Hypsodonty in ungulates: an adaptation for grass consumption or for foraging in open habitat?

Hypsodont (i.e. high-crowned) teeth have been interpreted as an indicator of feeding preferences and habitat selection in ungulates. For this reason, the degree of hypsodonty has been used for estimating the diet of ancient taxa and in palaeoenvironmental reconstructions. The goal of this study is to elucidate the relative importance of grass consumption and open habitat foraging in the development of hypsodont teeth, using novel computer techniques of knowledge discovery applied to a dataset of 134 species of artiodactyls and perissodactyls distributed among thirteen families. The results obtained suggest that high-crowned teeth represent an adaptation for feeding in an open habitat, although the minimum threshold of hypsodonty seems to increase with the relative length of the anterior part of the jaw. On the contrary, there is no direct relationship between the degree of hypsodonty and the percentage of grass consumed, except for the correspondence between grazing and dwelling in open habitats. A relatively wide muzzle evidences an adaptation for grass foraging in open and mixed habitats, but there are some non-grazing species from a closed habitat that also show wide muzzles. Thus, the hypsodonty index, combined with the length of the anterior part of the jaw and the width of the muzzle, allows accurate inferences on the ecological preferences of extinct ungulates.     

Body condition and ruminal morphology responses of free-ranging impala (Aepyceros melampus) to changes in diet

Variation of the intraruminal papillation pattern with diet quality has been described in many ruminant species, but the use of papillation measures as a proxy for habitat quality and nutritional status of animals has not been evaluated. We compared various measures of body condition (body mass, body condition score, kidney fat index, bone marrow fat index, adrenal mass, kidney to adrenal mass ratio), diet quality (%browse, protein and fibre content) and rumen papillation in 106 impalas (Aepyceros melampus) from four different locations in Zimbabwe. The various condition proxies indicated that periods of low diet quality are characterised by a high proportion of browse in the diet of this species. Animals with a high proportion of browse had more voluminous rumens, suggesting a compensation for lower diet quality by increased intake. Macroscopic papillation indices did not yield meaningful significant correlations with diet quality or body condition proxies, and hence, their use for estimating habitat or body condition cannot be advocated. In contrast to previous histological reports, ballooning cells of the Stratum corneum of the ruminal mucosa were more prominent in animals on lower-quality diets. There were significant correlations of the kidney to adrenal mass ratio with other body conditions and with diet quality indices, suggesting that poor body condition and low diet quality represent stressful situations.     

On the relationship between hypsodonty and feeding ecology in ungulate mammals,and its utility in palaeoecology

High‐crowned (hypsodont) teeth are widely found among both extant and extinct mammalian herbivores. Extant grazing ungulates (hoofed mammals) have hypsodont teeth (a derived condition), and so extinct hypsodont forms have usually been presumed to have been grazers. Thus, hypsodonty among ungulates has, over the past 150 years, formed the basis of widespread palaeoecological interpretations, and has figured prominently in the evolutionary study of the spread of grasslands in the mid Cenozoic. However, perceived inconsistencies between levels of hypsodonty and dental wear patterns in both extant and extinct ungulates have caused some workers to reject hypsodonty as a useful predictive tool in palaeobiology, a view that we consider both misguided and premature. Despite the acknowledged association between grazing and hypsodonty, the quantitative relationship of hypsodonty to the known ecology of living ungulate species, critical in making interpretations of the fossil record, was little studied until the past two decades. Also, much of the literature on ungulate ecology relevant to understanding hypsodonty has yet to be fully incorporated into the perspectives of palaeontologists. Here we review the history and current state of our knowledge of the relationship between hypsodonty and ungulate ecology, and reassert the value of hypsodonty for our understanding of ungulate feeding behaviour. We also show how soil consumption, rather than the consumption of grass plants per se, may be the missing piece of the puzzle in understanding the observed correlation between diets, habitats, and hypsodonty in ungulates. Additionally, we show how hypsodonty may impact life‐history strategies, and resolve some controversies regarding the relevance of hypsodonty to the prediction of the diets of extinct species. This in turn strengthens the utility of hypsodonty in the determination of past environmental conditions, and we provide a revised view of a traditional example of evolutionary trends in palaeobiology, that of the evolution of hypsodonty in horses and its correlation with the Miocene spread of grasslands in North America.     

