Modelling the nutritional ecology of ungulate herbivores: evolution of body size and competitive interactions

Summary A simulation model is used to quantify relationships between diet quality, digestive processes and body weight in ungulate herbivores. Retention time of food in the digestive tract is shown by regression to scale with W^0.27, and to be longer in ruminants than in hindgut fermenters. Allometric relationships between whole gut mean retention time (MRT, h) and weight (W) were: MRT=9.4 W^0.255 (r ²=0.80) for hindgut fermenters and MRT=15.3 W^0.251 (r ²=0.76) in ruminants. Longer retention of ingesta by large-bodied ruminants and hindgut fermenters increases digestive efficiency relative to small animals and permits them to survive on lower-quality foods. Compared with ruminants, hindgut fermenters' faster throughput is an advantage which outweighs their lower digestive efficiency, particularly on poor quality foods, provided that food resources are not limiting. This suggests that the predominance of ruminants in the middle range of body weights results from their more efficient use of scarce resources under conditions of resource depletion. Considering only physical limitations on intake, the model shows that the allometric coefficient which scales energy intake to body mass is 0.88 in ruminants and 0.82 in hindgut fermenters. The advantages of large body size are countered by disadvantages where food quantity is limited, and we suggest that the upper limit to ungulate body size is determined by the ability to extract nutrients from feeding niches during the nadir of the seasonal cycle of resource quality and abundance.     

Comparative chewing efficiency in mammalian herbivores

Although the relevance of particle size reduction in herbivore digestion is widely appreciated, few studies have investigated digesta particle size across species in relation to body mass or digestive strategy. We investigated faecal particle size, which reflects the size of ingesta particles after both mastication and specialized processes such as rumination. Particle size was measured by wet sieving samples from more than 700 captive individuals representing 193 mammalian species. Using phylogenetic generalized least squares, faecal particle size scaled to body mass with an exponent of 0.22 (95% confidence interval: 0.16–0.28). In comparisons among different digestive strategies, we found that (1) equids had smaller faecal particles than other hindgut fermenters, (2) non-ruminant foregut fermenters and hindgut fermenters had similar-sized faecal particles (not significantly different), and (3) ruminants had finer faecal particles than non-ruminants. These results confirm that the relationship between chewing efficiency and body mass is modified by morphological adaptations in dental design and physiological adaptations to chewing, such as rumination. This allometric relationship should be considered when investigating the effect of body size on digestive physiology, and digestion studies should include a measure of faecal particle size.     

哺乳动物的消化策略（英文）

理解动物的营养生态位是充分理解其整个生态学的基础,对于害兽控制和物种保护也很重要,食肉动物的小肠很发达,这可能与对食物的高消化能力有关;杂食性动物有更复杂的胃肠器官,其后端有可进行发酵的盲肠,消化物的平均滞留时间（mean retention times,MRTs)更长;最长的平均滞留时间见于食草动物,其消化道内高密度的微生物种群对不同滞留区内的消化物进行发酵,但是,并不是所有的食草动物都能够最大程度地消化植物纤维,只有反刍动物、骆驼和个体较大的后肠发酵动物（hindgut fermenter)能够具有这种能力,对比而言,许多其它的食草动物,如前肠发酵的有袋类和小型的后肠发酵动物如兔子、田鼠和负鼠等,它们具备可以使植物纤维消化效率最大的消化系统,可以在食物中的纤维素含量非常高的情况下仍能处理大量的食物。这些不同的消化策略使哺乳动物具有广幅的营养生态位。     

Fluid and particle retention times in the black rhinocerosDiceros bicornis, a large hindgut-fermenting browser

The mean retention time (MRT) of ingesta in the gastrointestinal tract is one of the major determinants of herbivore digestive physiology. We examined MRTs of fluids and particles in the gastrointestinal tract of six adult captive black rhinocerosesDiceros bicornis on conventional zoo diets. Fluid MRT ranged from 25–45 h and averaged 31 h. Particle MRT ranged from 28–59 h and averaged 38 h. In the six animals, both fluid and particle MRT declined as relative dry matter intake (g/kg metabolic body mass) increased. Black rhinoceroses, which are large hindgut-fermenting browsers, retained ingesta for a shorter period relative to their body size than grazing equids or grazing rhinoceros species. Our findings support the hypothesis that browsing hindgut fermenters have relatively shorter MRTs than grazing hindgut fermenters.     

