The maximum attainable body size of herbivorous mammals: morphophysiological constraints on foregut, and adaptations of hindgut fermenters AbstractAn oft-cited nutritional advantage of large body size is that larger animals have lower relative energy requirements and that, due to their increased gastrointestinal tract (GIT) capacity, they achieve longer ingesta passage rates, which allows them to use forage of lower quality. However, the fermentation of plant material cannot be optimized endlessly; there is a time when plant fibre is totally fermented, and another when energy losses due to methanogenic bacteria become punitive. Therefore, very large herbivores would need to evolve adaptations for a comparative acceleration of ingesta passage. To our knowledge, this phenomenon has not been emphasized in the literature to date. We propose that, among the extant herbivores, elephants, with their comparatively fast passage rate and low digestibility coefficients, are indicators of a trend that allowed even larger hindgut fermenting mammals to exist. The limited existing anatomical data on large hindgut fermenters suggests that both a relative shortening of the GIT, an increase in GIT diameter, and a reduced caecum might contribute to relatively faster ingesta passage; however, more anatomical data is needed to verify these hypotheses. The digestive physiology of large foregut fermenters presents a unique problem: ruminant-and nonruminant-forestomachs were designed to delay ingesta passage, and they limit food intake as a side effect. Therefore, with increasing body size and increasing absolute energy requirements, their relative capacity has to increase in order to compensate for this intake limitation. It seems that the foregut fermenting ungulates did not evolve species in which the intake-limiting effect of the foregut could be reduced, e.g. by special bypass structures, and hence this digestive model imposed an intrinsic body size limit. This limit will be lower the more the natural diet enhances the ingesta retention and hence the intake-limiting effect. Therefore, due to the mechanical characteristics of grass, grazing ruminants cannot become as big as the largest browsing ruminant. Ruminants are not absent from the very large body size classes because their digestive physiology offers no particular advantage, but because their digestive physiology itself intrinsically imposes a body size limit. We suggest that the decreasing ability for colonic water absorption in large grazing ruminants and the largest extant foregut fermenter, the hippopotamus, are an indication of this limit, and are the outcome of the competition of organs for the available space within the abdominal cavity. Our hypotheses are supported by the fossil record on extinct ruminant/tylopod species which did not, with the possible exception of the Sivatheriinae, surpass extant species in maximum body size. In contrast to foregut fermentation, the GIT design of hindgut fermenters allows adaptations for relative passage acceleration, which explai...

In the captive Indian rhinoceros (Rhinoceros unicornis), two disease complexes with a high incidence-chronic foot problems and uterine leiomyomas-may be linked to excess body weight (BW). In this study, intake and digestion trials were conducted (by means of 7-day weigh-backs, and 5-day total fecal collections, respectively) with I I Indian rhinoceroses at four zoological institutions in Europe and the United States to quantify energy and mineral nutrition on conventional or roughage-only diets. Diets comprising a variety of forages (grass hay only, a combination of grass hay and grass silage, straw, or a mixture of grass and legume hay) were offered as the roughage source, along with various concentrates, produce, and supplements. Water intake was quantified, and urine samples were obtained opportunistically. The animals consumed 0.5-1.1% of their BW in dry matter (DM) daily, with calculated digestible energy (DE, in megajoules MJ) values ranging from 0.27 to 0.99 MJ DE/kg BW0.75/day compared to an estimated requirement of 0.49-0.66 MJ DE/kg BW0.75/day. Seven of 11 rhinos (64%) fed restricted levels of concentrate plus forage consumed DE in excess of this estimate. Even on roughage-only diets, some individuals consumed energy well above the presumed metabolic requirements. Hence, restriction of both concentrates and roughage may be important for weight management in this species. Water intake ranged from 30 to 49 mL/kg BW daily (3.4-5.2 L/kg ingested DM), similar to values that have been reported for domestic equids. Excretion amounts and patterns also resembled those found in horses. Endogenous fecal losses measured for Ca, P, Cu, Fe, and Zn indicate that the maintenance requirements of these minerals should be met in Indian rhinoceroses by diets that meet recommendations for domestic horses. It is particularly important to evaluate dietary adequacy in mineral nutrition in this species in concert with the need for restricted energy intake, especially with regard to the hypothetical involvement of a low Zn supply in chronic foot problems

We performed intake, digestibility and ingesta passage studies in 11 Indian rhinoceroses (Rhinoceros unicornis) from four zoological institutions, using total faecal collection for the quantification of faecal output. The regularly fed zoo ration of roughage and concentrates (ration RC) and a roughage-only ration (ration R) were used; the roughage source differed between the facilities and comprised grass hay, grass silage, straw and lucerne hay. Dry matter intake ranged between 0.8 and 1.3% of body weight on ration RC and 0.5-1.2% on ration R. Digestibility coefficients achieved were similar to those reported for horses on diets of comparable composition. Endogenous losses as determined by linear regression analysis were within the range reported for horses. Measurements of faecal volatile fatty acids, faecal lactate and faecal pH also showed similarity to similar measurements in horses. The mean retention times of fluids (Co-EDTA) and particles (Cr-mordanted fibre <2 mm) in the whole gastrointestinal tract averaged 42 and 61 h, respectively, and were the longest ever recorded in a monogastric ungulate with this marker system. The results suggest that the horse is a useful model animal for designing diets for Indian rhinoceroses. Why digestive parameters are similar between these species in spite of enormous differences in body weight and retention times remains to be answered.