Correlations between lizard cranial shape and diet: a quantitative, phylogenetically informed analysis

Although the relationship between dietary and phenotypic specialization has been well documented for many vertebrate groups, it has been stated that few such general trends can be established for lizards. This is often thought to be due to the lack of dietary specialization in many lizards. For example, many species that are reported to be insectivorous may also consume a variety of plant materials, and the reverse is often true as well. In this study, we investigate whether a correlation exists between general cranial form and dietary niche in lizards. Additionally, we test previously proposed hypotheses suggesting that herbivorous lizards should be larger bodied than lizards with other diets. Our data indicate that lizards specializing in food items imposing different mechanical demands on the feeding system show clear patterns of morphological specialization in their cranial morphology. True herbivores (diet of fibrous and tough foliage) are clearly distinguished from omnivorous and carnivorous lizards by having taller skulls and shorter snouts, likely related to the need for high bite forces. This allows herbivores to mechanically reduce relatively less digestible foliage. Carnivores have relatively longer snouts and retroarticular processes, which may result in more efficient capture and processing of elusive prey. When analysed in an explicit phylogenetic context, only snout length and skull mass remained significantly different between dietary groups. The small number of differences in the phylogenetic analyses is likely the result of shared evolutionary history and the relative paucity of independent origins of herbivory and omnivory in our sample. Analyses of the relationship between diet and body size show that on average herbivores have a larger body size than carnivores, with omnivores intermediate between the two other dietary groups. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 86 , 433–466.     

The Gut Bacterial Community of Mammals from Marine and Terrestrial Habitats

After birth, mammals acquire a community of bacteria in their gastro-intestinal tract, which harvests energy and provides nutrients for the host. Comparative studies of numerous terrestrial mammal hosts have identified host phylogeny, diet and gut morphology as primary drivers of the gut bacterial community composition. To date, marine mammals have been excluded from these comparative studies, yet they represent distinct examples of evolutionary history, diet and lifestyle traits. To provide an updated understanding of the gut bacterial community of mammals, we compared bacterial 16S rRNA gene sequence data generated from faecal material of 151 marine and terrestrial mammal hosts. This included 42 hosts from a marine habitat. When compared to terrestrial mammals, marine mammals clustered separately and displayed a significantly greater average relative abundance of the phylum Fusobacteria. The marine carnivores (Antarctic and Arctic seals) and the marine herbivore (dugong) possessed significantly richer gut bacterial community than terrestrial carnivores and terrestrial herbivores, respectively. This suggests that evolutionary history and dietary items specific to the marine environment may have resulted in a gut bacterial community distinct to that identified in terrestrial mammals. Finally we hypothesize that reduced marine trophic webs, whereby marine carnivores (and herbivores) feed directly on lower trophic levels, may expose this group to high levels of secondary metabolites and influence gut microbial community richness.     

Revisiting the cost of carnivory in mammals

Predator–prey relationships play a key role in the evolution and ecology of carnivores. An understanding of predator–prey relationships and how this differs across species and environments provides information on how carnivorous strategies have evolved and how they may change in response to environmental change. We aim to determine how mammals overcame the challenges of living within the marine environment; specifically, how this altered predator–prey body mass relationships relative to terrestrial mammals. Using predator and prey mass data collected from the literature, we applied phylogenetic piecewise regressions to investigate the relationship between predator and prey size across carnivorous mammals (51 terrestrial and 56 marine mammals). We demonstrate that carnivorous mammals have four broad dietary groups: small marine carnivores (< 11 000 kg) and small terrestrial carnivores (< 11 kg) feed on prey less than 5 kg and 2 kg, respectively. On average, large marine carnivores (> 11 000 kg) feed on prey equal to 0.01% of the carnivore's body size, compared to 45% or greater in large terrestrial carnivores (> 11 kg). We propose that differences in prey availability, and the relative ease of processing large prey in the terrestrial environment and small prey in marine environment, have led to the evolution of these novel foraging behaviours. Our results provide important insights into the selection pressures that may have been faced by early marine mammals and ultimately led to the evolution of a range of feeding strategies and predatory behaviours.     

