Dietary classification of extant kangaroos and their relatives (Marsupialia: Macropodoidea)

Kangaroos and kin (superfamily Macropodoidea) are the principal endemic herbivores of Australia and the most diverse radiation of marsupial herbivores ever to have evolved. As is typical of other herbivore groups (e.g. bovids), dietary niches span fruit, fungi, dicot leaves and monocot grasses in both specialists and generalists, but to date dietary classification has been largely ad hoc and poorly tied to actual dietary ecological data. Here we provide a simple dietary classification of the Macropodoidea based on an extensive literature survey. Intake of four major dietary items – grasses, dicot leaves, fruits and seeds, and fungi – was assessed using proportional intake for 19 species and categorical (ranked intake) data for 37 species. Statistical comparisons with cluster and principal components analyses aligned species into four dietary groups. Members of the first group have diets that primarily consist of fungi and fruits. Relative proportions of grasses to dicot leaves separate the remaining species into browser (more than 70% dicots), grazer (more than 70% grasses) and mixed feeder groups. Comparison of our diet‐based classification with a prevailing scheme based on dental morphology suggests that most species with what has traditionally been viewed as a ‘browser‐grade dentition’ are actually mixed feeders. This suggests that either morphology and diet are not tightly linked or that morphological differences between the dentitions of browsers and mixed feeders are subtle and have been overlooked. A positive correlation was found between body mass and average proportional intake of grass in the diet of macropodoids. This parallels the situation found in bovids, as well as the percentage cut‐off between dietary groups. These trends suggest that some underlying ecophysiological constraints may influence food choice in mammalian herbivores providing useful pointers to the diets of extinct taxa.     

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.     

牛科(哺乳纲:偶蹄目)动物与食物有关的适应形态模式

利用逐步分辨分析方法（Stepwise discriminant analysis,SCDA）检测了广义牛科动物的颅齿部结构,这些结构特征可以作为采食行为生态适应特征。在本研究中,测量了72种广义牛科动物的28个颅齿部结构。逐步分辨分析方法得出了6种采食方式适应类型：一般粗食者、新鲜禾草粗食者、开阔生境混合型采食者、精食者、郁闭生境混合型采食者、食果者。用103个标本检测了分辨指标的预测能力,所用标本为缺损标本,大多数缺少一项或多项结构。从这些标本获得的分辨函数的平均预测能力为94％,比用72种广义牛科动物标本建立的分辨函数的平均预测能力（98％）低一些。从一个颅齿部结构小样本建立的分辨函数可以用于考古发掘物中不完整标本的研究。这些指标与用颅下结构测量建立的运动能力和生境选择的指标相结合,可以推断古牛科动物的个体生态学以及古环境重建。     

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.     

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.     

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.     

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.     

Quantifying rapid spatial and temporal variations of CO<Subscript>2</Subscript> fluxes from small,lowland freshwater ponds

In Lake Tanganyika, several lineages of cichlids have diversified into 200 species. Tribe Tropheini contains the greatest richness of herbivorous species that are classified into four ecomorphs: grazers, browsers, suckers, and pickers. All of these ecomorphs coexist on littoral rocky shores. To reveal the differences among species within ecomorphs, we analysed fish shape and functional morphology related to feeding habits, using geometric morphometrics, and measured intestine length. Furthermore, we tested the relationship between genetic and morphological distances. As a result, diversities of functional morphology among ecomorphs, and among species within ecomorphs, were revealed. In grazers, morphological diversity was greatest in jaws and the opening direction of mouth, suggesting that these species have adapted to feed on various substrata. In browsers, intestine length varied among species, indicating that this ecomorph consists of species with various specializations in herbivory. Morphological divergence was found only in the mandible and occlusal facet in grazers. No clear relation was found between morphological and phylogenetic distances in browsers, which are a polyphyletic group, indicating that their traits were acquired by convergence in multiple lineages. Our data suggest that the observed morphological variation among species enable sympatric species to separate their feeding niches.     

Mandible Form Relative to the Main Food Type in Ladybirds (Coleoptera: Coccinellidae)

The Coccinellidae is an economically important family within the Coleoptera. Some members are phytophagous pests, but many are beneficial predators and valuable biocontrol agents. This study investigates the morphology of the mandibles of adult Coccinellidae in relation to diet. Using scanning electron microscopy on 86 species of Coccinellidae, it was found that the morphology of the mandibles was dictated by the general feeding method, and could only be used to indicate a phytophagous, mycophagous or carnivorous diet. Phytophagous Coccinellidae of the subfamily Epilachninae had mandibles with denticulate apical teeth and setae for feeding on leaf material. The mandibles of the mycophagous Psylloborini had secondary teeth on the ventral apical tooth for collecting fungal spores. The mandibles of carnivorous Coccinellidae and Scymninae had either a bifid or unidentate apex. The unidentate mandible seemed to be restricted to coccidophagous species. Many species also had a mandibular groove along which prey body juices were conducted. Although mandible morphology could be related to the overall feeding method, there was no relationship between specific diet or food taxon and mandible shape. Mandible shape does not appear to be especially restricting for changes in diet either in the ecological sense or over evolutionary time. Mandible morphology is of limited use in determining diet and host specificity in Coccinellidae that are being selected as potential biocontrol agents.     

