Detecting Inter-Cusp and Inter-Tooth Wear Patterns in Rhinocerotids

Extant rhinos are the largest extant herbivores exhibiting dietary specialisations for both browse and grass. However, the adaptive value of the wear-induced tooth morphology in rhinos has not been widely studied, and data on individual cusp and tooth positions have rarely been published. We evaluated upper cheek dentition of browsing Diceros bicornis and Rhinoceros sondaicus, mixed-feeding R. unicornis and grazing Ceratotherium simum using an extended mesowear method adapted for rhinos. We included single cusp scoring (EM(R)-S) to investigate inter-cusp and inter-tooth wear patterns. In accordance with previous reports, general mesowear patterns in D. bicornis and R. sondaicus were attrition-dominated and C. simum abrasion-dominated, reflecting their respective diets. Mesowear patterns for R. unicornis were more attrition-dominated than anticipated by the grass-dominated diet, which may indicate a low intake of environmental abrasives. EM(R)-S increased differentiation power compared to classical mesowear, with significant inter-cusp and inter-tooth differences detected. In D. bicornis, the anterior cusp was consistently more abrasion-dominated than the posterior. Wear differences in cusp position may relate to morphological adaptations to dietary regimes. Heterogeneous occlusal surfaces may facilitate the comminution of heterogeneous browse, whereas uniform, broad grinding surfaces may enhance the comminution of physically more homogeneous grass. A negative tooth wear gradient was found in D. bicornis, R. sondaicus and R. unicornis, with wear patterns becoming less abrasion-dominated from premolars to molars. No such gradients were evident in C. simum which displayed a uniform wear pattern. In browsers, premolars may be exposed to higher relative grit loads, which may result in the development of wear gradients. The second premolar may also have a role in food cropping. In grazers, high absolute amounts of ingested abrasives may override other signals, leading to a uniform wear pattern and dental function along the tooth row, which could relate to the observed evolution towards homodonty.     

Coevolution of Tooth Crown Height and Diet in Oreodonts (Merycoidodontidae, Artiodactyla) Examined with Phylogenetically Independent Contrasts

The evolution of increased tooth crown height is considered to be an adaptation for coping with excessive rates of dental wear associated with abrasive herbivorous diets, such as grazing and(or high levels of exogenous grit (e.g. dust, sand, ash). Evolutionary trends in the crown heights of North American ungulates are grossly consistent with a transition from closed forests in the early Eocene to open grasslands in the late Miocene. However, the evolutionary proliferation of hypsodonty (high crowned teeth) in the early and middle Miocene occurs later than the apparent origin of open grassland habitats in North America. The paleoecology of species from the interval between the appearance of grasslands and the evolutionary proliferation of hypsodonty is critical to understanding the role of Cenozoic climate change in mammalian evolution. The paleodiets of late Eocene to middle Miocene oreodonts (Merycoidodontidae) were reconstructed by examining the relative facet development of molars (mesowear). A two-phase diet trend was discovered. Phase 1 suggests either an average reduction in the amount of exogenous grit from the late Eocene to early Oligocene or a decrease in fruit consumption related to the disappearance of more wooded habitats. Phase 2 is a gradual transition from early Oligocene low-abrasion browsing to high abrasion diets similar to mixed feeding and grazing in the Miocene. According to mesowear data, oreodont diets similar to those of modern grazers in terms of abrasion are not seen until the early Miocene (early Hemingfordian land mammal age). The coevolutionary relationship of molar crown height and diet, as represented by mesowear, was examined using phylogenetically independent contrasts. No significant coevolutionary relationship was found. In several instances, diet was found to shift over time despite morphological stasis (i.e. within a single species). These results do not clearly indicate that the overall trend of increasing dietary abrasion imposed sufficient selection to drive crown height evolution in oreodonts. Therefore, direct fossil evidence of dietary abrasion as a causal factor in the evolution of crown height, at least in this clade, is elusive.     