The relationship between diet and tooth complexity in living dentigerous saurians

Living saurian reptiles exhibit a wide range of diets, from carnivores to strict herbivores. Previous research suggests that the tooth shape in some lizard clades correlates with diet, but this has not been tested using quantitative methods. I investigated the relationship between phenotypic tooth complexity and diet in living reptiles by examining the entire dentary tooth row in over 80 specimens comprising all major dentigerous saurian clades. I quantified dental complexity using orientation patch count rotated (OPCR), which discriminates diet in living and extinct mammals, where OPCR‐values increase with the proportion of dietary plant matter. OPCR was calculated from high‐resolution CT‐scans, and I standardized OPCR‐values by the total number of teeth to account for differences in tooth count across taxa. In contrast with extant mammals, there appears to be greater overlap in tooth complexity values across dietary groups because multicusped teeth characterize herbivores, omnivores, and insectivores, and because herbivorous skinks have relatively simple teeth. In particular, insectivorous lizards have dental complexities that are very similar to omnivores. Regardless, OPCR‐values for animals that consume significant amounts of plant material are higher than those of carnivores, with herbivores having the highest average dental complexity. These results suggest reptilian tooth complexity is related to diet, similar to extinct and extant mammals, although phylogenetic history also plays a measurable role in dental complexity. This has implications for extinct amniotes that display a dramatic range of tooth morphologies, many with no modern analogs, which inhibits detailed dietary reconstructions. These data demonstrate that OPCR, when combined with additional morphological data, has the potential to be used to reconstruct the diet of extinct amniotes. J. Morphol. 278:500–522, 2017. © 2017 Wiley Periodicals, Inc.     

Evidence for a multi‐peak adaptive landscape in the evolution of trophic morphology in terapontid fishes

Using the Australian marine‐freshwater terapontid fishes as a model system, we examined the role of dietary phenotypic optima in an adaptive macro‐evolutionary landscape. Comparative modelling relying on both a priori and data‐driven identification of selective regimes suggested multi‐peak models as best describing much of the dietary phenotypic landscape of terapontids. Both approaches identified common phenotypic optima for different lineages of marine and freshwater herbivores, and minimal differentiation between carnivores and omnivores, irrespective of their phylogenetic relationships, as the model best describing morphological evolution. Significant correlations also existed between these phenotypic axes and proportions of non‐animal dietary items in species’ diets. While simulation results provided evidence for a multi‐peak adaptive landscape in the evolution of trophic morphology in terapontids, they could not rule out chance convergence in these adaptive peaks. However, they do provide scope for identifying areas for more detailed, functionally specific study of phenotypic convergence in herbivorous terapontid trophic habits. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 623–634.     

The effect of dietary protein quality on nitrogen isotope discrimination in mammals and birds

We tested the competing hypotheses that (1) nitrogen discrimination in mammals and birds increases with dietary nitrogen concentration or decreasing C:N ratios and, therefore, discrimination will increase with trophic level as carnivores ingest more protein than herbivores and omnivores or (2) nitrogen discrimination increases as dietary protein quality decreases and, therefore, discrimination will decrease with trophic level as carnivores ingest higher quality protein than do herbivores. Discrimination factors were summarized for five major diet groupings and 21 different species of birds and mammals. Discrimination did not differ between mammals and birds and decreased as protein quality (expressed as biological value) increased with trophic level (i.e., herbivores to carnivores). Relationships between discrimination factors and dietary nitrogen concentration or C:N ratios were either the opposite of what was hypothesized or non-significant. Dietary protein quality accounted for 72% of the variation in discrimination factors across diet groupings. We concluded that protein quality established the baseline for discrimination between dietary groupings, while other variables, such as dietary protein intake relative to animal requirements, created within-group variation. We caution about the care needed in developing studies to understand variation in discrimination and subsequently applying those discrimination factors to estimate assimilated diets of wild animals.     