Excretion patterns of fluid and different sized particle passage markers in banteng (Bos javanicus) and pygmy hippopotamus (Hexaprotodon liberiensis): two functionally different foregut fermenters

Processing of ingesta particles plays a crucial role in the digestive physiology of herbivores. In the ruminant forestomach different sized particles are stratified into a small and a large particle fraction and only the latter is regurgitated and remasticated to smaller, easier-to-digest particles. In contrast, it has been suggested that in non-ruminating foregut fermenters, such as hippopotamuses, larger particles should be selectively excreted since they tend to be digested at a slower rate and hence can be considered intake-limiting bulk. In our study we determined the mean retention time (MRT) of fluids and different sized particles (2 mm and 10 mm) in six pygmy hippos (Hexaprotodon liberiensis) and six banteng (Bos javanicus) on a diet of fresh grass at two intake levels. We used cobalt ethylendiamintetraacetate (Co-EDTA) as fluid and chromium (Cr)-mordanted fibre (2 mm) and cerium (Ce)-mordanted fibre (10 mm) as particle markers, mixed in the food. Average total tract MRT for fluid, small and large particles at the high intake level was 32, 76 and 73 h in pygmy hippos and 25, 56 and 60 h in banteng, and at the low intake level 39, 109, and 105 h in pygmy hippos and 22, 51 and 58 h in banteng, respectively. In accordance with the prediction, large particles moved faster than, or as fast as the small particles, through the gut of pygmy hippos. In contrast, large particles were excreted slower than the small particles in the ruminant of this study, the banteng. Pygmy hippos had longer retention times than the banteng, which probably compensate for the less efficient particle size reduction. Although the results were not as distinct as expected, most likely due to the fact that ingestive mastication of the larger particle marker could not be prevented, they confirm our hypothesis of a functional difference in selective particle retention between ruminating and non-ruminating foregut fermenters.     

A case of non-scaling in mammalian physiology? Body size, digestive capacity, food intake, and ingesta passage in mammalian herbivores

As gut capacity is assumed to scale linearly to body mass (BM), and dry matter intake (DMI) to metabolic body weight (BM(0.75)), it has been proposed that ingesta mean retention time (MRT) should scale to BM(0.25) in herbivorous mammals. We test these assumptions with the most comprehensive literature data collations (n=74 species for gut capacity, n=93 species for DMI and MRT) to date. For MRT, only data from studies was used during which DMI was also recorded. Gut capacity scaled to BM(1.06). In spite of large differences in feeding regimes, absolute DMI (kg/d) scaled to BM(0.76) across all species tested. Regardless of this allometry inherent in the dataset, there was only a very low allometric scaling of MRT with BM(0.14) across all species. If species were divided according to the morphophysiological design of their digestive tract, there was non-significant scaling of MRT with BM(0.04) in colon fermenters, BM(0.08) in non-ruminant foregut fermenters, BM(0.06) in browsing and BM(0.04) in grazing ruminants. In contrast, MRT significantly scaled to BM(0.24) (CI 0.16-0.33) in the caecum fermenters. The results suggest that below a certain body size, long MRTs cannot be achieved even though coprophagy is performed; this supports the assumption of a potential body size limitation for herbivory on the lower end of the body size range. However, above a 500 g-threshold, there is no indication of a substantial general increase of MRT with BM. We therefore consider ingesta retention in mammalian herbivores an example of a biological, time-dependent variable that can, on an interspecific level, be dissociated from a supposed obligatory allometric scaling by the morphophysiological design of the digestive tract. We propose that very large body size does not automatically imply a digestive advantage, because long MRTs do not seem to be a characteristic of very large species only. A comparison of the relative DMI (g/kg(0.75)) with MRT indicates that, on an interspecific level, higher intakes are correlated to shorter MRTs in caecum, colon and non-ruminant foregut fermenters; in contrast, no significant correlation between relative DMI and MRT is evident in ruminants.     