Comparing ungulate dietary proxies using discriminant function analysis

A variety of tooth‐wear and morphological dietary proxies have been proposed for ungulates. In turn, they have been applied to fossil specimens with the purpose of reconstructing the diets of extinct taxa. Although these dietary proxies have been used in isolation and in combination, a consistent set of statistical analyses has never been applied to all of the available datasets. The purpose of this study is to determine how well the most commonly used dietary proxies classify ungulates as browsers, grazers, and mixed feeders individually and in combination. Discriminant function analysis is applied to individual dietary proxies (hypsodonty, mesowear, microwear, and several cranial dietary proxies) and to combinations thereof to compare rates of successful dietary classification. In general, the tooth‐wear dietary proxies (mesowear and microwear) perform better than morphological dietary proxies, though none are strong proxies in isolation. The success rates of the cranial dietary proxies are not increased substantially when ruminants and bovids are analyzed separately, and significance among the three dietary guilds is reduced when controlling for phylogenetic relatedness. The combination of hypsodonty, mesowear, and microwear is found to have a high rate of successful dietary classification, but a combination of all commonly used proxies increases the success rate to 100%. In most cases, mixed feeders bear the greatest resemblance to browsers suggesting that a morphology intermediate to browsers and grazers may represent a fitness valley resulting from the inability to exploit both browse and graze efficiently. These results are important for future paleoecological studies and should be used as a guide for determining which dietary proxies are appropriate to the research question. J. Morphol., 2011. © 2011 Wiley‐Liss, Inc.     

The diet of <Emphasis Type="Italic">Metaschizotherium bavaricum</Emphasis> (Chalicotheriidae,Mammalia) from the MN 5 of Sandelzhausen (Germany) implied by the mesowear method

The genus Metaschizotherium is a Miocene member of the Schizotheriinae (Chalicotheriidae, Perissodactyla) and appears in the MN 5 of Southern Germany with the species M. bavaricum. The Chalicotheriidae have mostly been reconstructed as browsers. In this study, the mesowear method is applied to 11 upper premolars and molars of M. bavaricum from the Upper Freshwater Molasse locality of Sandelzhausen (MN 5). With this method the amount of abrasive and attritive dental wear is investigated and thus it provides a time-averaged signature of food abrasiveness of ungulates. Principal components analysis is performed on mesowear variables of M. bavaricum. This species is found to classify closest to extant mixed feeding ruminants. This indicates that the diet of M. bavaricum essentially included nonabrasive browse but also a certain amount of abrasive plant material. The composition of the diet of extant reference species indicates that this abrasiveness was most likely imposed by bark and whole branches. In the palaeoecological context of Sandelzhausen M. bavaricum occupied the dietary niche of a mixed feeder or an abrasion-dominated browser. A high degree of similarity is recognised between the mesowear patterns of M. bavaricum from Sandelzhausen and M. fraasi from the MN 6 of the Franconian Alb. This is interpreted as reflecting a similar dietary niche of the two species.      

Correlation of relative muzzle width and relative incisor width with dietary preference in ungulates

Qualitative observations suggest that grazing ungulates have relatively broader muzzles than browsing ones, and that grazers have incisors that are all of a similar size, as opposed to the large central and smaller lateral incisors seen in browsers. These differences may be correlated respectively with the need for grazing ungulates to maintain a large daily intake, or for browsing ungulates to forage selectively in a stand of vegetation. Quantitative examination of relative muzzle width and incisor width ratio in 95 species of living ungulates, correlated with seven different types of dietary preferences, substantiated these observations, although phylogenetic history may exhort a strong influence on morphological proportions. For example, equids have relatively narrower muzzles than grazing ruminants despite their less selective mode of feeding. The narrowest relative muzzle widths are not found in regular browsers, but in high level browsers and in mixed feeders in open habitats. Incisor width ratio can distinguish grazers from browsers, but can not be used to discriminate mixed feeders from other feeding types, and grazers appear to have incisors that are relatively broader overall than those of other dietary types, in correlation with their relatively broader muzzles.     

