Mammals inhabit all types of environments and have evolved chewing systems capable of processing a huge variety of structurally diverse food components. Surface textures of cheek teeth should thus reflect the mechanisms of wear as well as the functional traits involved. We employed surface textures parameters from ISO/DIS 25178 and scale-sensitive fractal analysis (SSFA) to quantify dental wear in herbivorous mammals at the level of an individual wear enamel facet. We evaluated cheek dentitions of two grazing ungulates: the Blue Wildebeest (Connochaetes taurinus) and the Grevy's Zebra (Equus grevyi). Both inhabit the east African grassland savanna habitat, but they belong to fundamentally different taxonomic units. We tested the hypothesis that the foregut fermenting wildebeest and the hindgut fermenting zebra show functional traits in their dentitions that relate to their specific mode of food-composition processing and digestion. In general, surface texture parameters from SSFA as well as ISO/DIS 25178 indicated that individual enamel ridges acting as crushing blades and individual wear facets of upper cheek teeth are significantly different in surface textures in the zebra when compared with the wildebeest. We interpreted the complexity and anisotropy signals to be clearly related to the brittle, dry grass component in the diet of the zebra, unlike the wildebeest, which ingests a more heterogeneous diet including fresh grass and herbs. Thus, SSFA and ISO parameters allow distinctions within the subtle dietary strategies that evolved in herbivorous ungulates with fundamentally different systematic affinities but which exploit a similar dietary niche.

(2008). The morphophysiological adaptations of browsing and grazing mammals. In: Gordon, I J [et al.]. The ecology of browsing and grazing. Berlin, 47-88. Postprint available at: http://www.zora.uzh.ch Posted at the Zurich Open Repository and Archive, University of Zurich. http://www.zora.uzh.ch Originally published at: Gordon, I J [et al.] 2008. The ecology of browsing and grazing. Berlin, 47-88. The morphophysiological adaptations of browsing and grazing mammals AbstractThere has been a continous debate whether there are fundametal morphophysiological differences in the ingestive apparatus (head, teeth) and the digestive tract between browsing and grazing herbivores. A particular characteristic of this debate appears to be that while there is a wealth of publications on such potential differences, the supposed undelying differences between browse and grass have rarely been analysed quantitatively. In this chapter, we first review the actual state of knowledge on those properties of browse and grass that appear relevant for the ingestive and digestive process, and then deduct hypotheses as to how one would assume that browsers and grazers differ due to these characteristics. We address the methodological issues involved in actually testing these hypotheses, with emphasis on the influence of body mass and phylogenetic descent. Finally, we present a literature compilation of statistical tests of differences between the feeding-types. Although in general, the published tests support many hypothesized differences, there is still both a lack of comparative data, and a lack of analyses with phylogenetic control, on different taxonomic levels. However, the published material appears to indicate that convergent evolutionary adaptations of browsing and grazing herbivores to their diet represent a rewarding area of research. The Morphophysiological Adaptations of Browsing and Grazing MammalsMarcus Clauss, Thomas Kaiser, and Jürgen Hummel IntroductionAnimals represent adaptations to particular ecological niches they occupy or once occupied.Studying the correlation between a given set of characteristics of an ecological niche and the morphological and physiological adaptations of organisms to these characteristics is one of the most basic approaches to comparative biology, and has fuelled scientific interest for generations (Gould 2002). However, current scientific standards cannot be met by mere descriptions of both the characteristics of the niche and the organism, and a (hypothetical) intuitive explanation for the adaptive relevance of the latter; the presence or absence of a characteristic must be demonstrated in sound statistical terms (Hagen 2003) 1 ideally supported by experimental data (from in vivo, in vitro, or model assays) on its adaptive relevance.In this chapter, we adopt an approach that first presents the relevant characteristics of the ecological niche of the 'grazer' (GR) and of the 'browser' (BR), outlines hypotheses based on these characteristics, and finally addresses examples where such hypotheses h...

