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Dietary reconstruction in vertebrates often relies on dental wear-based proxies. Although these proxies are widely applied, the contributions of physical and mechanical processes leading to meso- and microwear are still unclear. We tested their correlation using sheep (Ovis aries, n = 39) fed diets of varying abrasiveness for 17 months as a model. Volumetric crown tissue loss, mesowear change and dental microwear texture analysis (DMTA) were all applied to the same teeth. We hereby correlate: (i) 46 DMTA parameters with each other, for the maxillary molars (M1, M2, M3), and the second mandibular molar (m2); (ii) 10 mesowear variables to each other and to DMTA for M1, M2, M3 and m2; and (iii) volumetric crown tissue loss to mesowear and DMTA for M2. As expected, many DMTA parameters correlated strongly with each other, supporting the application of reduced parameter sets in future studies. Correlation results showed only few DMTA parameters correlated with volumetric tissue change and even less so with mesowear variables, with no correlation between mesowear and volumetric tissue change. These findings caution against interpreting DMTA and mesowear patterns in terms of actual tissue removal until these dental wear processes can be better understood at microscopic and macroscopic levels.

Dental microwear texture analysis (DMTA) is widely used for diet inferences in extant and extinct vertebrates. Often, a reference tooth position is analysed in extant specimens, while isolated teeth are lumped together in fossil datasets. It is therefore important to test whether dental microwear texture (DMT) is tooth position specific and, if so, what causes the differences in wear. Here, we present results from controlled feeding experiments with 72 guinea pigs, which received either fresh or dried natural plant diets of different phytolith content (lucerne, grass, bamboo) or pelleted diets with and without mineral abrasives (frequently encountered by herbivorous mammals in natural habitats). We tested for gradients in dental microwear texture along the upper cheek tooth row. Regardless of abrasive content, guinea pigs on pelleted diets displayed an increase in surface roughness along the tooth row, indicating that posterior tooth positions experience more wear compared with anterior teeth. Guinea pigs feedings on plants of low phytolith content and low abrasiveness (fresh and dry lucerne, fresh grass) showed almost no DMT differences between tooth positions, while individuals feeding on more abrasive plants (dry grass, fresh and dry bamboo) showed a gradient of decreasing surface roughness along the tooth row. We suggest that plant feeding involves continuous intake and comminution by grinding, resulting in posterior tooth positions mainly processing food already partly comminuted and moistened. Pelleted diets require crushing, which exerts higher loads, especially on posterior tooth positions, where bite forces are highest. These differences in chewing behaviour result in opposing wear gradients for plant versus pelleted diets.

Sharks are apex-predators that play an important role in past and present aquatic food webs. However, their diet - especially in extinct species - is often not well constrained. Dental microwear texture analysis (DMTA) has been successfully applied to reconstruct diet and feeding behaviours of different aquatic and terrestrial vertebrates. However, unlike in mammals, food-to-tooth contact in sharks is rather limited because only larger prey is manipulated before swallowing. Together with a fast tooth replacement rate, this reduces wear on individual teeth. Here, we present an explorative study of dental microwear texture on extant and extinct sharks to test whether ante-mortem wear is related to ingested diet or habitat preferences and resistant to post-mortem alteration processes. Shark teeth from 24 modern species and 12 fossil species from different localities were measured. As an additional comparison, extant shark teeth of Carcharhinus plumbeus were tumbled in sedimentwater suspensions to simulate post-mortem mechanical alteration by sediment transport. Only three of the twelve extant shark species with three or more specimens had significantly different dental surface textures. Furthermore, no clear relation between food or habitat preferences and ante-mortem dental wear features was detected for this sample set. Tumbling modern shark teeth with siliciclastic sediment of four different grain size fractions led to increasing complexity of the dental surface. Fossil specimens resemble these experimentally altered shark teeth more in complexity and roughness. Thus, fossil shark teeth seem to display either very different (e.g. harder) diet-related wear or a strong degree of post-mortem alteration. Based on our restricted sample size, dental wear of shark teeth does overall not seem to simply reflect dietary differences; hence, it is difficult to use DMTA as reliable dietary reconstruction, in either extant nor extinct sharks.