Paleoenvironment of Dryopithecus brancoi at Rudabánya, Hungary: evidence from dental meso- and micro-wear analyses of large vegetarian mammals

The environment of the hominoid Dryopithecus brancoi at Rudabánya (Late Miocene of Hungary) is reconstructed here using the dietary traits of fossil ruminants and equids. Two independent approaches, dental micro- and meso-wear analyses, are applied to a sample of 73 specimens representing three ruminants: Miotragocerus sp. (Bovidae), Lucentia aff. pierensis (Cervidae), Micromeryx flourensianus (Moschidae), and one equid, Hippotherium intrans (Equidae). The combination of meso- and micro-wear signatures provides both long- and short-term dietary signals, and through comparisons with extant species, the feeding styles of the fossil species are reconstructed. Both approaches categorize the cervid as an intermediate feeder engaged in both browsing and grazing. The bovid Miotragocerus sp. is depicted as a traditional browser. Although the dental meso-wear pattern of the moschid has affinities with intermediate feeders, its dental micro-wear pattern also indicates significant intake of fruits and seeds. Hippotherium intrans was not a grazer and its dental micro-wear pattern significantly differs from that of living browsers, which may suggest that the fossil equid was engaged both in grazing and browsing. However, the lack of extant equids which are pure browsers prevents any definitive judgment on the feeding habits of Hippotherium. Based on these dietary findings, the Rudabánya paleoenvironment is reconstructed as a dense forest. The presence of two intermediate feeders indicates some clearings within this forest; however the absence of grazers suggests that these clearings were most likely confined. To demonstrate the ecological diversity among the late Miocene hominoids in Europe, the diet and habitat of Dryopithecus brancoi and Ouranopithecus macedoniensis (Greece) are compared.     

The functional significance of the browser-grazer dichotomy in African ruminants

The allometric relationships for the fermentation rate of dry matter, the total energy concentration of volatile fatty acids (VFAs), the energy supplied from VFA production and the mass of the digesta contents within the rumen or caecum and proximal colon (hindgut) were used to test whether the digestive strategies of grazing and browsing African ruminants differ. The wet and dry mass of the contents of the rumen and hindgut were allometrically related to body mass (BM). These relationships did not differ between browsing and grazing ruminants. The fermentation rates in the rumen were strongly allometric and the intercepts of the relationships did not differ between browsers and grazers. The fermentation rates in the hindgut were not allometrically related to BM and did not differ between ruminants with different feeding habits. Likewise, the total energy concentration of the VFAs in the rumen and hindgut showed no allometric scaling and did not differ between browsing and grazing ruminants. The energy supplied by VFA production in both the rumen and hindgut of African ruminants scaled at around 0.8 with BM. Only in the case of the energy supplied by VFAs in the rumen were there significantly different intercepts for browsing and grazing ruminants. The energy supplied by VFA production in the rumen was inadequate to meet the energy requirements for maintenance of browsers and small grazers. The retention time of digesta in the alimentary tract was positively related to BM although there was no difference in the allometric relationships for grazers and browsers. The results of these analyses suggest that, after controlling for the effects of body mass, there is little difference in digestive strategy between African ruminants with different morphological adaptations of the gut.     

Differences in fecal particle size between free‐ranging and captive individuals of two browser species

Data from captive animals indicated that browsing (BR) ruminants have larger fecal particles—indicative of lesser chewing efficiency—than grazers (GR). To answer whether this reflects fundamental differences between the animal groups, or different reactions of basically similar organisms to diets fed in captivity, we compared mean fecal particle size (MPS) in a GR and a BR ruminant (aurox Bos primigenius taurus, giraffe Giraffa camelopardalis) and a GR and a BR hindgut fermenter (Przewalski's horse Equus ferus przewalskii, lowland tapir Tapirus terrestris), both from captivity and from the wild. As would be expected owing to a proportion of finely ground, pelleted feeds in captive diets, MPS was smaller in captive than free‐ranging GR. In contrast, MPS was drastically higher in captive than in free‐ranging BR of either digestion type. Thus, the difference in MPS between GR and BR was much more pronounced among captive than free‐ranging animals. The results indicate that BR teeth have adapted to their natural diet so that in the wild, they achieve a particle size reduction similar to that of GR. However, although GR teeth seem equally adapted to food ingested in captivity, the BR teeth seem less well suited to efficiently chew captive diets. In the case of ruminants, less efficient particle size reduction could contribute to potential clinical problems like “rumen blockage” and bezoar formation. Comparisons of MPS between free‐ranging and captive animals might offer indications for the physical suitability of zoo diets. Zoo Biol 27:70–77, 2008. © 2007 Wiley‐Liss, Inc.     