Elemental and isotopic analyses of mammalian fauna from Southern Africa and their implications for paleodietary research

Elemental analyses of mammalian bone (e.g., strontium-calcium ratios, or Sr/Ca) distinguish between herbivores and carnivores; however, the relationships among herbivores are unclear. To study this question, a modern faunal sample from the Nagupande Tsetse Control Area (Zambezi drainage, Northwestern Zimbabwe) was used. This collection has the advantage of well-established geographical controls in addition to a varied fauna, which includes both bovids and suids. The grazing/browsing dietary status of each species was ascertained by means of isotopic analysis of carbon. Clear differences were seen in the δ¹³C of grazing and browsing animals; a specialized grazer was found to have significantly lower Sr/Ca than less specialized grazers and browsers. In this study it was also possible to examine differences in Sr/Ca by sex; female warthogs were found to have significantly lower Sr/Ca than males. The variation in certain animal groups was found to be abnormal. Implications for reconstruction of prehistoric human diets using trace-element techniques are discussed.     

Functional morphology of comminuting feeding structures of Trichodactylus borellianus (Brachyura,Decapoda, Trichodactylidae), an omnivorous freshwater crab

Crustaceans exhibit great diversity of feeding structures with morphological traits that are useful to infer the general trophic habits of species. In this study, we analyzed the functional morphology of comminuting feeding structures (mandibles, chelipeds, gastric mill) of the freshwater crab Trichodactylus borellianus directly related with the food fragmentation. The heterochely and mechanical advantage (MA) of the chelae were also studied. In both analyses, we considered the relationship between morphology and the natural diet. We expected to find a consistent relation between feeding habits and morphological traits. In general, we found simple structures armed with uniform setal systems and feeding appendages without pronounced teeth or spines. Mandibles have primarily cutting functions, helping with the food anchoring and fragmentation with mandibular palps armed with pappose setae. Chelipeds were covered with spines and simple setae. Adult males exhibited right-handedness with high MA of the major chelae. The ingested, relatively large pieces of food are finally chewed by a gastric mill equipped with sharp cusps characteristic of decapods with low ingestion of crude fiber material. The morphology of the feeding apparatus revealed that it is well adapted to an omnivorous diet, being able to cope with dietary changes.     

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.     

Combining isotopic and ecomorphological data to refine bovid paleodietary reconstruction: a case study from the Makapansgat Limeworks hominin locality

The relationship between environmental change and hominin evolution remains obscure. For the most part, this stems from the difficulty of reconstructing ancient hominin habitats. Bovids are among the most frequently utilized paleoenvironmental indicators, but little is known about the habitat preferences of extinct taxa. It is generally assumed that fossil bovids both ate the same things and occupied the same habitats as their closest extant relatives. We test the first part of this assumption by reconstructing the diets of seven bovids from Makapansgat Limeworks, South Africa. Since diet and habitat are linked, these reconstructions have implications for our understanding of fossil bovid habitat tolerances. Ecomorphological and stable carbon isotope analyses are employed, allowing us to take advantage of the strengths and overcome the weaknesses of both. In most cases, fossil bovids did have similar diets to their extant relatives, and probably occupied similar habitats. Gazella vanhoepeni and Aepyceros sp., however, were almost exclusive browsers, and not mixed feeders like their living counterparts.     

