Is there an influence of body mass on digesta mean retention time in herbivores? A comparative study on ungulates.

The relation between body mass (BM) and digesta mean retention time (MRT) in herbivores was the focus of several studies in recent years. It was assumed that MRT scaled with BM(0.25) based on the isometric scaling of gut capacity (BM(1.0)) and allometric scaling of energy intake (BM(0.75)). Literature studies that tested this hypothesis produced conflicting results, arriving sometimes at higher or lower exponents than the postulated 0.25. This study was conducted with 8 ruminants (n=2-6 per species) and 6 hindgut fermenting species/breeds (n=2-6, warthog n=1) with a BM range of 60-4000 kg. All animals received a ration of 100% grass hay with ad libitum access. Dry matter intake was measured and the MRT was estimated by the use of a solute and a particle (1-2 mm) marker. No significant scaling of MRT(particle) with BM was observed for all herbivores (32 BM(0.04), p=0.518) and hindgut fermenters (32 BM(0.00), p=1.00). The scaling exponent for ruminants only showed a tendency towards significance (29 BM(0.12), p=0.071). Ruminants on average had an MRT(particle) 1.61-fold longer than hindgut fermenters. Whereas an exponent of 0.25 is reasonable from theoretical considerations, much lower exponents were found in this and other studies. The energetic benefit of increasing MRT is by no means continuous, since the energy released from a given food unit via digestion decreases over time. The low and non-significant scaling factors for both digestion types suggest that in ungulates, MRT is less influenced by BM (maximal allometric exponent ≤0.1) than often reported.

Methane output of tortoises: its contribution to energy loss related to herbivore body mass.

An increase in body mass (M) is traditionally considered advantageous for herbivores in terms of digestive efficiency. However, recently increasing methane losses with increasing M were described in mammals. To test this pattern in non-mammal herbivores, we conducted feeding trails with 24 tortoises of various species (M range 0.52-180 kg) fed a diet of grass hay ad libitum and salad. Mean daily dry matter and gross energy intake measured over 30 consecutive days scaled to M(0.75 (95%CI 0.64-0.87)) and M(0.77 (95%CI 0.66-0.88)), respectively. Methane production was measured over two consecutive days in respiration chambers and scaled to M(1.03 (95%CI 0.84-1.22)). When expressed as energy loss per gross energy intake, methane losses scaled to 0.70 (95%CI 0.47-1.05) M(0.29 (95%CI 0.14-0.45)). This scaling overlaps in its confidence intervals to that calculated for nonruminant mammals 0.79 (95%CI 0.63-0.99) M(0.15 (95%CI 0.09-0.20)), but is lower than that for ruminants. The similarity between nonruminant mammals and tortoises suggest a common evolution of the gut fauna in ectotherms and endotherms, and that the increase in energetic losses due to methane production with increasing body mass is a general allometric principle in herbivores. These findings add evidence to the view that large body size itself does not necessarily convey a digestive advantage.

Herbivorous reptiles and body mass: effects on food intake, digesta retention, digestibility and gut capacity, and a comparison with mammals.

Differences in the allometric scaling between gut capacity (with body mass, BM¹·°°) and food intake (with BM°·75) should theoretically result in a scaling of digesta retention time with BM°·²5 and therefore a higher digestive efficiency in larger herbivores. This concept is an important part of the so-called 'Jarman-Bell principle' (JBP) that explains niche differentiation along a body size gradient in terms of digestive physiology. Empirical data in herbivorous mammals, however, do not confirm the scaling of retention time, or of digestive efficiency, with body mass. Here, we test these concepts in herbivorous reptiles, adding data of an experiment that measured food intake, digesta retention, digestibility and gut capacity in 23 tortoises (Testudo graeca, T. hermanni , Geochelone nigra, G. sulcata, Dipsochelys dussumieri) across a large BM range (0.5-180 kg) to a literature data collection. While dry matter gut fill scaled to BM¹·°7 and dry matter intake to BM°·76, digesta mean retention time (MRT) scaled to BM°·¹7; the scaling exponent was not significantly different from zero for species > 1 kg. Food intake level was a major determinant of MRT across reptiles and mammals. In contrast to dietary fibre level, BM was not a significant contributor to dry matter digestibility in a General Linear Model. Digestibility coefficients in reptiles depended on diet nutrient composition in a similar way as described in mammals. Although food intake is generally lower and digesta retention longer in reptiles than in mammals, digestive functions scale in a similar way in both clades, indicating universal principles in herbivore digestive physiology. The reasons why the theoretically derived JBP has little empirical support remain to be investigated. Until then, the JBP should not be evoked to explain niche differentiation along a body size axis in terms of digestive physiology.