Food preferences and tooth wear in the sand gazelle (Gazella marica)

Food preferences of the sand gazelle (Gazella marica) from the Mahazat as-Sayd Protected Area in Saudi Arabia were evaluated using focal animal sampling in conjunction with an eco-morphological method examining two parameters of tooth wear, i.e., occlusal relief and cusp shape. Observations of live, free-ranging animals (n = 53) showed that sand gazelles generally consumed more grass (58.4%) than browse (41.6%). However, during the dry season, gazelles spent significantly more time browsing (51.0%) and less time grazing (49.0%) than under wet conditions (browsing: 17.6%; grazing: 82.4%). Thus, consistent with predictions, sand gazelles are intermediate feeders but shift towards browsing when grass is scarce. The mesowear signature of the sand gazelle is consistent with a grazing signal in other ruminants. In other words, the browse component of the diets of live animals was not reflected in the tooth wear. This could have occurred because browse is less abrasive than grass, but more likely because all food types are heavily abrasive in this dusty habitat. We conclude that the sand gazelle population in Mahazat as-Sayd encounters a highly abrasive diet, which has implications for their ability to meet nutritional demands.     

Microwear evidence for Plio-Pleistocene bovid diets from Makapansgat Limeworks Cave, South Africa

Makapansgat Limeworks Cave is a well-known Australopithecus africanus bearing locality that has spawned a considerable amount of paleoecological research because of its hominin component. Most recently, the paleoecology of this Plio-Pleistocene site has been studied by determining the diet and habitat of other extinct taxa, particularly the bovids. The diets of seven bovids (Aepyceros sp., Gazella vanhoepeni, Makapania broomi, Parmularius braini, Redunca darti, Tragelaphus sp. aff. T. angasii, and Tragelaphus pricei) have now been classified using taxonomic uniformitarianism, ecomorphology, stable carbon isotopes, and mesowear analysis. Here, dental microwear is applied to the same bovids for additional comparison and to further elucidate the strengths and weaknesses of each method. The different dietary proxy methods noted provide a temporal continuum, with genetic signals such as ecomorphology and taxonomic uniformitarianism indicating behavioral adaptations over geologic time, while nongenetic data such as stable carbon isotopes and mesowear reflect different aspects of average diet over extended portions of an animal's life, and dental microwear provides dietary snapshots.Microwear separated an extant baseline of ten bovid species into expected dietary categories and the Makapansgat bovids clearly fell into two groups with the same degree of separation as between extant grazers and browsers. The results indicate that a multidisciplinary approach produces a more accurate and robust reconstruction of past diets. In sum, the microwear analysis is in-line with the isotope and mesowear results, which suggest a stronger browsing component than either taxonomic uniformitarianism or ecomorphology imply.     

Ruminant diets and the Miocene extinction of European great apes

The successful evolutionary radiations of European hominoids and pliopithecoids came to an end during the Late Miocene. Using ruminant diets as environmental proxies, it becomes possible to detect variations in vegetation over time with the potential to explain fluctuations in primate diversity along a NW–SE European transect. Analysis shows that ruminants had diverse diets when primate diversity reached its peak, with more grazers in eastern Europe and more browsers farther west. After the drop in primate diversity, grazers accounted for a greater part of western and central European communities. Eastwards, the converse trend was evident with more browsing ruminants. These opposite trends indicate habitat loss and an increase in environmental uniformity that may have severely favoured the decline of primate diversity.     