1. The evolution of high-crowned teeth or hypsodonty in herbivorous mammals is widely interpreted as a species-specific adaptation to increasingly wear-inducing diets and environments at evolutionary time scales, with internal abrasives (such as phytoliths in grasses) and/or external abrasives (such as dust or grit) as putative causative factors. The mesowear score (MS) instead describes tooth wear experienced by individual animals during their lifetime. 2. Under the assumption that the abrasiveness that causes the MS in individuals is the same abrasiveness to which species adapted by evolving hypsodonty, one would expect a close correlation between the MS and the hypsodonty index (HI). Alternatively, if these two measures reflect different aspects of wear, one would expect differences in the way that proxies of diet or environment/climate correlate with each parameter. 3. In order to test these hypotheses, we collated a dataset on the HI, MS, percentage of grass in the natural diet (%grass), habitat (open, intermediate, closed) and annual precipitation (PREC) in extant mammalian herbivores. The availability of a quantitative MS constrained the dataset to 75 species. Data were analysed with and without phylogenetic generalized least squares (PGLS). 4. Correlations with PREC were stronger for HI than for MS, whereas correlations with %grass were similar for HI and MS. Habitat had a significant influence on the relationship with %grass for HI but not for MS. Habitat also had a significant influence on the relationship between HI and MS. MS improved the predictive power of HI for %grass, but not for PREC. 5. These results suggest that while the MS indicates predominantly the wear effect of the diet (internal abrasives), HI represents an adaptation to a wear effect that comprises both diet and environment (external abrasives). The additional environmental wear effect must reduce tooth height without causing macroscopic changes in tooth facet development as described by the MS. 6. The most parsimonious explanation for the apparent discrepancy between HI and MS is that external abrasives of very fine particle size play a major role in naturally occurring tooth wear. The experimental testing of this hypothesis will enhance our understanding of the processes involved in tooth wear. ABSTRACT1. The evolution of high-crowned teeth or hypsodonty in herbivorous mammals is widely interpreted as a species-specific adaptation to increasingly wear-inducing diets and environments at evolutionary time scales, with internal abrasives (such as phytoliths in grasses) and/or external abrasives (such as dust or grit) as putative causative factors. The mesowear score (MS) instead describes tooth wear experienced by individual animals during their lifetime. 2. Under the assumption that the abrasiveness that causes the MS in individuals is the same abrasiveness to which species adapted by evolving hypsodonty, one would expect a close correlation between the MS and the hypsodonty index (HI). Alternatively, if these two measures...

No abstract

Dental microwear and 3D surface texture analyses are useful in reconstructing herbivore diets, with scratches usually interpreted as indicators of grass dominated diets and pits as indicators of browse. We conducted feeding experiments with four groups of rabbits (Oryctolagus cuniculus) each fed a different uniform, pelleted diet (lucerne, lucerne & oats, grass & oats, grass). The lowest silica content was measured in the lucerne and the highest in the grass diet. After 25 weeks of exposure to the diets, dental castings were made of the rabbit's lower molars. Occlusal surfaces were then investigated using dental microwear and 3D areal surface texture analysis. In terms of traditional microwear, we found our hypothesis supported, as the grass group showed a high proportion of (long) “scratches” and the lucerne group a high proportion of “pits”. Regardless of the uniform diets, variability of microwear and surface textures was higher when silica content was low. A high variability in microwear and texture analysis thus need not represent dietary diversity, but can also be related to a uniform, low-abrasion diet. The uniformity or variability of microwear/texture analysis results thus might represent varying degrees of abrasion and attrition rather than a variety of diet items per se.

No abstract

The mesowear method evaluates the wear patterns of herbivore cheek teeth by 2 visually evaluating the facet development of the occlusal surfaces. It thus allows 3 classification of most herbivorous ungulates into browsers, grazers or intermediate 4 feeders, due to the fact that in grazers, tooth wear is characterized by a 5 comparatively high degree of abrasion, most probably due to the presence of 6 silicacious phytoliths in grasses, a higher amount of dust and grit adhering to their 7 forage, or both. It has been suggested that excessive tooth wear could be a 8 particularly limiting factor in the husbandry of captive large browsing species, and 9 major tooth wear was demonstrated in captive as compared to free-ranging giraffe. If 10 this increased tooth wear in captivity was an effect of feeding type and diets fed, then 11 it would be expected that other browsing species are affected in a similar manner. In 12 order to test this hypothesis, we investigated the dental mesowear pattern in captive 13 individuals of 19 ruminant species and compared the results to data on free-ranging 14 animals. Compared to free-ranging populations, captive browsers show a 15 significantly more abrasion-dominated tooth wear signal. The reverse applies to 16 captive grazers, which tend to show a less abrasion-dominated wear in captivity. 17Captive ruminants were generally more homogenous in their wear signature than 18 free-ranging ruminants. If grit contamination in the natural habitat is a major cause of 19 dental wear in grazers, then diets in captivity, although similar in botanical 20 composition, most likely contain less abrasives due to feeding hygiene. If dental wear 21 is one of the major factors limiting longevity, then captive grazers should achieve 22 longer lifespans than both captive browsers and free-ranging grazers. In particular 23 with respect to browsers, the results suggest that captive feeding regimes could be 24 improved.

SignificanceIngesta leave characteristic wear features on the tooth surface, which enable us to reconstruct the diet of extant and fossil vertebrates. However, whether dental wear is caused by internal (phytoliths) or external (mineral dust) silicate abrasives is controversially debated in paleoanthropology and biology. To assess this, we fed guinea pigs plant forages of increasing silica content (lucerne < grass < bamboo) without any external abrasives, both in fresh and dried state. Abrasiveness and enamel surface wear increased with higher forage phytolith content. Additionally, water loss altered plant material properties. Dental wear of fresh grass feeding was similar to lucerne browsing, while dried grass caused more grazer-like wear. Fresh grass grazing could be confounded with browsing, being a major pitfall for paleodietary reconstructions.