Fibre digestibility in large herbivores as related to digestion type and body mass — An in vitro approach

The coexistence of different ungulate species in a given ecosystem has been the focus of many studies. Differences between ruminant foregut fermenters and hindgut fermenters were remarkable for example in the way they ingest and digest high fibre diets. Digestion trials based on total collections are difficult to conduct or are sometimes even not possible for wild animals in the field or in zoos. To gain information on the fibre digestion achieved by these animals and the influence of body mass (BM) thereon, a method using spot sampling is desirable. In this study, in vitro fermentation of faecal neutral detergent fibre (NDF) was used as a measure of fibre digestion in large ungulates. Food and faecal samples of 10 ruminant foregut fermenting and 7 hindgut fermenting species/breeds were collected. All animals received 100% grass hay with ad libitum access. The NDF of food and faeces was fermented in vitro in a Hohenheim gas test (HGT) for 96 h. The digestion type generally had an effect on the gas production (GP) of faecal NDF in the HGT with hindgut fermenters showing higher values than ruminant foregut fermenters. At any time interval of incubation, BM had no influence on GP. The results are in accordance with both findings that ruminant foregut fermenters have longer mean retention times and more comprehensive particle reduction and findings of a lack of influence of BM on digesta mean retention time. It can be stated that the HGT (96 h) is a useful and quick method to show also small differences within groups in fibre digestion.     

The passage of digesta, particle size, and in vitro fermentation rate in the three-toed sloth Bradypus tridactylus (Edentata: Bradypodidae)

The digestive physiology of six three-toed sloths ( Bradypus tridactylus ) fed exclusively on Cecropia palmata foliage was studied. The mass of digesta in the gut was between 17 and 37% of body mass. This was between 133 and 282% of that expected in an animal of this size, based on published allometric equations. The concentration of total short chain fatty acids in the stomach was similar to that in the fermentative regions of other foregut fermenting herbivores but the rate of fermentation measured in vitro was very slow (6–12 mmol.l^-1.h^-1) and substantially lower than that observed using similar techniques in other small foregut fermenters.
The overall (dose to excretion) mean retention time of particulate and solute digesta markers was about 150 h. Most of this (73%) occurred in the stomach but a substantial proportion (17%) could be attributed to the storage of faeces in the rectum.
The slow rate of passage of digesta through the gut together with the slow rate of fermentation in the stomach is not typical of small foregut fermenting herbivores. However, such a pattern is feasible in Bradypus tridactylus because of the large volume of digesta retained in the gut and the very low metabolic rate of these mammals.     

Modelling digestive constraints in non-ruminant and ruminant foregut-fermenting mammals

It has been suggested that large foregut-fermenting marsupial herbivores, the kangaroos and their relatives, may be less constrained by food intake limitations as compared with ruminants, due mainly to differences in their digestive morphology and management of ingesta particles through the gut. In particular, as the quality of forage declines with increasing contents of plant fibre (cellulose, hemicelluloses and lignin; measured as neutral-detergent fibre, NDF), the tubiform foregut of kangaroos may allow these animals to maintain food intakes more so than ruminants like sheep, which appear to be limited by fibrous bulk filling the foregut and truncating further ingestion. Using available data on dry matter intake (DMI, g kg(-0.75) d(-1)), ingesta mean retention time (MRT, h), and apparent digestibility, we modelled digestible dry matter intake (DDMI) and digestible energy intake (DEI) by ruminant sheep (Ovis aries) and by the largest marsupial herbivore, the red kangaroo (Macropus rufus). Sheep achieved higher MRTs on similar DMIs, and hence sheep achieved higher DDMIs for any given level of DMI as compared with kangaroos. Interestingly, MRT declined in response to increasing DMI in a similar pattern for both species, and the association between DMI and plant NDF contents did not support the hypothesis that kangaroos are less affected by increasing fibre relative to sheep. However, when DEI was modelled according to DDMIs and dietary energy contents, we show that the kangaroos could meet their daily maintenance energy requirements (MER) at lower levels of DMI and on diets with higher fibre contents compared with sheep, due largely to the kangaroos' lower absolute maintenance and basal energy metabolisms compared with eutherians. These results suggest that differences in the metabolic set-point of different species can have profound effects on their nutritional niche, even when their digestive constraints are similar, as was the case for these ruminant and non-ruminant foregut fermenters.     