A herbivore's diet can affect its teeth by causing different types of wear. Browsers typically have sharper, higher cusps, while grazers show lower, blunter cusps, presumably due to the more abrasive nature of their diet. On the macroscopic scale, this allows the reconstruction of herbivore diets based on the shape of the tooth's profile, using the mesowear method. However, the timeframe involved in constituting a stable dietary signal represented by mesowear has not been precisely defined. To obtain a more precise delimitation of this timeframe, sheep (Ovis aries, n = 39) were fed pelleted diets containing external abrasives of different sizes (⌀ 4 μm, ⌀ 50 μm, and ⌀ 130 μm), and concentrations (4%, or 8%) for 17 months. Mesowear was scored on the skulls at the end of the experiment, as well as on computer-tomographic scans and on resin casts of the teeth taken at different timepoints along the experiment. These datasets were compared, and changes in score were calculated using the CT and cast data from the beginning and end of the experiment. Overall, even though a visual trend appeared of more wear on higher abrasion diets, it was of an extremely small magnitude, and the dietary effect on the mesowear scores or the change of the mesowear scores in these animals was never significant. This leads us to conclude that, at least in small ruminants, mesowear is more of a general signal than a seasonal one, and needs to be considered as such for tooth wear-based palaeodietary reconstructions. Experiments with natural forages are required to corroborate this conclusion.

A new study by Fraser et al (2018) urges the use of phylogenetic comparative methods, whenever possible, in analyses of mammalian tooth wear. We are concerned about this for two reasons. First, this recommendation may mislead the research community into thinking that phylogenetic signal is an artifact of some sort rather than a fundamental outcome of the evolutionary process. Secondly, this recommendation may set a precedent for editors and reviewers to enforce phylogenetic adjustment where it may unnecessarily weaken or even directionally alter the results, shifting the emphasis of analysis from common patterns manifested by large clades to rare cases.

Tooth wear induced by abrasive particles is a key process affecting dental function and life expectancy in mammals. Abrasive particles may be plant endogenous opal phytoliths, exogene wind-blown quartz dust or rain borne mineral particles ingested by mammals. Nano-indentation hardness of abrasive particles and dental tissues is a significant yet not fully established parameter of this tribological system. We provide consistent nano-indentation hardness data for some of the major antagonists in the dental tribosystem (tooth enamel, tooth dentine and opaline phytoliths from silica controlled cultivation). All indentation data were gathered from native tissues under stable and controlled conditions and thus maximize comparability to natural systems. Here we show that native (hydrated) wild boar enamel exceeds any hardness measures known for dry herbivore tooth enamel by at least 3 GPa. The native tooth enamel is not necessarily softer then environmental quartz grit, although there is little overlap. The native hardness of the tooth enamel exceeds that of any silica phytolith hardness recently published. Further, we find that native reed phytoliths equal native suine dentine in hardness, but does not exceed native suine enamel. We also find that native suine enamel is significantly harder than dry enamel and dry phytoliths are harder than native phytoliths. Our data challenge the claim that the culprit of tooth wear may be the food we chew, but suggest instead that wear may relates more to exogenous than endogenous abrasives.

Grasses are an important part of the forage of many herbivorous mammals and their phytoliths have long been regarded as the most important agent of tooth wear. Recent work has challenged this “paradigm” in finding evidence 1. of native phytoliths to be much softer then tooth enamel and 2. indicating, that phytolith hardness is highly variable, 3. prone to methodology and 4. not easy to be related to habitat conditions. We conduct controlled silica-cultivations measuring SiO2 content in the common forage grass Themeda triandra. Phytoliths are extracted natively, and nano-indentation values are measured. Phytolith hardness in Themeda triandra is found to be independent of silicate availability in the substrate. We further investigate the phytolith ultrastructure of Hordeum vulgare phytoliths. Phytoliths are shown to be an anisotropic composite of at least 3 components, silica bodies, inter-body matrix (both mineralised) and globular inclusions (likely non-mineralised). It can be argued, that indentation will be largely influenced by the heterogeneity of the structure and thus nano-indentation measurements will largely reflect the matrix and its mechanical properties but not necessarily the silicate bodies, which make up the vast majority of a phytolith.