Nutritional content of savanna plant foods: implications for browser/grazer models of ungulate diversification

Models of herbivore diversification rely heavily on adaptations that reflect the nutritional quality of foods consumed. In particular, browsers and grazers are expected to show dichotomous adaptations to deal with high quality (concentrate) browse-based and poor quality grass-based diets, respectively. In this study, we test the widespread assumption that browse represents a higher quality food source than grass. We analyzed plants from a South African savanna, collected over one dry and one wet season across several habitat types, for percent nitrogen (%N), neutral detergent fiber (NDF), acid detergent fiber (ADF), and acid detergent lignin (ADL) to compare variations in nutritional value of different food types. Results show consistently higher %N and lower NDF and ADF of tree foliage and forbs compared to monocots, but the former have consistently higher ADL, implying a higher fiber digestibility in grass compared with browse. Some fruit species have a high NDF and ADL content, implying poorer nutritional value than is commonly assumed. Our findings are in agreement with several other studies depicting relatively poor digestibility of browse (tree foliage and fruit) compared to grass. Reference to browse as high quality foods is therefore misleading, and models of herbivory that rest on this assumption require revision. The more efficient fiber digestibility recorded in grazers compared to browsers cannot be treated as an adaptation to poor quality diets, but rather to maximize benefits of higher fiber digestibility of grass. Spatio-seasonal variations in plant nutritional seem to reflect seasonal and spatial diet changes expected for grazers and intermediate (mixed) feeders. We propose that future studies require further detail on variations in diet, diet quality, and digestive efficiency to properly understand mechanisms of adaptation.     

Methodological considerations for the use of faecal nitrogen to assess diet quality in ungulates: The Alpine ibex as a case study

Faecal indices are used to describe spatiotemporal patterns of diet quality in wild herbivores, and near infrared spectroscopy methods (NIRS) distinctively reduce costs and labour compared to conventional chemical analyses. In this study we compared the prediction accuracy of faecal nitrogen (N), ash and fibrous fractions of a laboratory instrument (NIRS) with that of a lightweight portable instrument (VisNIRS) using faecal samples of Alpine ibex (Capra ibex). We then compared how different indices based on faecal N and fibrous fractions were able to detect the expected temporal trend and difference between sexes in diet quality, and verified whether the presumably lower accuracy of NIRS and VisNIRS (compared to wet chemistry) influenced the ability of indices to detect such trends. The laboratory NIRS was more accurate than the portable VisNIRS, especially for fibrous fractions. Predicted data from both instruments clearly identified the temporal trend identified by chemically analysed data, but only NIRS identified the small difference between sexes. Expressing faecal N as a proportion of organic matter (OM) or neutral detergent fibre (NDF) improved the power of the index as compared to expressing it as a proportion of dry matter (DM), irrespective of the method used to estimate the data. Faecal NDF contents followed the patterns expected from the variation of faecal N, while faecal acid detergent fibre (ADF) and especially lignin (ADL) followed contradictory patterns, which might be due to a selection of different forage sources rather than to changing quality of a homogenous forage source. We conclude that expressing faecal N as a proportion of OM is advised with wild herbivores faeces, where ash content can be biased by accidental soil contamination or ingestion. Faecal fibrous fractions are useful to complement faecal N and to avoid simplified interpretations of the results. For this purpose, NIRS methods can be extremely useful to predict multiple indices on large numbers of samples. VisNIRS portable instruments might be unable to detect subtle biological patterns in small numbers of samples. However, developing procedures for portable instruments outside laboratory facilities might further improve practicability, cost effectiveness, and, through allowing on-field predictions on large numbers of samples, power of statistical testing of diet quality indices.     

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.     