Differences in selective reticulo-ruminal particle retention as a key factor in ruminant diversification

The measurement of passage rate is important for the concept of ruminant diversification. While supporters of Hofmann's 1989 feeding type classification claim that browsing ruminants have faster passage rates than grazing ruminants, other researchers consider the passage rate to depend on body size alone. To date, no convincing comparison of ruminant passage rates has been put forward. For comparative purposes, we suggest the use of the "selectivity factor", which is an expression of how much longer particles of a defined size (<2 mm) are retained in the ruminant digestive tract than fluids. From the limited data available, it seems that grazing ruminants display selectivity factors between 1.56 and 3.80, whereas browsers have a much narrower range of 1.14-1.80. This suggests that browsers are not able to selectively retain particles as long as grazers. Intake of browsers, on the other hand, may not be limited by physical fill of the forestomach to the same degree as in grazers. This result can explain several observations on the digestive physiology of browsers, some of which have been linked to a rumen bypass mechanism. We propose that the ability for selective particle retention is a key factor for understanding the physiological consequences of ruminant diversification.     

Phylogenetic topology mapped onto dietary ecospace reveals multiple pathways in the evolution of the herbivorous niche in African Bovidae

Understanding the evolutionary history of the herbivore niche within African bovids has traditionally relied on examining anatomical adaptations to diet, particularly those related to digestive strategy. More recently, mesowear and stable isotope analyses have been used to great effect to reconstruct dietary preferences. We use these dietary proxies to construct a morphology‐free dietary ecospace and examine the topology of the phylogenetic relationships of African bovids mapped onto this ecospace. The reconstructed dietary ecospace provides evidence for four distinct dietary classes: species with C3‐ or C4‐dominated diets that produce low or high occlusal relief, likely related to diets high or low in abrasives, respectively. We detected no evidence for a discrete mixed feeder category; the species often categorized as such represent the end members of groups of species with either C3‐ or C4‐ dominated diets. Our analysis reveals high variability within the C4 grazing ecospace, and phylogenetic evidence indicates at least two pathways to grazing, likely related to the abrasive qualities of ingested food, which may be determined by the moisture content or the height of consumed grasses. These different pathways probably contribute to the high diversity of African grazers, both today and in the fossil record. C3 browsers (non‐frugivores) also display a high degree of variation, but there are no species associated with highly abrasive diets and there is evidence for only a single evolutionary pathway. We find evidence for only one evolutionary route towards frugivory, which includes species with diets that produce both high and low occlusal reliefs. The cause of abrasive wear in frugivores may be related to grit and/or the hard parts of fruits, but this requires further examination.     

Molar proportions of volatile fatty acids in the gastrointestinal tract of East African wild ruminants

The molar proportions of seven individual VFA's were determined at select sites along the gastrointestinal tract of sixteen species of East African wild ruminants. The resulting data were statistically analyzed for species effect, and for effects due to major feeding groups (browsers, grazers, fresh grass grazers, etc.) and for body weight groups (5-750 kg animals). Present data suggest that body weight, rather than diet, is the more influential factor in reticulo-rumen fermentation rate, and in the molar proportion of fatty acids present. The molar proportions of VFA's observed in the mid and hindgut of these wild ruminants appeared more responsive to diet and body weight of the animal than did foregut VFA values.     