Guild‐specific shifts in visitation rates of frugivores with habitat loss and plant invasion

Habitat loss and plant invasions are two major drivers of global change in subtropical and tropical ecosystems. Both lead to a loss of biodiversity and alter species interactions, which may imperil vital ecosystem processes such as seed dispersal by frugivores. Reponses of frugivores to disturbance are often linked to their specialization on certain habitats or resources. Yet, it is poorly understood how habitat loss and plant invasion structure interactions between plants and different habitat or feeding guilds. Here we investigated whether visitation rates of frugivores change guild‐specifically with increasing habitat loss and invasion level in a heterogeneous subtropical landscape. In 756 h of observations, we recorded 1446 plant–frugivore interactions among 18 plant species and 42 avian frugivore species. Visitation rates of forest specialists decreased with increasing habitat loss, but not with changes in invasion level. In contrast forest generalists and forest visitors were unaffected by either driver. Similarly, obligate frugivores that overall showed a generalized fruit choice were unaffected by habitat loss and changes in invasion level. Contrary, visitation rates of specialized partial and opportunistic frugivores decreased with higher invasion level. Importantly, the negative effect of plant invasion on partial frugivores was more pronounced as habitat loss in the same study site increased, indicating a synergistic effect of the two drivers. The implications of our study are twofold: first, frugivores respond guild‐specifically to habitat loss and plant invasion. Thereby forest dependency is mainly related to habitat loss, and degree of frugivory mainly related to plant invasion. Forest generalists and obligate frugivores in turn may play a key‐role for forest regeneration in disturbed forest landscapes. Second, particularly frugivores with a specialized fruit choice may be threatened by synergistic effects between habitat loss and plant invasion.     

3D Morphometric Analysis Reveals Similar Ecomorphs for Early Kangaroos (Macropodidae) and Fanged Kangaroos (Balbaridae) from the Riversleigh World Heritage Area,Australia

Understanding feeding ecology of extinct kangaroos is fundamental to understanding the evolution of kangaroos and the Australia paleoenvironment during the Oligo-Miocene. Comparisons with extant species have suggested that the macropodiforms of the Oligo/Miocene (kangaroos and allies) from the Riversleigh World Heritage Area, northern Australia, were predominantly folivorous browsers or fungivores, unlike the majority of extant species. To further test this hypothesis, we investigate the relationship between variation in cranial and mandibular shape of extant and extinct macropodiforms and ecological factors such as diet, locomotion, and body mass using 3D geometric morphometric analysis of 42 living species and eight extinct species from two radiations (the extinct clade of Balbaridae and some early representatives of the extant Macropodidae. Dietary class (fungivore, browser, grazer, and mixed feeder) correlated strongly with variation in cranial shape (20–25% of variance explained). There was also significant association between cranial shape, and both locomotor mode and body mass. In a principal component analysis of shape variation for crania (including the shape of the molar row), Riversleigh macropodiforms cluster with extant folivorous browsers on principal components (PC) 1 and 3, providing support for previous interpretations of these species as browsing kangaroos. However, as a group and regardless of phylogenetic association, the shape centroid of extinct species differs significantly from that of extant species. Riversleigh macropodiforms cluster with regular hoppers or arboreal tree kangaroos, but this may be a result of the correlation between diet and locomotor mode in kangaroos. Their similarity to extant browsers supports previous interpretations of rainforest and woodland environments at Riversleigh during the early and middle Miocene, respectively. Procrustes ANOVA Analysis of the full shape dataset and diet also shows that diet accounts for a significant portion of variation; however, when phylogeny is taken into account these results become nonsignificant. In analyses of dentary shape, some balbarid species cluster with extant mixed feeders, although this may reflect phylogenetic differences rather than ecological signal.

     

Morphology and locomotor adaptations of the bovid femur in relation to habitat

Extant bovids inhabit a wide diversity of environments that range from forest to savanna and display locomotor patterns that are habitat specific. I report here on an investigation of the linkage between these locomotor patterns and habitat based on a study of the morphology of the bovid femur. Femoral head shape, shaft dimensions, and knee structure are examined and support a statistically significant separation of the different morphological complexes present in bovids from forest, broken cover, and savanna habitats. Morphological differences are primarily related to locomotor patterns as reflected in the degree of cursoriality displayed by bovids in different habitats. Cursorial bovids from savanna environments have laterally expanded femoral heads that act to limit the degree of abduction and axial rotation at the hip, and elliptically shaped distal femora that increase the moment arm of the extensor muscles that cross the knee. Forest bovids have spherically shaped femoral heads. This morphology permits a much higher degree of abduction and axial rotation at the hip and appears to provide greater maneuverability in a vegetationally complex habitat. Bovids living in broken cover environments that fall between the extremes of closed canopy forest and savanna display an intermediate set of femoral characters. This approach to the relationship between habitat and locomotion offers a potentially powerful means with which to examine the interplay between structural form and function in bovid evolution.     