Fibre digestibility in large herbivores as related to digestion type and body mass — An in vitro approach

The coexistence of different ungulate species in a given ecosystem has been the focus of many studies. Differences between ruminant foregut fermenters and hindgut fermenters were remarkable for example in the way they ingest and digest high fibre diets. Digestion trials based on total collections are difficult to conduct or are sometimes even not possible for wild animals in the field or in zoos. To gain information on the fibre digestion achieved by these animals and the influence of body mass (BM) thereon, a method using spot sampling is desirable. In this study, in vitro fermentation of faecal neutral detergent fibre (NDF) was used as a measure of fibre digestion in large ungulates. Food and faecal samples of 10 ruminant foregut fermenting and 7 hindgut fermenting species/breeds were collected. All animals received 100% grass hay with ad libitum access. The NDF of food and faeces was fermented in vitro in a Hohenheim gas test (HGT) for 96 h. The digestion type generally had an effect on the gas production (GP) of faecal NDF in the HGT with hindgut fermenters showing higher values than ruminant foregut fermenters. At any time interval of incubation, BM had no influence on GP. The results are in accordance with both findings that ruminant foregut fermenters have longer mean retention times and more comprehensive particle reduction and findings of a lack of influence of BM on digesta mean retention time. It can be stated that the HGT (96 h) is a useful and quick method to show also small differences within groups in fibre digestion.     

THE SHAPE OF CONTENTION: ADAPTATION,HISTORY, AND CONTINGENCY IN UNGULATE MANDIBLES

Mandibles and teeth of ungulates have been extensively studied to discern the functional significance of their design. Grazing ungulates have deeper mandibles, longer coronoid processes, flatter incisor arcades, and more hypsodont molars in comparison to browsers. If the functional significance of both mandible and teeth shapes is well‐established, it remains uncertain to what extent mandible shapes are really adapted to grazing, meaning that they evolved either to serve their current biological function or just as a structural requirement to accommodate higher crowned molars. Here, we address this question by studying the contribution of phylogeny, hypsodonty, and body size to mandibular shape variation. The mandible shape appeared to be significantly influenced by hypsodonty but not by body size. Interestingly, hypsodonty‐related changes influenced the tooth row in artiodactyls and perissodactyls significantly but in the opposite directions, which is ultimately related to their different digestive strategies. Yet, we obtained a strong phylogenetic effect in perissodactyls, suggesting that their mandible shape should be strongly inherited. The strength of this effect was not significant within artiodactyls (where hypsodonty explained much more variance in mandible shape). Digestive strategy is deemed to interplay with hypsodonty to produce different paths of adaptation to particular diets in ungulates.     

The influence of fibre in the diet on growth rates and the digestibility of nutrients in the greater cane rat (Thryonomys swinderianus).

The greater cane rat Thryonomys swinderianus is a coprophagous rodent in which fermentation occurs in the large caecum. The extent to which a 45% increase in the fibre component of the diet influenced growth rates of cane rats and the digestibility of nutrients and energy was investigated in two feeding trials. Higher fibre levels in the diet reduced the digestibility of dry matter, protein and fat, while animals digested fibre components (neutral-detergent fibre, acid detergent fibre, hemicellulose and cellulose) with a comparable efficiency to those maintained on a low fibre diet. In one of the trials animals fed the high fibre diet exhibited significantly lower growth rates than animals fed the low fibre diet. Digestibility coefficients of the cane rats for neutral-detergent fibre and protein seem to be intermediate to high when compared to reported values for the porcupine, guinea-pig, degu and rabbit. It is suggested that the ability of cane rats to utilise large quantities of fibre enable them to survive periods when only dry grass is available.     