Methane output of rabbits (Oryctolagus cuniculus) and guinea pigs (Cavia porcellus) fed a hay-only diet: implications for the scaling of methane production with body mass in non-ruminant mammalian herbivores.

It is assumed that small herbivores produce negligible amounts of methane, but it is unclear whether this is a physiological peculiarity or simply a scaling effect. A respiratory chamber experiment was conducted with six rabbits (Oryctolagus cuniculus, 1.57±0.31 kg body mass) and six guinea pigs (Cavia porcellus, 0.79±0.07 kg) offered grass hay ad libitum. Daily dry matter (DM) intake and DM digestibility were 50±6 g kg⁻°·75 d⁻¹ and 55±6% in rabbits and 59±11 g kg⁻°·75 d⁻¹ and 61±3% in guinea pigs, respectively. Methane production was similar for both species (0.20±0.10 L d⁻¹ and 0.22±0.08L d⁻¹ and represented 0.69±0.32 and 1.03±0.29% of gross energy intake in rabbits and guinea pigs, respectively. In relation to body mass (BM) guinea pigs produced significantly more methane. The data on methane per unit of BM obtained in this study and from the literature on the methane output of elephant, wallabies and hyraxes all lay close to a regression line derived from roughage-fed horses, showing an increase in methane output with BM. The regression, including all data, was nearly identical to that based on the horse data only (methane production in horses [L d⁻¹]=0.18 BM [kg]°·97(95%CI °·9²â»¹·°²)) and indicates linear scaling. Because feed intake typically scales to BM°·75, linear scaling of methane output translates into increasing energetic losses at increasing BM. Accordingly, the data collection indicates that an increasing proportion of ingested gross energy is lost because relative methane production increases with BM. Different from ruminants, such losses (1%-2% of gross energy) appear too small in non-ruminant herbivores to represent a physiologic constraint on body size. Nevertheless, this relationship may represent a physiological disadvantage with increasing herbivore body size.

Allometry of visceral organs in living amniotes and its implications for sauropod dinosaurs.

Allometric equations are often used to extrapolate traits in animals for which only body mass estimates are known, such as dinosaurs. One important decision can be whether these equations should be based on mammal, bird or reptile data. To address whether this choice will have a relevant influence on reconstructions, we compared allometric equations for birds and mammals from the literature to those for reptiles derived from both published and hitherto unpublished data. Organs studied included the heart, kidneys, liver and gut, as well as gut contents. While the available data indicate that gut content mass does not differ between the clades, the organ masses for reptiles are generally lower than those for mammals and birds. In particular, gut tissue mass is significantly lower in reptiles. When applying the results in the reconstruction of a sauropod dinosaur, the estimated volume of the coelomic cavity greatly exceeds the estimated volume of the combined organ masses, irrespective of the allometric equation used. Therefore, substantial deviation of sauropod organ allometry from that of the extant vertebrates can be allowed conceptually. Extrapolations of retention times from estimated gut contents mass and food intake do not suggest digestive constraints on sauropod dinosaur body size.

The Coolidge effect, individual recognition and selection for distinctive cuticular signatures in a burying beetle.

The ability to recognize individuals is an important aspect of social interactions, but it can also be useful to avoid repeated matings with the same individual. The Coolidge effect is the progressive decline in a male's propensity to mate with the same female combined with a heightened sexual interest in new females. Although males that recognize previous partners and show a preference for novel females should have a selective advantage as they can distribute sperm evenly among the females they encounter, there are few invertebrate examples of the Coolidge effect. Here we present evidence for this effect in the burying beetle Nicrophorus vespilloides and examine the mechanism underlying the discrimination between familiar and novel mates. Burying beetles feed and reproduce on vertebrate carcasses, where they regularly encounter conspecifics. Males showed greater sexual interest in novel females (virgin or mated) than in females they had inseminated before. The application of identical cuticular extracts allowed us to experimentally create females with similar odours, and male responses to such females demonstrated that they use female cuticular patterns for discrimination. The chemical analysis of the cuticular profile revealed greater inter-individual variation in female than in male cuticular patterns, which might be due to greater selection on females to signal their individual identity.