Comparative finite element analysis of the cranial performance of four herbivorous marsupials

Marsupial herbivores exhibit a wide variety of skull shapes and sizes to exploit different ecological niches. Several studies on teeth, dentaries, and jaw adductor muscles indicate that marsupial herbivores exhibit different specializations for grazing and browsing. No studies, however, have examined the skulls of marsupial herbivores to determine the relationship between stress and strain, and the evolution of skull shape. The relationship between skull morphology, biomechanical performance, and diet was tested by applying the finite element method to the skulls of four marsupial herbivores: the common wombat (Vombatus ursinus), koala (Phascolarctos cinereus), swamp wallaby (Wallabia bicolor), and red kangaroo (Macropus rufus). It was hypothesized that grazers, requiring stronger skulls to process tougher food, would have higher biomechanical performance than browsers. This was true when comparing the koala and wallaby (browsers) to the wombat (a grazer). The cranial model of the wombat resulted in low stress and high mechanical efficiency in relation to a robust skull capable of generating high bite forces. However, the kangaroo, also a grazer, has evolved a very different strategy to process tough food. The cranium is much more gracile and has higher stress and lower mechanical efficiency, but they adopt a different method of processing food by having a curved tooth row to concentrate force in a smaller area and molar progression to remove worn teeth from the tooth row. Therefore, the position of the bite is crucial for the structural performance of the kangaroo skull, while it is not for the wombat which process food along the entire tooth row. In accordance with previous studies, the results from this study show the mammalian skull is optimized to resist forces generated during feeding. However, other factors, including the lifestyle of the animal and its environment, also affect selection for skull morphology to meet multiple functional demands. J. Morphol. 276:1230–1243, 2015. © 2015 Wiley Periodicals, Inc.     

Using carbon isotope analysis of the diet of two introduced Australian megaherbivores to understand Pleistocene megafaunal extinctions

Aim Australia lost a diverse assemblage of large marsupial herbivores in the late Pleistocene, with suggestions that the extinctions were biased towards browsers. In modern times two bovines, the Asian water buffalo (Bubalus bubalis) and banteng (Bos javanicus), have established feral populations in the Northern Territory, Australia. Buffalo have aggressively expanded throughout the savanna landscape, yet banteng remain near their point of introduction on the Cobourg Peninsula. We hypothesized that this difference is related to feeding ecology, possibly reflecting a legacy of the Pleistocene extinctions. Location Western Arnhem Land, Northern Territory, Australia. Methods Analysing a previously published dataset of body mass and feeding ecology of extinct and extant marsupial herbivores, we evaluated whether browsers were at greater risk of extinction than grazers. We compared the carbon isotope composition and nitrogen content of banteng and buffalo dung in order to evaluate the hypotheses that the differences in invasion success are related to feeding ecology, and that seasonal variation in browse consumption is linked to changing nutritional quality of grass. Results Controlling for body mass, the Pleistocene extinctions were clearly biased towards browsers. Introduced banteng appear to be primarily browsers, with their diets comprising 40% grass in the wet season and 15% in the late dry season. Buffalo have a more variable diet, with an increasing proportion of browse from the wet (30%) to the late dry season (75%), and can therefore be described as switching from grazer to browser. The decline of grass in the diet of both species appears to reflect the decline in the nutritional value of grass through the dry season, an inference supported by the negative relationship between δ¹³C values and the nitrogen content of dung. Main conclusions Banteng and buffalo are much larger than extant native herbivores, of which browsers are restricted to isolated rocky habitats. This suggests that banteng and buffalo have filled niches made vacant following the Pleistocene extinctions. The success of buffalo appears to be related to their greater dietary breadth, which enables them to graze and browse in eucalypt savannas, whilst the browsing banteng remain tethered to a mosaic of rain forest patches. The restriction of browsers may be a long‐range consequence of habitat transformations associated with Aboriginal landscape burning.     

Phylogenetic analysis of stomach adaptation in digestive strategies in African ruminants

The stomach morphology of 28 species of artiodactyls that differ in feeding style (browser, mixed feeder, grazer) was analysed using a multivariate approach and phylogenetic correction in order to test whether stomach morphology was correlated with feeding style when body mass was controlled for. A total of 25 morphological traits of the stomach were used in the analysis. After the effects of body mass and phylogeny on stomach morphology were taken into account, there was no significant grouping of species according to feeding style. When information about the feeding style of each species was included in the analysis, the set of morphological traits separated the mixed feeders from the other two feeding styles, but grazers and browsers had similar morphological features. Most of the variance in stomach morphology was explained by body mass and a lesser proportion by phylogeny. The main morphological features that have previously been proposed as being adaptations in grazing species, namely, lengthening of the retention time of ingesta to achieve an increase in their fibre digestion capability by means of a larger relative stomach capacity, a greater subdivision of chambers and smaller openings, are not supported by the findings of this study. Thus, there is no consistent evidence to support a significant adaptive effect of stomach morphology to different diets in the Artiodactyla.