Comparative omasum anatomy in ruminants: Relationships with natural diet,digestive physiology,and general considerations on allometric investigations

The omasum is the third forestomach compartment of pecoran ruminants. It is assumed that the re-absorption of fluid present in the forestomach digesta (that facilitates particle sorting, digestion, and harvest of microbes) is its main function, so that less diluted digesta is submitted to enzymatic digestion in the lower digestive tract. Here, we evaluate measures of omasum size (representing 84 ruminant species in the largest data set) against body mass and proxies of the natural diet (%grass) or forestomach physiology (fluid throughput), using phylogenetically controlled models. The origin of specimens (free-ranging or captive) did not have an effect in the data set. Models with the best support invariably either included %grass or a physiology proxy in addition to body mass. These effects were not necessarily additive (affecting the intercept of the allometric regression), but often indicated a change in the allometric body mass-exponent with diet or physiology. Only models that allowed an influence on the allometric exponent yielded basic exponents compatible with predictions derived from geometry. Species that include more grass in their natural diet, or that have a “cattle-type” physiology marked by a high forestomach fluid throughput, generally have larger omasa. However, the existence of outliers, as well as the overall data pattern, suggest that this is not an obligatory morphophysiological condition. Circumstantial evidence is presented leading to the hypothesis that the comparatively small and less complex omasa of “moose-type” species do not necessarily represent an “original” state, but may be derived from more complex states by ontogenetic reduction and fusion of omasal laminae.     

Relationships between oral morphology and feeding style in the Ungulata: a phylogenetically controlled evaluation

In ungulates it is argued that specialization in the consumption of a particular type of food (feeding style) is reflected in morphological adaptations of the organs involved in the selection, processing and digestion of food. We analysed the differences in size and morphology of some oral traits that have been functionally related to food-selection ability (muzzle width, incisor-arcade shape, incisor shape), prehension of food (incisor protrusion), food comminution (molar occlusal surface area, hypsodonty (high-crowned molars)) and intake rate (incisor breadth) between ungulate species with different feeding styles (browser, mixed feeder, grazer). Grazers were characterized by large-body-size species. After controlling only for body mass, we found that grazers had wider muzzles and incisors, more-protruding incisors and more-bulky and higher-crowned molars than did mixed feeders and browsers. When the analyses took into account both body mass and phylogeny, only body mass and two out of the three hypsodonty indexes used remained significantly different between feeding styles. Browsers were smaller, on average, than mixed feeders and grazers, whilst grazers and mixed feeders did not differ in size. Also, browsers had shorter and less-bulky molars than did mixed feeders and grazers; the latter two feeding styles did not differ from each other in any of the hypsodonty indexes. We conclude that the adaptation to different dietary types in most of the oral traits studied is subsumed by the effects of body mass and the sharing of common ancestors. We hypothesize that differences in the ability to exploit different food resources primarily result from differences in body mass between species, and also discuss why hypsodonty characterizes feeding styles.     

Hypsodonty and tooth facet development in relation to diet and habitat in herbivorous ungulates: implications for understanding tooth wear

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Modelling ungulate dependence on higher quality forage under large trees in African savannahs

In African savannahs, large trees improve grass quality, particularly in dry and nutrient poor areas. Enhanced below-canopy grass nutrients, such as nitrogen and phosphorus contents should therefore attract and benefit grazers. To predict whether ungulates really need these forage quality islands we focused on four grazer species, i.e., zebra, buffalo, wildebeest, and warthog, differing in body size and digestive system. We confronted literature estimations of their feeding requirements with forage availability and quality, observed in three South African savannah systems, through linear modelling. The model predicted the proportion of below-canopy grass that grazers should include in their diet to meet their nutritional requirements.During the wet season, the model predicted that all animals could satisfy their daily nutrient requirements when feeding on a combination of below- and outside-canopy grasses. However, wildebeest, having relatively high nutrient demands, could meet their nutrient requirements only by feeding almost exclusively below canopies.During the dry season, all animals could gain almost twice as much digestible protein when feeding on below – compared to outside-canopy forage. Nonetheless, only warthogs could satisfy their nutrient requirements – when feeding almost exclusively on below-canopy grasses. The other ungulate species could not meet their phosphorus demands by feeding at either site without exceeding their maximum fibre intake, indicating the unfavourable conditions during the dry season.We conclude that grazing ungulates, particularly warthog, zebra, and buffalo, actually depend on the available below-canopy grass resources. Our model therefore helps to quantify the importance of higher quality forage patches beneath savannah trees. The composition of grazer communities depending on below-canopy grasses can be anticipated if grazer food requirements and the abundance of large trees in savannahs are known. The model suggests that the conservation of large single-standing trees in savannahs is crucial for maintenance of locally grazing herbivores.     

