The herbivorous sauropod dinosaurs of the Jurassic and Cretaceous periods were the largest terrestrial animals ever, surpassing the largest herbivorous mammals by an order of magnitude in body mass. Several evolutionary lineages among Sauropoda produced giants with body masses in excess of 50 metric tonnes by conservative estimates. With body mass increase driven by the selective advantages of large body size, animal lineages will increase in body size until they reach the limit determined by the interplay of bauplan, biology, and resource availability. There is no evidence, however, that resource availability and global physicochemical parameters were different enough in the Mesozoic to have led to sauropod gigantism.We review the biology of sauropod dinosaurs in detail and posit that sauropod gigantism was made possible by a specific combination of plesiomorphic characters (phylogenetic heritage) and evolutionary innovations at different levels which triggered a remarkable evolutionary cascade. Of these key innovations, the most important probably was the very long neck, the most conspicuous feature of the sauropod bauplan. Compared to other herbivores, the long neck allowed more efficient food uptake than in other large herbivores by covering a much larger feeding envelope and making food accessible that was out of the reach of other herbivores. Sauropods thus must have been able to take up more energy from their environment than other herbivores.The long neck, in turn, could only evolve because of the small head and the extensive pneumatization of the sauropod axial skeleton, lightening the neck. The small head was possible because food was ingested without mastication. Both mastication and a gastric mill would have limited food uptake rate. Scaling relationships between gastrointestinal tract size and basal metabolic rate (BMR) suggest that sauropods compensated for the lack of particle reduction with long retention times, even at high uptake rates.The extensive pneumatization of the axial skeleton resulted from the evolution of an avian-style respiratory system, presumably at the base of Saurischia. An avian-style respiratory system would also have lowered the cost of breathing, reduced specific gravity, and may have been important in removing excess body heat. Another crucial innovation inherited from basal dinosaurs was a high BMR. This is required for fueling the high growth rate necessary for a multi-tonne animal to survive to reproductive maturity.The retention of the plesiomorphic oviparous mode of reproduction appears to have been critical as well, allowing much faster population recovery than in megaherbivore mammals. Sauropods produced numerous but small offspring each season while land mammals show a negative correlation of reproductive output to body size. This permitted lower population densities in sauropods than in megaherbivore mammals but larger individuals.Our work on sauropod dinosaurs thus informs us about evolutionary limits to body size in other groups of herbivorous terrestrial tetrapo...

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.

Differences in allometric scaling of physiological characters have the appeal to explain species diversification and niche differentiation along a body mass (BM) gradient -because they lead to different combinations of physiological properties, and thus may facilitate different adaptive strategies. An important argument in physiological ecology is built on the allometries of gut fill (assumed to scale to BM1.0) and energy requirements/intake (assumed to scale to BM0.75) in mammalian herbivores. From the difference in exponents, it has been postulated that the mean retention time (MRT) of digesta should scale to BM1.0-0.75 = BM0.25. This has been used to argue that larger animals have an advantage in digestive efficiency and hence can tolerate lower-quality diets. However, empirical data does not support the BM0.25 scaling of MRT, and the deduction of MRT scaling implies, according to physical principles, no scaling of digestibility; basing assumptions on digestive efficiency on the thus-derived MRT scaling amounts to circular reasoning. An alternative explanation considers a higher scaling exponent for food intake than for metabolism, allowing larger animals to eat more of a lower quality food without having to increase digestive efficiency; to date, this concept has only been explored in ruminants. Here, using data for 77 species in which intake, digestibility and MRT were measured (allowing the calculation of the dry matter gut contents DMC), we show that the unexpected shallow scaling of MRT is common in herbivores and may result from deviations of other scaling exponents from expectations. Notably, DMC have a lower scaling exponent than 1.0, and the 95% confidence intervals of the scaling exponents for intake and DMC generally overlap. Differences in the scaling of wet gut contents and dry matter gut contents confirm a previous finding that the dry matter concentration of gut contents decreases with body mass, possibly compensating for the less favourable volume-surface ratio in the guts of larger organisms. These findings suggest that traditional explanations for herbivore niche differentiation along a BM gradient should not be based on allometries of digestive physiology. In contrast, they support the recent interpretation that larger species can tolerate lower-quality diets because their intake has a higher allometric scaling than their basal metabolism, allowing them to eat relatively more of a lower quality food without having to increase digestive efficiency. MRT scaling amounts to circular reasoning. An alternative explanation considers a higher 38 scaling exponent for food intake than for metabolism, allowing larger animals to eat more of a 39 lower quality food without having to increase digestive efficiency; to date, this concept has 40 only been explored in ruminants. Here, using data for 77 species in which intake, digestibility 41 and MRT were measured (allowing the calculation of the dry matter gut contents DMC), we 42show that the unexpected shallow scaling of MRT is common in herbi...

Comparative physiology applies methods established in domestic animal science to a wider variety of species. This can lead to improved insight into evolutionary adaptations of domestic animals, by putting domestic species into a broader context. Examples include the variety of responses to seasonally fluctuating environments, different adaptations to heat and drought, and in particular adaptations to herbivory and various herbivore niches. Herbivores generally face the challenge that a high food intake compromises digestive efficiency (by reducing ingesta retention time and time available for selective feeding and for food comminution), and a variety of digestive strategies have evolved in response. Ruminants are very successful herbivores. They benefit from potential advantages of a forestomach without being constrained in their food intake as much as other foregut fermenters, because of their peculiar reticuloruminal sorting mechanism that retains food requiring further digestion but clears the forestomach of already digested material; the same mechanism also optimises food comminution. Wild ruminants vary widely in the degree to which their rumen contents 'stratify', with little stratification in 'moose-type' ruminants (which are mostly restricted to a browse niche) and a high degree of stratification into gas, particle and fluid layers in 'cattle-type' ruminants (which are more flexible as intermediate feeders and grazers). Yet all ruminants uniformly achieve efficient selective particle retention, suggesting that functions other than particle retention played an important role in the evolution of stratification-enhancing adaptations. One interesting emerging hypothesis is that the high fluid turnover observed in 'cattle-type' ruminants -which is a prerequisite for stratification -is an adaptation that not only leads to a shift of the sorting mechanism from the reticulum to the whole reticulorumen, but also optimises the harvest of microbial protein from the forestomach. Although potential benefits of this adaptation have not been quantified, the evidence for convergent evolution toward stratification suggests that they must be substantial. In modern production systems, the main way in which humans influence the efficiency of energy uptake is by manipulating diet quality. Selective breeding for conversion efficiency has resulted in notable differences between wild and domestic animals. With increased knowledge on the relevance of individual factors, that is fluid throughput through the reticulo-rumen, more specific selection parameters for breeding could be defined to increase productivity of domestic ruminants by continuing certain evolutionary trajectories.Keywords: herbivory, hindgut fermenter, foregut fermenter, browser, grazer ImplicationsUnderstanding evolutionary adaptations of ruminants will have an impact on (i) husbandry of captive wild ruminants, many of which cannot be kept or fed as domestic ruminants and (ii) research for continuous refinement of the production potential of domestic ruminants, ...