminants. The energy supplied by VFA production in the rumen was inadequate to meet the energy requirements for maintenance of browsers and small grazers. The retention time of digesta in the alimentary tract was positively related to BM although there was no difference in the allometric relationships for grazers and browsers. The results of these analyses suggest that, after controlling for the effects of body mass, there is little difference in digestive strategy between African ruminants with different morphological adaptations of the gut.

The ecology of browsing and grazing

Globally, many terrestrial ecosystems have been and are being heavily influenced by human activity, both directly and indirectly. Humanity and our domestic animals (1.4 billion cattle, 1.2 billion sheep and 0.5 billion goats, but only some 120 million horses and 13 million camels; Encyclopedia.com) have now so much impact on global ecosystems that we have entered the Anthropocene (Lewis and Maslin 2015). Wild ruminants number at least 75 million (Hackmann and Spain 2010), and are native to all continents except Antarctica. In such ecosystems extensive grazing and browsing by domestic and wild large mammalian herbivores (hereafter called large herbivores) and, in places, burning have shaped vegetation composition, structure and dynamics. Through their grazing, browsing, trampling and defecation large herbivores not only shape the structure and distribution of the vegetation but also affect nutrient flows and the responses of associated fauna. Consequently, it is the interactions between management or population dynamics of large herbivores and the vegetation they consume that shape the biodiversity, structure and dynamics of these ecosystems, covering vast parts of the globe. Therefore, a knowledge of the determinants of the distribution, movements and activities of herbivores, and how these interact with vegetation composition and dynamics, is required in order to predict the broader impact of these animals, now and into the future.

DIET SELECTION IN GOATS: A TEST OF INTAKE‐RATE MAXIMIZATION

The mechanisms of diet choice by herbivores are poorly understood. We tested whether the preference of goats among five grass species was accounted for by differences in intake rate or differences in species-specific attributes of the grasses. When offered a choice between grass species, the goats selected diets that tended to maximize intake rate. Only a small amount of the residual variation was explained by the individual preference of goats for each grass species. The animals exhibited partial preferences, only leaving the lower intake-rate alternative ungrazed in a quarter of the trials. It is argued that the most likely explanation of this is the difficulty of discriminating between alternatives of similar intake rate. Discrimination error is likely to be a general explanation of mixed diets in herbivores, regardless of whether other explanations, such as rate maximization subject to nutrient constraints, also apply.