External abrasives ingested along with the herbivore diet are considered main contributors to dental wear, though how the different sizes and concentrations of these abrasives influence wear remains unclear. Dental microwear texture analysis (DMTA) is an established method for dietary reconstruction which describes a tooth9s surface topography on a micrometre scale. The method has yielded conflicting results as to the effect of external abrasives. In the present study, a feeding experiment was performed on sheep (Ovis aries) fed seven diets of different abrasiveness. Our aim was to discern the individual effects of size (4, 50 and 130 µm) and concentration (0%, 4% and 8% of dry matter) of abrasives on dental wear, applying DMTA to four tooth positions. Microwear textures differed between individual teeth, but surprisingly, showed no gradient along the molar tooth row, and the strongest differentiation of experimental groups was achieved when combining data of all maxillary molars. Overall, a pattern of increasing height, volume and complexity of the tooth9s microscopic surface appeared with increasing size of dietary abrasives, and when compared with the control, the small abrasive diets showed a polishing effect. The results indicate that the size of dietary abrasives is more important for dental microwear texture traces than their concentration, and that different sizes can have opposing effects on the dietary signal. The latter finding possibly explains conflicting evidence from previous experimental DMTA applications. Further exploration is required to understand whether and how microscopic traces created by abrasives translate quantitatively to tissue loss.

There is an ongoing discourse about whether or not external abrasives influence the microscopic wear in herbivore teeth, including a statement that “dust does not matter”. We submitted the maxillary and mandibular second molar of 28 goats (Capra aegagrus hircus) to dental microwear texture analysis (DMTA). The study animals were divided into four groups, which received diets of increasing phytolith-based abrasiveness (L: lucerne based pellets, very low phytolith abrasion diet, acting as control; G: grass-based pellets, medium abrasive phytolith diet; GR: grass and rice husk pellets, high abrasion phytolith diet), or a diet with added external abrasives (GRS: the GR diet with addition of large-grained sand as source of external abrasives), for 6 months. Generally, the three diets without sand (L, G, GR) were ranked in the expected sequence by DMTA parameters. For some parameters, in particular those relating to area, complexity, peak sharpness and plateau size of the enamel surface, diet GRS differed from diet GR, whereas the two were not distinct in other parameters, mostly relating to direction, height or volume. Additionally, diets triggered different patterns on the maxillary or mandibular molar for several parameters. For some DMTA parameters, the range covered in the experimental goats corresponded to that observed in free-ranging browsing giraffe (Giraffa camelopardalis) and grazing wildebeest (Connochaetes taurinus), with goats receiving diet L resembling the giraffe, while the other diets G, GR and GRS more resembled the wildebeest. However, the previously reported mesowear in the goats did not cover the browser-grazer range, and DMTA measurements showed no correlation to the development of mesowear scores or measures of absolute tissue loss in the same animals. In summary, the results confirm DMTA measurements as generally recording properties related to both internal and external abrasives, but also indicate that our understanding of how DMTA characteristics actually originate, and how they relate to other proxies of dental wear, is still very limited.

Food processing wears down teeth, thus affecting tooth functionality and evolutionary success. Other than intrinsic silica phytoliths, extrinsic mineral dust/grit adhering to plants causes tooth wear in mammalian herbivores. Dental microwear texture analysis (DMTA) is widely applied to infer diet from microscopic dental wear traces. The relationship between external abrasives and dental microwear texture (DMT) formation remains elusive. Feeding experiments with sheep have shown negligible effects of dust-laden grass and browse, suggesting that intrinsic properties of plants are more important. Here, we explore the effect of clay- to sand-sized mineral abrasives (quartz, volcanic ash, loess, kaolin) on DMT in a controlled feeding experiment with guinea pigs. By adding 1, 4, 5, or 8% mineral abrasives to a pelleted base diet, we test for the effect of particle size, shape, and amount on DMT. Wear by fine-grained quartz (>5/ abrasive-free control diet. Fine silt-sized quartz (∼5 μm) results in higher surface anisotropy and lower roughness (polishing effect). Coarse-grained volcanic ash leads to significantly higher complexity, while fine sands (130 to 166 μm) result in significantly higher roughness. Complexity and roughness values exceed those from feeding experiments with guinea pigs who received plants with different phytolith content. Our results highlight that large (>95-μm) external silicate abrasives lead to distinct microscopic wear with higher roughness and complexity than caused bymineral abrasive-free herbivorous diets. Hence, high loads of mineral dust and grit in natural diets might be identified by DMTA, also in the fossil record.