植食性哺乳动物牙齿磨痕分析方法简介及其在古食性恢复中的应用前景

通过研究古哺乳动物的食性来探讨哺乳动物演化与古生态环境变化之间的关系是目前古生物学研究领域的一个热点,而牙齿磨痕分析是恢复古食性和重建古生态环境的重要手段。牙齿磨痕(dental wear)分析包括微痕(microwear)分析和中痕(mesowear)分析,两种方法均强调食性与牙齿磨痕模式的严格对应,即不同食性的动物具有不同的牙齿磨痕特征模式。近年来,牙齿磨痕分析方法以其简单、快捷和高效等优点已被广泛应用于奇蹄类、偶蹄类、啮齿类、长鼻类和食肉类等哺乳动物的食性研究。但哺乳动物的食性和摄食习性比较复杂,很可能会影响微痕和中痕分析对食性的分辨率。所以,为了获得更加详细的古食性信息和更高的食性分辨率,一方面要对微痕和中痕分析方法进行改进,增添稳定并具有食性识别意义的观测变量,另一方面,需要同时结合微痕和中痕分析,从而获得更加全面的食性信息。虽然牙齿磨痕分析目前主要应用于植食性哺乳动物的食性研究,但其原理对哺乳动物的其它类群也是适用的,随着磨痕分析方法的不断改进和其它类群磨痕数据库的建立,未来的牙齿磨痕分析将可以恢复更多类群的古食性,从而可以更加全面和准确地揭示古食性与古环境信息。     

An in vitro evaluation of browser and grazer fermentation efficiency and microbiota using European moose spring and summer foods

Evolutionary morphological and physiological differences between browsers and grazers contribute to species‐specific digestion efficiency of food resources. Rumen microbial community structure of browsers is supposedly adapted to characteristic nutrient composition of the diet source. If this assumption is correct, domesticated ruminants, or grazers, are poor model animals for assessing the nutritional value of food consumed by browsing game species. In this study, typical spring and summer foods of the European moose (Alces alces) were combined with rumen fluid collected from both dairy cows (Bos taurus) and from moose, with the aim of comparing fermentation efficiency and microbial community composition. The nutritional value of the food resources was characterized by chemical analysis and advanced in vitro measurements. The study also addressed whether or not feed evaluation based on in vitro techniques with cattle rumen fluid as inoculum could be a practical alternative when evaluating the nutritional value of plants consumed by wild browsers. Our results suggest that the fermentation characteristics of moose spring and summer food are partly host‐specific and related to the contribution of the bacterial phyla Firmicutes and Bacteriodetes to the rumen microbial community. Host‐specific adaptations of the ruminal microbial community structure could be explained from the evolutionary adaptations related to feeding habitats and morphophysiological differences between browsers and grazers. However, the observed overall differences in microbial community structure could not be related to ruminal digestion parameters measured in vitro. The in vitro evaluation of digestion efficiency reveals that equal amounts of methane were produced across all feed samples regardless of whether the ruminal fluid was from moose or dairy cow. The results of this study suggested that the nutritional value of browsers' spring and summer food can be predicted using rumen fluid from domesticated grazers as inoculum in in vitro assessments of extent of digestion when excluding samples of the white water lily root, but not of fermentation characteristics as indicated by the proportions of individual fermentation fatty acids to the total of volatile fatty acids.     

Adaptation of ruminants to browse and grass diets: are anatomical-based browser-grazer interpretations valid?

As a result of pioneering work of Hofmann (1973, 1989), nutritional ecologists classify ruminants into three feeding-type categories: browsers (concentrate feeders), grazers, and intermediate or mixed feeders. Hofmann proposed that these feeding types result from evolutionary adaptations in the anatomy of the digestive system and that one consequence is shorter retention of the digesta in the rumen of browsers, and thus a decreased efficiency of fiber digestion relative to that of grazers. We examined the hypotheses that (1) fiber digestion of browsers is lower than that of grazers, (2) salivary gland size is larger in all browsers than in grazers, (3) the browser's larger salivary glands produce larger volumes of thin serous saliva than those of grazers, and (4) thus, browsers have higher liquid passage rates than do grazers. We found that the extent of fiber digestion is not significantly different between browsers and grazers, although fiber digestion is positively related to herbivore size. In general, salivary gland size is approximately 4 times larger in browsers than grazers, but some browsers (e.g., greater kudu) have small, grazer-sized salivary glands. Resting (non-feeding or ruminating) saliva flow rates of mule deer (browser) and domestic sheep and cattle (grazers) were not significantly different from each other. Finally, ruminal liquid flow rates were not different between feeding types. We conclude that many of Hofmann's nutritional and physiological interpretations of anatomical differences amongst ruminants are not supportable.     