Similar Associations of Tooth Microwear and Morphology Indicate Similar Diet across Marsupial and Placental Mammals

Low-magnification microwear techniques have been used effectively to infer diets within many unrelated mammalian orders, but the extent to which patterns are comparable among such different groups, including long extinct mammal lineages, is unknown. Microwear patterns between ecologically equivalent placental and marsupial mammals are found to be statistically indistinguishable, indicating that microwear can be used to infer diet across the mammals. Microwear data were compared to body size and molar shearing crest length in order to develop a system to distinguish the diet of mammals. Insectivores and carnivores were difficult to distinguish from herbivores using microwear alone, but combining microwear data with body size estimates and tooth morphology provides robust dietary inferences. This approach is a powerful tool for dietary assessment of fossils from extinct lineages and from museum specimens of living species where field study would be difficult owing to the animal’s behavior, habitat, or conservation status.     

Intestine length in the fishes of a tropical stream: 2. Relationships to diet — the long and short of a convoluted issue

Synopsis We examined the relationship between the intestine length and the amount of plant material in the diet of 21 species of fish from forest streams in Panama. Alimentary tract analyses supplemented by literature reports showed that four loricariid catfish species and one poeciliid were specialized herbivores consuming almost exclusively periphyton and detritus. Four species, including one erythrinid, one characid, one trichomyctycterid and one eleotrid, were carnivores consuming almost entirely food of animal origin. Twelve species, including five characids, one lebiasinid, two pimelodelids, three cichlids and one poeciliid, were omnivores consuming food of both plant and animal origin, but the average proportion of food of plant origin (detritus and algae plus higher plant parts) varied from 4–60%. Most omnivores increased plant food consumption with increasing size. Because intestine length increases allometrically with body size and the pattern of increase differs considerably among species and is influenced by length:mass relationships, we compared species at the same size and took both length and mass into account. At a given size, intestine lengths of herbivores were longer than those of omnivores, and these were longer than those of carnivores. Differences in intestine length among the dietary categories were greater at larger body sizes and when the common size was defined by body mass than when it was defined by body length. There was no trend for the average proportion of plant material consumed to be related to intestine length among the omnivores, when confounding effects of body mass were taken into account. The slopes of the allometric equations relating log₁₀ intestine length to log₁₀ body size for herbivores tended to be higher than for omnivores and higher for omnivores than for carnivores, but both herbivores and omnivores showed extensive variation and overlap with the other dietary categories. Among the omnivores, there was no trend for slopes to be steeper for species consuming more plant material on average or for species showing larger ontogenetic increases in plant consumption. These results permit increased precision in describing diet-intestine length relationships, but indicate that the widely held belief that intestine length reflects diet in fishes should only be applied to broad dietary categories and not to finer divisions among omnivores.     

Climate and mammalian life histories

Mammals display considerable geographical variation in life history traits. To understand how climatic factors might influence this variation, we analysed the relationship between life history traits – adult body size, litter size, number of litters per year, gestation length, neonate body mass, weaning age and age at sexual maturity – and several environmental variables quantifying the seasonality and predictability of temperature and precipitation across the distribution range of five terrestrial mammal groups. Environmental factors correlated strongly with each other; therefore, we used principal components analysis to obtain orthogonal climatic predictors that could be used in multivariate models. We found that in bats, primates and even‐toed ungulates adult body size tends to be larger in species inhabiting cold, dry, seasonal environments, whereas in carnivores and rodents a smaller body size is characteristic of warm, dry environments, suggesting that low food availability might limit adult size. Species inhabiting cold, dry, seasonal habitats have fewer, larger litters and shorter gestation periods; however, annual fecundity in these species is not higher, implying that the large litter size of mammals living at high latitudes is probably a consequence of time constraints imposed by strong seasonality. On the other hand, the number of litters per year and annual fecundity were greater in species inhabiting environments with higher seasonality in precipitation. Lastly, we found little evidence for specific effects of environmental variability. Our results highlight the complex effects of environmental factors in the evolution of life history traits in mammals. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 719–736.     