One reason for the mammalian clade's success is the evolutionary diversity of their teeth. In herbivores, this is represented by high-crowned teeth evolved to compensate for wear caused by dietary abrasives like phytoliths and grit. Exactly how dietary abrasives wear teeth is still not understood completely. We fed four different pelleted diets of increasing abrasiveness (L: Lucerne; G: grass; GR: grass and rice husks; GRS: grass, rice husks, and sand) to four groups of a total of 28 adult goats, all with completely erupted third molars, over a six-month period. Tooth morphology was captured by medical computed tomography scans at the beginning and end of the controlled feeding experiment, and separation lines between the crown and root segments were defined in the upper right second molar (M2), to gauge absolute wear. Using bootstrapping, significant differences in volume loss between diets L/G and GR/GRS were detected. A small but nevertheless consistent volume gain was noted in the roots, and there was a significant, positive correlation between crown volume loss and root volume gain. This growth could possibly be attributed to the well-known process of cementum deposition and its relation with a putative feedback mechanism, in place to attenuate wear caused by abrasive diets.

Dental mesowear is applied as a proxy to determine the general diet of mammalian herbivores based on tooth-cusp shape and occlusal relief. Low, blunt cusps are considered typical of grazers and high, sharp cusps typical of browsers. However, how internal or external abrasives impact mesowear, and the time frame the wear signature takes to develop, still need to be explored. Four different pelleted diets of increasing abrasiveness (lucerne, grass, grass and rice husks, and grass, rice husks and sand) were fed to four groups of a total of 28 adult goats in a controlled feeding experiment over a 6-month period. Tooth morphology was captured by medical CT scans at the beginning and end of the experiment. These scans, as well as the crania obtained post mortem, were scored using the mesowear method. Comparisons between diet groups showed few significant differences after 6 months, irrespective of whether CT scans or the real teeth were scored. Only when assessing the difference in signal between the beginning and the end of the experiment did relevant, significant diet-specific effects emerge. Diets containing lower phytolith content caused a more pronounced change in mesowear towards sharper cusps/higher reliefs, while the feed containing sand did not result in more extreme changes in mesowear when compared with the same feed without sand. Our experiment suggests that the formation of a stable and hence reliable mesowear signal requires more time to develop than 6 months.

Reasons for performing study: Captive breeding has played a crucial role in the conservation of threatened equid species. Grazing ruminants and rhinoceros in captivity have less abrasion-dominated tooth wear than their free-ranging conspecifics, with potential negative consequences for their health. However, a similar study on wild equids in captivity is missing. Objectives: The aim was to establish if different tooth wear patterns are exhibited by free-ranging and captive equids. Study design: Cross-sectional study of museum specimens comparing free-ranging and captive equids. Methods: Dental casts of maxillary cheek teeth of 228 museum specimens (122 from free-ranging and 106 from captive individuals) of 7 wild equid species were analysed using the extended mesowear method. Although teeth showing specific abnormalities were not scored, the presence of focal overgrowths (hooks) of the rostral premolars (106, 206) was recorded. Results: Captive Equus ferus przewalskii, E. grevyi, E. hemionus, E. quagga boehmi and E. zebra hartmannae have less abrasion-dominated tooth wear on their premolars than their free-ranging conspecifics (P

Dental mesowear is applied as a proxy to determine the general diet of mammalian herbivores based on tooth-cusp shape and occlusal relief. Low, blunt cusps are considered typical of grazers and high, sharp cusps typical of browsers. However, how internal or external abrasives impact mesowear, and the time frame the wear signature takes to develop, still need to be explored. Four different pelleted diets of increasing abrasiveness (lucerne, grass, grass and rice husks, and grass, rice husks and sand) were fed to four groups of a total of 28 adult goats in a controlled feeding experiment over a 6-month period. Tooth morphology was captured by medical CT scans at the beginning and end of the experiment. These scans, as well as the crania obtained post mortem, were scored using the mesowear method. Comparisons between diet groups showed few significant differences after 6 months, irrespective of whether CT scans or the real teeth were scored. Only when assessing the difference in signal between the beginning and the end of the experiment did relevant, significant diet-specific effects emerge. Diets containing lower phytolith content caused a more pronounced change in mesowear towards sharper cusps/higher reliefs, while the feed containing sand did not result in more extreme changes in mesowear when compared with the same feed without sand. Our experiment suggests that the formation of a stable and hence reliable mesowear signal requires more time to develop than 6 months.