The influence of natural diet composition, food intake level, and body size on ingesta passage in primates

An important component of digestive physiology involves ingesta mean retention time (MRT), which describes the time available for digestion. At least three different variables have been proposed to influence MRT in herbivorous mammals: body mass, diet type, and food intake (dry matter intake, DMI). To investigate which of these parameters influences MRT in primates, we collated data for 19 species from trials where both MRT and DMI were measured in captivity, and acquired data on the composition of the natural diet from the literature. We ran comparative tests using both raw species values and phylogenetically independent contrasts. MRT was not significantly associated with body mass, but there was a significant correlation between MRT and relative DMI (rDMI, g/kg(0.75)/d). MRT was also significantly correlated with diet type indices. Thus, both rDMI and diet type were better predictors of MRT than body mass. The rDMI-MRT relationship suggests that primate digestive differentiation occurs along a continuum between an "efficiency" (low intake, long MRT, high fiber digestibility) and an "intake" (high intake, short MRT, low fiber digestibility) strategy. Whereas simple-stomached (hindgut fermenting) species can be found along the whole continuum, foregut fermenters appear limited to the "efficiency" approach.     

The potential interplay of posture, digestive anatomy, density of ingesta and gravity in mammalian herbivores: why sloths do not rest upside down

1. The interaction between the density of ingesta and gravity observed in the digestive systems of ruminant herbivores should receive attention in other non‐ruminant herbivorous mammals. The resting postures adopted by non‐ruminants are of particular interest. 2. A new interpretation of established findings regarding the digestive tract of sloths illustrates that the interplay of posture, anatomy, the density of ingesta and gravity can provide a novel explanation of behavioural and morphological adaptations in herbivores, as the average particle size and dry‐matter content increases within their forestomach from its caudal towards its cranial portion. In sloths, this could be indicative of a stratification of ingesta occurring in the upright sitting posture adopted while resting, as opposed to their characteristic upside down posture when moving. 3. The sitting resting posture of sloths could therefore be an adaptation to exploit the tendency of the forestomach contents to stratify in order to pass larger, more difficult‐to‐digest particles faster from the fermentation chamber.     

Intake, ingesta retention, particle size distribution and digestibility in the hippopotamidae

Although several aspects of the digestive physiology of the hippopotamidae-non-ruminating foregut fermenters-have been described, ingesta kinetics and passage characteristics of these species are not well understood. The most outstanding feature of the hippo digestive physiology reported so far is the very long mean ingesta retention times (MRTs) measured by Foose [Foose, T., 1982. Trophic strategies of ruminant versus nonruminant ungulates. PhD dissertation, University of Chicago, Chicago.]. Since those data had been investigated with animals without water access, we intended to measure MRT in hippos which were allowed to enter water pools during the night. MRT parameters as well as dry matter (DM) digestibility were determined in four common (Hippopotamus amphibius) and four pygmy hippos (Hexaprotodon liberiensis) on two different diets each using cobalt ethylendiamintetraacetate (Co-EDTA) as a fluid, chromium (Cr)-mordanted fibre (<2 mm) as a particle and acid detergent lignin (ADL) as an internal digestibility marker. Four of the animals additionally received cerium (Ce)-mordanted fibres (2-10 mm) as particle markers. Total MRTs for fluids and particles ranged between 20-35 and 48-106 h in the common and between 13-39 and 32-107 h in the pygmy hippos. The difference between fluid and particle retention was greater than usually reported in ruminants. Excretion patterns of the markers differed from those usually observed in ruminants but resembled those reported for macropods (kangaroos), indicating a plug-flow reactor-like physiology in the hippo forestomach (FRST). This finding complements other described similarities between the macropod and the hippo forestomach. The measurements of larger particle retention profiles suggest that in the hippo, larger particles might be excreted either faster or at the same rate as smaller particles, indicating a general difference between ruminants and hippos with respect to differential particle retention. The digestive physiology of hippos is characterised by a generally low food intake, long ingesta retention times and dry matter digestibilities lower than reported in ruminants. Moderate digestibilities in spite of long retention times might be the result of the generally high average ingesta particle size in hippos. The comparatively easy management of pygmy hippos, together with the significant correlations between food intake, MRT and digestibility in the pygmy hippos of this study, recommends this species for further studies on the interplay of these parameters in herbivore digestive physiology.     

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

Summary Ruminants 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.     