Higher masseter muscle mass in grazing than in browsing ruminants

Using cranioskeletal measurements, several studies have generated evidence that grazing ruminants have a more pronounced mastication apparatus, in terms of muscle insertion areas and protuberances, than browsing ruminants, with the resulting hypothesis that grazers should have larger, heavier chewing muscles than browsers. However, the only investigation of this so far [Axmacher and Hofmann (J Zool 215:463-473, 1988)] did not find differences between ruminant feeding types in the masseter muscle mass of 22 species. Here, we expand the dataset to 48 ruminant species. Regardless of phylogenetic control in the statistical treatment, there was a significant positive correlation of body mass and masseter mass, and also a significant association between percent grass in the natural diet and masseter mass. The results support the concept that ruminant species that ingest more grass have relatively larger masseter muscles, possibly indicating an increased requirement to overcome the resistance of grass forage. The comparative chewing resistance of different forage classes may represent a rewarding field of ecophysiological research.     

The ecology and evolution of tooth wear in red deer and moose

Differences in body size and diet type (browser–grazer continuum) have formed functional traits of ruminants, including tooth design. Grazers and mixed-feeders eat a more fibrous diet than browsers, which arguably increase tooth wear. Tooth wear has also been suggested to increase with body size. Moreover, for species with large distribution ranges, different populations may be exposed to very different ecological factors affecting diet and thus tooth wear rates. Therefore, evolutionary history and contemporary ecological conditions, both operating through diet, may be important for patterns of tooth wear. Here, we compare inter- and intraspecific rates of tooth wear in multiple populations of one large browser (moose Alces alces ) and one mixed-feeder (red deer Cervus elaphus ) covering the main distribution range of each species in Norway. We found that the mixed-feeding red deer wore teeth faster than the larger and browsing moose, suggesting that feeding-type was more important than body size for patterns of wear. There was substantial spatial variation in tooth wear rates, but the inter-specific difference in wear was consistent. Molar wear rates, but not incisors wear rates, in the browser were less variable between populations than in the mixed-feeder. There was no close link between incisor and molar wear rates at the population level. Our findings are consistent with the view that both evolution related to diet type and current ecological conditions (being proxies for within-species variation in diet quality) are important for patterns of tooth wear.     

Effects of grass and browse consumption on the winter mass dynamics of elk

Much of the research into herbivore ecology and evolution has focused on patterns and mechanisms of niche partitioning in the diets of specialized grazers and browsers, but the significance of the balance of grazing and browsing within intermediate feeders present unique questions that have received less attention. We explored the nutritional effects of varying the balance of grass and browse in the winter diets of elk (Cervus elaphus). We compiled published data from three similar studies that monitored the mass dynamics of captive elk fed diets of pure grass, pure browse or 1 of 14 mixed diets in winter feeding trials. Elk lost mass (up to 22% of initial body weight) in 29 of 33 feeding trials, similar to wild elk in winter. We used regression models of mass dynamics, considering the linear, quadratic or logarithmic effects of the proportion of the diet that was grass (≈1 − proportion of the diet that was browse) and the additive and interactive effects of nitrogen intake. Diet composition had strong effects on mass dynamics, and all models explained ≥73% of the variation (adjusted r ²) in mass dynamics. Nitrogen intake had uniformly positive effects on mass balance, and increasing grass intake caused improved maintenance of body mass up to a point, but further increases in the proportion of grass in the diet had neutral or negative effects on body mass. Overall, the data suggest that elk are adapted to consuming mixed diets. Nonetheless, data on the foraging behavior and diet selection of wild elk in many populations show that elk often consume grass-dominated diets and sometimes consume browse-dominated diets, but rarely consume mixed diets. Physiological adaptations to mixed diets may place unique spatio-temporal constraints on diet selection in intermediate feeders and impose large penalties for a sub-optimal balance of grazing and browsing.