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.

Dietary traits of nine extant Equus species (E. africanus, E. ferus, E. grevyi, E. hart- mannae, E. hemionus, E. khur, E. kiang, E. quagga and E. zebra) were reconstructed and ranked in a continuum reflecting the relative proportions of abrasion and attrition. 2. In order to match them automatically with climatic data, localities were refer- enced within a 2° worldwide grid system using geographical information system software. The mesowear score was used as a proxy variable to test the covariance of diet with mean annual precipitation, evapotranspiration, mean annual climatic water balance and mean annual temperature. 3. Seven of the nine equid species presented mesowear signatures, indicating a grass-dominated diet, and we found interspecific as well as intraspecific variability. Non-gramineous food components shifted the equilibrium towards the attrition- dominated end in E. africanus and E. khur, indicating an intermediate diet. Especially in E. zebra, additional sources of exogenous abrasives (grit and soil) might increase dietary abrasiveness. In E. quagga, dietary flexibility and the capability to cope with increased abrasiveness are considered keystone factors allowing its large geographi- cal range. 4. The broad, species-specific range of climate parameters associated with the habitats was used to define the thresholds of the species' feeding niches. Our data suggest that climate as represented by the variables we used does not, in the first instance, relate to the tooth wear signature of equids. 5. We thus conclude that the tooth wear signature is more specific to the feeding strategy and should be regarded a dietary trait rather than an immediate imprint of climate. However, tooth wear can be used to infer local dietary traits and might therefore become an important tool in conservation management.

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.

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.

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

In mammals, tooth function, and its efficiency, depends both on the mechanical properties of the food and on chewing dynamics. These aspects have rarely been studied in combination and/or at the intra-specific level. Here we applied 3D dental surface texture analysis to a sample of field voles (Microtus agrestis) trapped from Finnish Lapland at different seasons and localities to test for inter-population variations. We also explored intra-individual variation in chewing dynamics by analysing two facets on the second upper molars. Our results confirm that the two localities have similar environments and that the voles feed on the same items there. On the other hand, the texture data suggest that diets are seasonally variable, probably due to varying concentrations of abrasives. Lastly, the textures on the buccal facets are more isotropic and their direction deviates more from the mesial chewing direction than the lingual facets. We interpret these results as reflecting food, rather than chewing, movements, where food particles are more guided on the lingual side of the molars. This has implications for the application of dental microwear analysis to fossils: only homologous facets can be compared, even when the molar row seems to constitute a functional unit.

Captive giraffe (Giraffa camelopardalis) mostly do not attain the longevity possible for this species and frequently have problems associated with low energy intake and fat storage mobilization. Abnormal tooth wear has been among the causes suggested as an underlying problem. This study utilizes a tooth wear scoring method ("mesowear") primarily used in paleobiology. This scoring method was applied to museum specimens of free-ranging (n=20) and captive (n=41) giraffes. The scoring system allows for the differentiation between attrition--(typical for browsers, as browse contains little abrasive silica) and abrasion--(typical for grazers, as grass contains abrasive silica) dominated tooth wear. The dental wear pattern of the free-ranging population is dominated by attrition, resembles that previously published for free-ranging giraffe, and clusters within browsing herbivores in comparative analysis. In contrast, the wear pattern of the captive population is dominated by abrasion and clusters among grazing herbivores in comparative analyses. A potential explanation for this difference in tooth wear is likely related to the content of abrasive elements in zoo diets. Silica content (measured as acid insoluble ash) is low in browse and alfalfa. However, grass hay and the majority of pelleted compound feeds contain higher amounts of silica. It can be speculated that the abnormal wear pattern in captivity compromises tooth function in captive giraffe, with deleterious long-term consequences.

