Gross intestinal morphometry and allometry in ruminants.

While some descriptions of ruminants' dietary adaptations suggest that the length of the intestinal tract reflects the proportion of grass or browse in the diet, this assumption has been questioned. We collated data on body mass (BM), as well as small intestine, caecum, colon/rectum, large and total intestine length in 68 ruminant species, and, while accounting for the phylogenetic structure of the dataset, evaluated both allometric scaling and the potential influence of diet, digestive physiology or climate proxies on measures of intestine length. Intestinal length generally scaled to BM at an exponent higher than the 0.33 expected due to geometry. Diet or digestive physiology proxies did not have an influence on any intestinal length measures, though some proxies indicating more arid natural habitats were positively correlated with measures of the large intestine. The relative size of a forestomach compartment, the omasum, was negatively correlated with intestine length. The results indicate that intestine length measures provide little indication of feeding type or digestive physiology, but rather indicate adaptations to aridity. Higher-than-geometry scaling of intestinal length may be related to the necessity of maintaining geometric (or metabolic) scaling of intestinal surface area while keeping gut diameter, and hence the diffusion distances, small. The way in which space trade-offs determine the macroanatomy of different organs in the abdominal cavity, such as the omasum and the intestine, deserves further investigation.

Comparative omasum anatomy in ruminants: Relationships with natural diet, digestive physiology, and general considerations on allometric investigations.

The omasum is the third forestomach compartment of pecoran ruminants. It is assumed that the re-absorption of fluid present in the forestomach digesta (that facilitates particle sorting, digestion, and harvest of microbes) is its main function, so that less diluted digesta is submitted to enzymatic digestion in the lower digestive tract. Here, we evaluate measures of omasum size (representing 84 ruminant species in the largest data set) against body mass and proxies of the natural diet (%grass) or forestomach physiology (fluid throughput), using phylogenetically controlled models. The origin of specimens (free-ranging or captive) did not have an effect in the data set. Models with the best support invariably either included %grass or a physiology proxy in addition to body mass. These effects were not necessarily additive (affecting the intercept of the allometric regression), but often indicated a change in the allometric body mass-exponent with diet or physiology. Only models that allowed an influence on the allometric exponent yielded basic exponents compatible with predictions derived from geometry. Species that include more grass in their natural diet, or that have a "cattle-type" physiology marked by a high forestomach fluid throughput, generally have larger omasa. However, the existence of outliers, as well as the overall data pattern, suggest that this is not an obligatory morphophysiological condition. Circumstantial evidence is presented leading to the hypothesis that the comparatively small and less complex omasa of "moose-type" species do not necessarily represent an "original" state, but may be derived from more complex states by ontogenetic reduction and fusion of omasal laminae.

Convergence of macroscopic tongue anatomy in ruminants and scaling relationships with body mass or tongue length.

Various morphological measures demonstrate convergent evolution in ruminants with their natural diet, in particular with respect to the browser/grazer dichotomy. Here, we report quantitative macroanatomical measures of the tongue (length and width of specific parts) of 65 ruminant species and relate them to either body mass (BM) or total tongue length, and to the percentage of grass in the natural diet (%grass). Models without and with accounting for the phylogenetic structures of the dataset were used, and models were ranked using Akaike's Information Criterion. Scaling relationships followed geometric principles, that is, length measures scaled with BM to the power of 0.33. Models that used tongue length rather than BM as a body size proxy were consistently ranked better, indicating that using size proxies that are less susceptible to a wider variety of factors (such as BM that fluctuates with body condition) should be attempted whenever possible. The proportion of the freely mobile tongue tip of the total tongue (and hence also the corpus length) was negatively correlated to %grass, in accordance with concepts that the feeding mechanism of browsers requires more mobile tongues. It should be noted that some nonbrowsers, such as cattle, use a peculiar mechanism for grazing that also requires long, mobile tongues, but they appear to be exceptions. A larger corpus width with increasing %grass corresponds to differences in snout shape with broader snouts in grazers. The Torus linguae is longer with increasing %grass, a finding that still warrants functional interpretation. This study shows that tongue measures covary with diet in ruminants. In contrast, the shape of the tongue (straight or "hourglass-shaped" as measured by the ratio of the widest and smallest corpus width) is unrelated to diet and is influenced strongly by phylogeny.

The intraruminal papillation gradient in wild ruminants of different feeding types: Implications for rumen physiology.

Browsing and grazing ruminants are thought to differ in the degree their rumen contents are stratified-which may be due to different characteristics of their respective forages, to particular adaptations of the animals, or both. However, this stratification is difficult to measure in live animals. The papillation of the rumen has been suggested as an anatomical proxy for stratification-with even papillation indicating homogenous contents, and uneven papillation (with few and small dorsal and ventral papillae, and prominent papillae in the atrium ruminis) stratified contents. Using the surface enlargement factor (SEF, indicating how basal mucosa surface is increased by papillae) of over 55 ruminant species, we demonstrate that differences between the SEF(dorsal) or SEF(ventral) and the SEF(atrium) are significantly related to the percentage of grass in the natural diet. The more a species is adapted to grass, the more distinct this difference, with extreme grazers having unpapillated dorsal and ventral mucosa. The relative SEF(dorsal) as anatomical proxy for stratification, and the difference in particle and fluid retention in the rumen as physiological proxy for stratification, are highly correlated in species (n = 9) for which both kind of data are available. The results support the concept that the stratification of rumen contents varies among ruminants, with more homogenous contents in the more browsing and more stratified contents in the more grazing species.