The diet of Impala (Aepyceros melampus) in the Sengwa Wildlife Research Area, Rhodesia

The seasonal variation in the plant morphological part composition of the dict of Impala in the Sengwa Wildlife Research Area, Rhodesia, was determined by the analysis of rumen content samples from shot animals. The grass species composition of the diet was examined by microscopic identification of grass fragments found in the rumens. The seasonal variation in the dicotyledonous species composition of the diet was studied by direct observation of feeding animals. The protein contents of the rumen contents and faeces were used as measures of diet quality. In the wet season, grass was selected in preference to dicotyledonous plants, and grass leaf was the preferred plant part. In the late wet and early dry seasons, forbs were the principal food. The proportion of woody dicotyledons in the diet was at a maximum in the mid-dry season, when diet quality was at a minimum. Diet quality was directly related to the proportion of grass in the diet. Female Impala had a significantly higher quality diet than males, probably as a result of differential habitat selection.     

The temporal scale of diet and dietary proxies

Diets estimated from different proxies such as stable isotopes, stomach contents, and dental microwear often disagree, leading to nominally well‐supported but greatly differing estimates of diet for both extinct and extant species that complicate our understanding of ecology. We show that these perceived incongruences can be caused by proxies recording diet over vastly different timescales. Field observations reveal a diet averaged over minutes or hours, whereas dental morphology may reflect the diet of a lineage over millions of years of evolution. Failing to explicitly consider the scale of proxies and the potentially large temporal variability in diet can cause erroneous predictions in any downstream analyses such as conservation planning or paleohabitat reconstructions. We propose a cross‐scale framework for conceptualizing diet suitable for both modern ecologists and paleontologists and provide recommendations for any studies involving dietary data. Treating diet in this temporally explicit framework and matching the scale of our questions with the scale of our data will lead to a much richer and clearer understanding of ecological and evolutionary processes.     

Fungal endophytes of a forage grass reduce faecal degradation rates

Mutualisms between fungal endophytes and forage grasses can exert broad-reaching effects on grassland communities and ecosystem processes. We hypothesised that endophytes of grasses would retard the process of faecal degradation since grazing animals consume primarily live plant material and excrete a large portion of the herbage they consume as faeces. We examined the degradation rates of faeces from sheep that had consumed pure swards of perennial ryegrass containing a range of unique strains (AR1, AR37, or Wild type) of the fungal endophyte, Neotyphodium lolii, or no endophyte. Ultimately, the presence of endophytes in perennial ryegrass resulted in slower faecal decay rates compared to the nil endophyte treatment, although only consistently for the C concentration decay rates that were approximately 2× to 4× slower in the endophyte-derived faecal matter. The decay rate of dry matter content was significantly slower (ca. 1.5×) in the novel endophyte-derived faeces (AR1 and AR37) compared to the nil endophyte-derived faeces. The N decay rates differed significantly only in the AR1 treatment that was approximately 4× slower than the nil endophyte group. The reduced decay rates are attributed to the presence of endophyte-derived alkaloids in the faeces, and a greater proportion of more easily degraded hemicellulose in faeces from sheep that consumed the endophyte-free grass. There were no significant differences in the faecal carbon and nitrogen decay rates among the three endophyte strain treatments. This suggests that all the strain-specific alkaloids might have similar effects, or that N. lolii has a general effect that is not strain-specific, such as altered fibre composition, as reported here. This is the first report of a fungal endophyte affecting the rate of faecal degradation, and the first report of the alkaloids peramine, lolitrem B and epoxy–janthitrems in faecal matter. This study shows that a common agronomic grass–endophyte mutualism can have effects on ecosystem processes that have not previously been considered.     