Dietary characteristics and environmental variables are important selective factors directing ecological diversification in rodents. On Madagascar, the introductions and spread of the commensal black rat (Rattus rattus) can be seen as example cases to study dietary niche occupation and dietary adaptation in an insular environment. We investigate how tooth wear as a measure of dietary adaptation of black rats differs between four distinct habitats (village, manioc fields, spiny forest, and rainforest) with different dietary resources. We use the 3D surface texture analysis (3DST, using 30 parameters according to ISO 25178) as a measure of dietary abrasiveness. 3DST is applied on the occlusal surface of the upper first molar of 37 black rat specimens. The rainforest sample displays less rough and less voluminous surface textures compared to the village samples as indicated by smaller values for height parameters (Sa, Sp, Sq), inverse areal material ratio (Smc), and volume parameters (Vm, Vmc, Vmp, Vv, and Vvc). We therefore rank sampling areas from highest to lowest abrasiveness (village>manioc fields/spiny forest>rainforest). The rats from villages and rainforest differ to such an extent that one could have interpreted them to belong to different species. This indicates a high degree of variability in terms of ingesta abrasiveness. Furthermore, the pronounced difference between rats from human habitations compared to rats from associated fields or natural vegetation is interpreted to clearly indicate shifts in dietary niche occupation in relation to human impact.

Unworn teeth of herbivorous mammals are not immediately functional. They have to get in wear to expose enamel ridges which can then act as shear-cutting blades to disintegrate the food. We use the Plains Zebra (Equus quagga) as a hypsodont, herbivorous model organism to investigate how initial wear of the tooth crown is controlled by underlying structures. We find that the enamel proportion is smaller at the apical part of the tooth crown in all upper tooth positions. Measurements of enamel thickness on the first molar show that the outer enamel band is widest in the lower half of the tooth crown, where enamel content is also highest. We therefore find evidence that the distribution of enamel within the tooth crown is uneven and lower enamel content at the apex promotes early wear. This gradient in enamel distribution is less pronounced in the last molar (txM3), which has also a higher overall enamel content. The M3 is thus hypothesised to have a slightly different functional trait in mastication, resisting highest bite forces along the tooth row and maintaining functionality while anterior teeth are already worn down.

Unworn teeth of herbivorous mammals are not immediately functional. They have to be partially worn to expose enamel ridges which can then act as shear-cutting blades to break the food down. We use the Plains Zebra (Equus quagga) as a hypsodont, herbivorous model organism to investigate how initial wear of the tooth crown is controlled by underlying structures. We find that the enamel proportion is smaller at the apical half of the tooth crown in all upper tooth positions and suggest that lower enamel content here could promote early wear. Besides this uneven enamel distribution, we note that the third molar has a higher overall enamel content than any other tooth position. The M3 is thus likely to have a slightly different functional trait in mastication, resisting highest bite forces along the tooth row and maintaining functionality when anterior teeth are already worn down.

8 pages; International audience; Evolution in isolated island has shaped a variety of endemic taxa with outstanding characteristics. Amongst them is the extinct bovid genus Myotragus, endemic to Mallorca and Menorca Island, for which six succeeding species have been described: M. palomboi, M. pepgonellae, M. antiquus, M. kopperi, M. batei and M. balearicus. Myotragus has developed special cranial and post-cranial adaptations to meet the specific ecological demands of its insular habitat, like progressive dwarfing and fused limb elements. During its evolution, the dentition of Myotragus underwent subsequent changes: firstly a reduction in the number of teeth, and secondly an increase in hypsodonty. The ecological conditions inducing this dental evolution, especially Myotragus' diet, remain unknown. In this study, methods of 3D-dental topometry, enamel surface texture analysis according to ISO/FDIS 25178-2, and Scale-Sensitive Fractal Analysis (SSFA) are applied in order to infer palaeodiets of M. pepgonellae, M. kopperi, M. batei and M. balearicus, and to test the hypothesis that a dietary change may have occurred in the Myotragus lineage which relates to gradual morphological changes on upper second molars. We detect changes in the enamel/dentin ratio, enamel ridge length and enamel surface area within the lineage. Furthermore, Myotragus balearicus has enamel surface texture characteristics also present in extant browsing ungulates, while the three antecedent Myotragus species show an enamel surface texture signal similar to extant grazers. These results suggest a dietary change and are interpreted as a successive adaptation to limited resources in an isolated, insular environment. They can either be a consequence of a change in plant community structure or a successive expansion of Myotragus' dietary range due to increased intraspecific competition.