Tooth root morphology as an indicator for dietary specialization in carnivores (Mammalia: Carnivora)

The Carnivora occupy a wide range of feeding niches in concordance with the enormous diversity in their skull and dental form. It is well established that differences in crown morphology are linked to variations in the material properties of the foods ingested and masticated. However, how tooth root form is related to dietary specialization is less well known. In the present study, we investigate the relationship between tooth root morphology and dietary specialization in terrestrial carnivores (canids, felids, hyaenids, and ursids). We specifically address the question of how variation in tooth root surface area is related to bite force potentials as one of the crucial masticatory performance parameters in feeding ecology. We applied computed tomography imaging to reconstruct and quantify dental root surface area in 17 extant carnivore species. Moreover, we computed maximal bite force at several tooth positions based on a dry skull model and assessed the relationship of root surface area to skull size, maximal bite force, food properties, and prey size. We found that postcanine tooth root surface areas corrected for skull size serve as a proxy for bite force potentials and, by extension, dietary specialization in carnivores. Irrespective of taxonomic affinity, species that feed on hard food objects have larger tooth roots than those that eat soft or tough foods. Moreover, carnivores that prey on large animals have larger tooth root surface areas. Our results show that tooth root morphology is a useful indicator of bite force production and allows inferences to be made about dietary ecology in both extant and extinct mammals. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105, 456–471.     

Dental root size in bats with diets of different hardness

The relationship between tooth roots and diet is relatively unexplored, although a logical relationship between harder diets and increased root surface area (RSA) is suggested. This study addresses the interaction between tooth morphology, diet, and bite force in small mammals, phyllostomid bats. Using micro computed tomography (microCT), tooth root morphology of two fruit‐eating species (Carollia perspicillata and Chiroderma villosum) and two insect‐eating species (Mimon bennettii and Macrotus californicus) was compared. These species did not differ in skull or estimated body size. Food hardness, rather than dietary classification, proved to be the strongest grouping factor, with the two insectivores and the seed‐processing frugivore (C. villosum) having significantly larger RSAs. Bite force was estimated using skull measurements; bite force significantly correlated with tooth RSA but not with body size. Although the three durophagous species did exhibit larger crowns, the area of the occlusal surface did not vary among the four species. There was a linear relationship between root size and crown size, indicating that the roots were not expanded disproportionately; instead the entire tooth was larger in the hard diet species. MicroCT allows the nondestructive quantification of previously difficult‐to‐access tooth morphology; this method shows the potential for tooth roots to provide valuable dietary, behavioral, and ecological information in small mammals. J. Morphol. 276:1065–1074, 2015. © 2015 Wiley Periodicals, Inc.     

Non-linear relationship between body size of terrestrial carnivores and their trophic niche breadth and overlap

Studying food partitioning of mammalian predators is important for understanding trophic structures and interactions between coexisting carnivore species. This is particularly pertinent in the light of expanding ranges of populations of generalist species whose habitat and diet overlap with more specialized species. Here, we tested the resource partitioning hypothesis in terrestrial carnivores, predicting that trophic niche breadth and overlap relate positively to body mass. We used dietary data from 18 terrestrial carnivore taxa in four families (Canidae, Mustelidae, Felidae and Ursidae; body mass 0.1–173.6 kg) in three regions in Central and Eastern Europe, i.e. deciduous forest and forest-steppe region (DFR), temperate deciduous and mixed forest region (MFR) and transitory mixed forest regions (TFR). We ranked carnivores along an axis of trophic niche (breadth and overlap), and analysed the relationship between trophic niche and body mass (or pair-wise difference in body mass). A hierarchical cluster analysis of diet composition divided carnivores into four ecological groups: wild ungulate predators; small-mammal predators; amphibians and small mammal predators and omnivores. The relationship between body mass of predators and both trophic niche breadth and trophic niche overlap were hump-shaped. The trophic niche breadth to body mass ratio was significantly lower in DFR than in TFR and trophic niche overlap was significantly higher in DFR than in MFR and TFR. The predominant food resource is small mammals whose abundance is related to local agricultural and forestry management practices. Modifications of management techniques can affect population dynamics and community composition of carnivore species, especially in the case of small-mammal predators.     