Reliability of δ13C and δ15N in faeces for reconstructing savanna herbivore diet

We tested the reliability of herbivore faecal δ¹³C and δ¹⁵N values for reconstructing diet through review of an extensive database derived from a 3-year study of ungulates in South Africa's Kruger National Park. Faeces are a useful material for stable isotope studies of diet because they record dietary turnover at very short time scales, and because sampling is non-invasive. However, the validity of faecal isotope proxies may be questioned because they represent only undigested food remains. Results from Kruger Park confirm that free-ranging browsers have faecal δ¹³C consistent with C₃ feeding, grazer faeces are C₄, and mixed-feeder faeces intermediate. Although the respective ranges do not overlap, there is significant variation in faecal δ¹³C of browsers and grazers (～2.0–4.0‰) across space and through time. We demonstrate that most (～70%) of this variation can be ascribed to corresponding patterns of variation in the δ¹³C of C₃ and C₄ plants, respectively, re-enforcing the fidelity of faecal isotope proxies for diet but highlighting a need for mixing models that control for variations in plant δ¹³C in order to achieve accurate diet reconstructions. Predictions for the effects of climate (rainfall) and ecophysiology on ¹⁵N-abundance variations in mammals do not persist in faeces. Rather, faecal δ¹⁵N tracks changes in plant δ¹⁵N, with further fractionation occurring primarily due to variations in dietary protein (reflected by %N). Controlling for these effects, we show that a dual-isotope multiple source mixing model (Isosource) can extend diet reconstructions for African savanna herbivores beyond simplified C₃/C₄ distinctions, although further understanding of variations in mammal δ¹⁵N are needed for greater confidence in this approach.     

Coefficients of digestibility and nutritional values of geophytes and tubers eaten by southern African mole-rats (Rodentia: Bathyergidae)

Subterranean rodents have high energy requirements when they are excavating their burrows. This study investigates the energy available to, and the efficiency with which it can be extracted by, four species of bathyergid mole-rats fed natural diets ranging from the underground storage organs of geophytes to grass roots and leaves.
The digestibility coefficients of geophytes ranged from 53% for the fibrous tuber of the gemsbok cucumber to 95–7% for corms and bulbs. One species, Bathyergus suillus whose diet consists of over 80% grass, had a digestibility coefficient of 87% on an all grass diet.
All species had similar coefficients of digestibility of > 90% when fed on a uniform diet of sweet potato.
Bulbs and corms had a low fibre content (3–3–4%), high calorific value (15–16kJ/g) and high digestibility coefficients (95–7–96%) and on this diet the mole-rats maintained their body mass. Food of lower digestibility tended to have a high fibre content (8–2–45%) and, with the exception of B. suillus , although the mole-rats consumed a greater quantity of food, they lost mass. The sweet potato had a low fibre content (4–1 %) but was energetically very similar to bulbs and corms (15–5 kJ/g).
Geophytes which have low fibre contents are generally small (1–20 g), whereas geophytes with high fibre contents are much larger (30–2000 g) and often occur in more arid zones.     

Metabarcoding reveals diet diversity in an ungulate community in Thailand

The diverse large mammal communities found in Asian dry forests and savannas should segregate based on their diet selection. We examined the diet composition of sympatric ungulate species using metabarcoding to determine whether their diet was segregated and whether obvious attributes (i.e., body size, phylogeny, ecology) explained the structure. We collected fecal samples from eight ungulate species in Huai Kha Khaeng Wildlife Sanctuary in the western forest complex of Thailand. The fecal collections occurred around a plot where all woody species were codified within a genetic barcode library, and this library was supplemented with samples from plant species known to be consumed by these species. Of 273 plant species tested, at least 93 were found within the fecal samples. Over half of the identified species were not previously known by experts as forage species. All ungulate species showed a strong consumption of grasses and forbs. For the three species with sufficient sample size (sambar, banteng, and guar), there were seasonal differences in their diet, with each showing increased occurrence of woody plants during the dry season. The pattern of forage consumption did not follow obvious paradigms of body size or taxonomy, with significant diet differences found in two similar‐sized bovids (gaur, banteng), while the diet of sambar was more similar to bovids than to the other deer species. Asian ungulates differ in their forage consumption and metabarcoding should allow for testing of diet shifts in response to seasonal rains and fires which dominate the phenology of Asian dry forests and savannas.