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.

The circumstances of the evolution of hypsodonty (= high-crowned teeth) are a bone of contention. Hypsodonty is usually linked to diet abrasiveness, either from siliceous phytoliths (monocotyledons) or from grit (dusty environments). However, any empirical quantitative approach testing the relation of ingested silica and hypsodonty is lacking. In this study, faecal silica content was quantified as acid detergent insoluble ash and used as proxy for silica ingested by large African herbivores of different digestive types, feeding strategies and hypsodonty levels. Separate sample sets were used for the dry ( n = 15 species) and wet ( n = 13 species) season. Average faecal silica contents were 17–46 g kg −1 dry matter (DM) for browsing and 52–163 g kg −1 DM for grazing herbivores. No difference was detected between the wet (97.5 ± 14.4 g kg −1 DM) and dry season (93.5 ± 13.7 g kg −1 DM) faecal silica. In a phylogenetically controlled analysis, a strong positive correlation (dry season r = 0.80, p < 0.0005; wet season r = 0.74, p < 0.005) was found between hypsodonty index and faecal silica levels. While surprisingly our results do not indicate major seasonal changes in silica ingested, the correlation of faecal silica and hypsodonty supports a scenario of a dominant role of abrasive silica in the evolution of high-crowned teeth.

Tooth wear can affect body condition, reproductive success and life expectancy. Poor dental health is frequently reported in the zoo literature, and abrasion-dominated tooth wear, which is typical for grazers, has been reported in captive browsing ruminants. The aim of this study was to test if a similar effect is evident in captive rhinoceros species. Dental casts of maxillary cheek teeth of museum specimens of captive black (Diceros bicornis; browser), greater one-horned (Rhinoceros unicornis; intermediate feeder) and white rhinoceroses (Ceratotherium simum; grazer) were analysed using the recently developed extended mesowear method for rhinoceroses. Captive D. bicornis exhibited significantly more abrasion-dominated tooth wear than their free-ranging conspecifics (pC. simum exhibited significantly less abrasion-dominated tooth wear, particularly in the posterior cusp of the second molar (p=0.005). In R. unicornis, fewer differences were exhibited between free-ranging and captive animals, but tooth wear was highly variable in this species. In both free-ranging and captive D. bicornis, anterior cusps were significantly more abrasiondominated than posterior cusps (pD. bicornis exhibited more homogenous tooth wear than their free-ranging conspecifics, which may be caused by an increase in the absolute dietary abrasiveness and a decrease in relative environmental abrasiveness compared to their freeranging conspecifics. The opposite occurred in C. simum. The results of this study suggest that diets fed to captive browsers are too abrasive, which could result in the premature loss of tooth functionality, leading to reduced food acquisition and processing ability and, consequently, malnourishment.

Lepidosauria show a large diversity in dietary adaptations, both among extant and extinct tetrapods. Unlike mammals, Lepidosauria do not engage in sophisticated mastication of their food and most species have continuous tooth replacement, further reducing the wear of individual teeth. However, dietary tendency estimation of extinct lepidosaurs usually rely on tooth shape and body size, which allows only for broad distinction between faunivores and herbivores. Microscopic wear features on teeth have long been successfully applied to reconstruct the diet of mammals and allow for subtle discrimination of feeding strategies and food abrasiveness. Here, we present, to our knowledge, the first detailed analysis of dental microwear texture on extant lepidosaurs using a combination of 46 surface texture parameters to establish a framework for dietary tendency estimation of fossil reptilian taxa. We measured dental surface textures of 77 specimens, belonging to herbivorous, algaevorous, frugivorous, carnivorous, ovivorous, insectivorous, molluscivorous, as well as omnivorous species. Carnivores show low density and shallow depth of furrows, whereas frugivores are characterized by the highest density of furrows. Molluscivores show the deepest wear features and highest roughness, herbivores have lower surface roughness and shallower furrows compared to insectivores and omnivores, which overlap in all parameters. Our study shows that despite short food–tooth interaction, dental surface texture parameters enable discrimination of several feeding strategies in lepidosaurs. This result opens new research avenues to assess diet in a broad variety of extant and extinct non-mammalian taxa including dinosaurs and early synapsids.