More efficient mastication allows increasing intake without compromising digestibility or necessitating a larger gut: Comparative feeding trials in banteng (Bos javanicus) and pygmy hippopotamus (Hexaprotodon liberiensis)

The digestion of plant material in mammalian herbivores basically depends on the chemical and structural composition of the diet, the mean particle size to which the forage is processed, and the ingesta retention time. These different factors can be influenced by the animal, and they can presumably compensate for each other. The pygmy hippopotamus, a non-ruminating foregut fermenter, has longer mean retention times than ruminants; however hippos do not achieve higher (fibre) digestibilities on comparable diets, which could be due to ineffective mastication. We performed feeding trials with six pygmy hippos (Hexaprotodon liberiensis) and six banteng cattle (Bos javanicus) on a grass diet. As predicted, both species achieved similar dry matter, organic matter, crude protein and gross energy digestibilities. However, neutral and acid detergent fibre digestibility was lower in pygmy hippos. Apparently, in these species, fibre digestibility was more influenced by particle size, which was larger in pygmy hippos compared to banteng, than by retention time. In spite of their higher relative food intake, the banteng in this study did not have greater relative gut fills than the hippos. Ruminants traditionally appear intake-limited when compared to equids, because feed particles above a certain size cannot leave the rumen. But when compared to nonruminating foregut fermenters, rumination seems to free foregut fermenters from an intrinsic food intake limitation. The higher energy intakes and metabolic rates in wild cattle compared to hippos could have life-history consequences, such as a higher relative reproductive rate.     

Fibrobacter communities in the gastrointestinal tracts of diverse hindgut‐fermenting herbivores are distinct from those of the rumen

The genus Fibrobacter contains cellulolytic bacteria originally isolated from the rumen. Culture‐independent investigations have since identified Fibrobacter populations in the gastrointestinal tracts of numerous hindgut‐fermenting herbivores, but their physiology is poorly characterized due to few representative axenic cultures. To test the hypothesis that novel Fibrobacter diversity exists in hindgut fermenters, we performed culturing and 16S rRNA gene amplicon sequencing on samples collected from phylogenetically diverse herbivorous hosts. Using a unique approach for recovering axenic Fibrobacter cultures, we isolated 45 novel strains from 11 different hosts. Full‐length 16S rRNA gene sequencing of these isolates identified nine discrete phylotypes (cutoff = 0.03%) among them, including several that were only isolated from hindgut‐fermenting hosts, and four previously unrepresented by axenic cultures. Our phylogenetic analysis indicated that six of the phylotypes are more closely related to previously described subspecies of Fibrobacter succinogenes, while the remaining three were more closely related to F. intestinalis. Culture‐independent bacterial community profiling confirmed that most isolates were representative of numerically dominant phylotypes in their respective samples and strengthened the association of certain phylotypes with either ruminants or hindgut‐fermenters. Despite considerable phylogenetic diversity observed among the Fibrobacter strains isolated here, phenotypic characterization suggests a conserved specialization for growth on cellulose.     

Differences in selective reticulo-ruminal particle retention as a key factor in ruminant diversification

The measurement of passage rate is important for the concept of ruminant diversification. While supporters of Hofmann's 1989 feeding type classification claim that browsing ruminants have faster passage rates than grazing ruminants, other researchers consider the passage rate to depend on body size alone. To date, no convincing comparison of ruminant passage rates has been put forward. For comparative purposes, we suggest the use of the "selectivity factor", which is an expression of how much longer particles of a defined size (<2 mm) are retained in the ruminant digestive tract than fluids. From the limited data available, it seems that grazing ruminants display selectivity factors between 1.56 and 3.80, whereas browsers have a much narrower range of 1.14-1.80. This suggests that browsers are not able to selectively retain particles as long as grazers. Intake of browsers, on the other hand, may not be limited by physical fill of the forestomach to the same degree as in grazers. This result can explain several observations on the digestive physiology of browsers, some of which have been linked to a rumen bypass mechanism. We propose that the ability for selective particle retention is a key factor for understanding the physiological consequences of ruminant diversification.     