Green world maintained by adaptation

An ecological theory (HSS hypothesis) predicts that carnivores maintain the terrestrial ecosystem with abundant plants (green world) by regulating herbivore abundance. However, a weak density dependence of herbivores will make the equilibrium unstable and results in population oscillations with a large amplitude. Here, we study a possibility that the dynamics can be stabilized if defence trait by herbivores and offence trait by carnivores change in an adaptive manner. When the cost constraints on adaptation are strong in both the herbivores and the carnivores, the equilibrium is more likely to be stable if the herbivore adapts more quickly than the carnivore. When the constraints on the adaptation are asymmetric between species, the equilibrium is likely to be unstable. We conclude that the green world may be maintained by fast and costly adaptation by the herbivore through mechanisms such as phenotypic plasticity and behavioural change. Plant defence which is poisonous and prickly has been proposed as one of explanations, however, world can be green through adaptation in higher trophic levels even without plant’s defence.     

How do food passage rate and assimilation differ between herbivorous lizards and nonruminant mammals?

Summary What digestive adaptations permit herbivorous nonruminant mammals to sustain much higher metabolic rates than herbivorous lizards, despite gross similarity in digestive anatomy and physiology? We approached this question by comparing four herbivorous species eating the same diet of alfalfa pellets: two lizards (chuckwalla and desert iugana) and two mammals (desert woodrat and laboratory mouse). The mammals had longer small and large intestines, greater intestinal surface area, much higher (by an order of magnitude) food intake normalized to metabolic live mass, and much faster food passage times (a few hours instead of a few days). Among both reptiles and mammals, passage times increase with body size and are longer for herbivores than for carnivores. The herbivorous lizards, despite these much slower passage times, had slightly lower apparent digestive efficiencies than the mammals. At least for chuckwallas, this difference from mammals was not due to differences in body temperature regime. Comparisons of chuckwallas and woodrats in their assimilation of various dietary components showed that the woodrat's main advantage lay in greater assimilation of the dietary fiber fraction. Woodrats achieved greater fiber digestion despite shorter residence time, but possibly because of a larger fermentation chamber, coprophagy, and/or different conditions for microbial fermentation. We conclude with a comparative overview of digestive function in herbivorous lizards and mammals, and with a list of four major unsolved questions.     

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.     

Examining predator–prey body size,trophic level and body mass across marine and terrestrial mammals

Predator–prey relationships and trophic levels are indicators of community structure, and are important for monitoring ecosystem changes. Mammals colonized the marine environment on seven separate occasions, which resulted in differences in species'' physiology, morphology and behaviour. It is likely that these changes have had a major effect upon predator–prey relationships and trophic position; however, the effect of environment is yet to be clarified. We compiled a dataset, based on the literature, to explore the relationship between body mass, trophic level and predator–prey ratio across terrestrial (n = 51) and marine (n = 56) mammals. We did not find the expected positive relationship between trophic level and body mass, but we did find that marine carnivores sit 1.3 trophic levels higher than terrestrial carnivores. Also, marine mammals are largely carnivorous and have significantly larger predator–prey ratios compared with their terrestrial counterparts. We propose that primary productivity, and its availability, is important for mammalian trophic structure and body size. Also, energy flow and community structure in the marine environment are influenced by differences in energy efficiency and increased food web stability. Enhancing our knowledge of feeding ecology in mammals has the potential to provide insights into the structure and functioning of marine and terrestrial communities.     

Generic species richness and body mass in North American mammals: Support for the inverse relationship of body size and speciation rate

Generic species richness, the number of species per genus, is examined as a function of mean generic body mass for extant North American mammals. Species richness decreases as an inverse power function with increased mass, and the Spearman rank correlation coefficient of the logio transformed data is significant (r_s= ‐0.37). When the data are partitioned by trophic level, the relationship is not statistically significant for carnivores but strengthens for herbivores (r_s= ‐0.46). This interesting but incidental effect is due to the negligible number of diminutive and excessively large carnivores, which is in turn determined by foraging strategies. Alternate hypotheses for the “right‐skewed”; size distribution of modern North American mammals, such as disproportionate extinction of large species, differential species longevity, and a geographical scaling function, are rejected in favor of the proposition that elevated levels of speciation are restricted to animals of small body mass, as originally proposed by Gould and Eldredge (1977). This phenomenon is explained as a function of habitat restriction and particularly in herbivores, limited home range size. Aquatic mammals, regardless of body size, speciate rarely. Cope's Rule, the tendency of many animal groups to evolve towards large size, is understood as a probabilistic statement reflecting the phylogenetic tendencies of a disproportionately high number of small species alive at any given point in time.     

Mandible shape in marsupial and placental carnivorous mammals: a morphological comparative study using geometric morphometrics

We analysed mandible shape of the orders Dasyuromorpha, Didelphimorphia, and Carnivora using two‐dimensional geometric morphometrics, in order to explore the relationship between shape, size, and phylogeny. We studied 541 specimens, covering most of the genera of the terrestrial Carnivora (115 species) and a wide sample of marsupials (36 species). The observed shape variation had an ecological component. As an example, omnivorous carnivores have thick mandibles and large talonids in the carnassials, while hypercarnivores possess short mandibles and reduced talonids. There is also a discrimination between different taxonomic groups (i.e. marsupials and Carnivora), indicating some kind of constraint. Size explains a large percentage of total variance (large species had shorter and stronger mandibles, with anteriorly displaced carnassials), was significant when phylogeny was taken into account with a comparative method, but not when size and shape were optimized on the phylogeny. Carnivora presents a larger disparity and variation in body size, which could be related to the difference in teeth replacement. The optimization of mandible shape on the phylogenetic tree indicates that functional aspects, such as diet, are a key factor in the evolution of the carnivore mandible, but also that there is a phylogenetic pattern that cannot be explained by differences in diet alone. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164 , 836–855.     

Optimal foraging and community structure: implications for a guild of generalist grassland herbivores

Summary A particular linear programming model is constructed to predict the diets of each of 14 species of generalist herbivores at the National Bison Range, Montana. The herbivores have body masses ranging over seven orders of magnitude and belonging to two major taxa: insects and mammals. The linear programming model has three feeding constraints: digestive capacity, feeding time and energy requirements. A foraging strategy that maximizes daily energy intake agrees very well with the observed diets. Body size appears to be an underlying determinant of the foraging parameters leading to diet selection. Species that possess digestive capacity and feeding time constraints which approach each other in magnitude have the most generalized diets. The degree that the linear programming models change their diet predictions with a given percent change in parameter values (sensitivity) may reflect the observed ability of the species to vary their diets. In particular, the species which show the most diet variability are those whose diets tend to be balanced between monocots and dicots. The community-ecological parameters of herbivore body-size ranges and species number can possibly be related to foraging behavior.     

Does plant richness influence animal richness?: the mammals of Catalonia (NE Spain)

Although it has long been held that plant diversity must influence animal diversity, the nature of this relationship remains poorly understood at large spatial scales. We compare the species richness patterns of vascular plants and mammals in north‐eastern Spain using a 100‐km² grain size to examine patterns of covariation. We found that the total mammal richness pattern, as well as those of herbivores and carnivores considered separately, only weakly corresponded to the pattern of plants. Rather, mammal richness was best described by climatic variables incorporating water inputs, and after adding these variables to multiple regression models, plant and mammal richness were virtually independent. We conclude that the observed association, although weak, is explained by shared responses of both groups to climate, and thus, plant richness has no influence on the richness pattern of Catalan mammals.