Phylogenetic constraints on digesta separation: Variation in fluid throughput in the digestive tract in mammalian herbivores

The relevance of the mean retention time (MRT) of particles through the gastrointestinal tract (GIT) is well understood and MRT(particle)GIT is an important parameter in digestion models. Solute markers have been used to estimate MRT(solute)GIT (or 'fluid passage') in animals, but the relevance of this measure is less evident and is usually sought in its relation to MRT(particle)GIT. The ratio between the two measures indicates the degree of 'digesta washing', with little washing occurring at ratios of 1, aborad washing at ratios >1 (where the solute marker travels faster than the particle marker), and orad (retrograde) washing at ratios <1 (where the solute marker travels slower than the particle marker). We analysed digesta washing in a dataset of 98 mammalian species including man of different digestion types (caecum, colon and nonruminant foregut fermenters, and ruminants), controlling for phylogeny; a subset of 72 species allowed testing for the influence of food intake level. The results indicate that MRT(solute)GIT and the degree of digesta washing are related to digestion type, whereas variation in MRT(particle)GIT is influenced mainly by effects of body mass and food intake. Thus, fluid throughput and digesta washing emerge as important correlates of digestive anatomy. Most importantly, primates appear constrained to little digesta washing compared to non-primate mammalian herbivores, regardless of their digestion type. These results may help explain the absence of primates from certain herbivore niches and represent a drastic example of a physiologic limitation in a phylogenetic group. More experimental research is required to illuminate relative benefits and costs of digesta washing.     

The relative merits of foregut and hindgut fermentation

Mammalian herbivores cannot break down cellulose except by fermentation, and may have Termentation chambers at either end of the gut: ruminants have their principal fermentation chamber in the stomach but horses ferment only in the hindgut. A mathematical model (Alexander, 1991) predicted that foregut fermenters should do better than hindgut fermenters on poor foods, and the reverse on richer. less fibrous foods. Further, the optimum gut for poor foods would have the hindgut fermentation chamber only a little smaller than the foregut chamber. However. it has been claimed that horses do better than ruminants on poor food, and the hindgut of ruminants is much smaller than the rumen.
In this paper, the basic model is modified in ways designed to make it more realistic and the effects are investigated. None of the modifications alters the conclusion that the optimum gut for poor food has a large foregut fermentation chamber. However, the optimum proportions of fore-to hindgut, for poor diets, become more like those of real ruminants when account is taken of the diminishing volume of the food passing through the gut, and of incomplete mixing in the rumen.     

Habitat heterogeneity as a driver of ungulate diversity and distribution patterns: interaction of body mass and digestive strategy

Aim Classic island biogeographical theory predicts that reserves have to be large to conserve high biodiversity. Recent literature, however, suggests that habitat heterogeneity can counterbalance the effect of small reserve size. For savanna ungulates, body mass is said to drive habitat selection and facilitate species coexistence, where large species use a higher proportion of the landscape than smaller species, because a wider food quality tolerance allows them to use a higher diversity of habitat types. In this case, high habitat heterogeneity would facilitate diverse assemblages of different‐sized ungulates. Digestive physiology should further modify this relationship, because non‐ruminants have a wider diet tolerance than ruminants. We tested this hypothesis with an empirical dataset on distribution and habitat preference of different‐sized African grazers. Location Hluhluwe‐iMfolozi Park, Republic of South Africa. Methods We recorded herbivore dung and habitat type on 24 line transects varying between 4 and 11 km with a total length of 190 km to determine habitat selection and landscape distribution of six grazer species, three ruminants and three non‐ruminants. Results Larger ruminant grazers were more evenly distributed than smaller ruminants, had a more diverse use of habitats and used more low quality habitat. In contrast, non‐ruminant grazers were more evenly distributed than similar‐sized ruminants and body mass did not clearly influence diversity of habitat use and use of low quality habitat. Main conclusions We confirm that body mass influences diversity of habitat use of large herbivores but digestive strategy potentially modifies this relationship. Hence, habitat heterogeneity might facilitate herbivore diversity in savanna ecosystems and high heterogeneity might counterbalance the effects of fragmentation and declining reserve size. Concluding, processes that homogenize the landscape, such as fire (mis)management and artificial waterholes, might be as threatening to biodiversity as landscape fragmentation, especially for smaller ruminant herbivores.     

Aspects of digestion and in vitro fermentation in the caecum of some East African herbivores

Caecal digestive functions were compared in 22 species of East African herbivores. Comparisons were made between ruminant pseudo-ruminant, and non-ruminant herbivores to assess the relative in vitro fermentation rate and composition of caecal contents from these species observed in their natural habitat. Measurements were made of caecal fermentation rate, organic acid composition, osmolality, pH and dry matter content. The data were compared by foregut structure, feed preference and body weight of the herbivores.