Seasonal diet and prey selection of black‐backed jackals on a small‐livestock farm in South Africa

The extent to which black‐backed jackals (Canis mesomelas) selectively consume domestic sheep (Ovis aries) compared to wild prey is unknown. Using faecal analysis and prey surveys, we determined the seasonal diet and prey selection of jackals on a small‐livestock farm in South Africa. Sheep comprised 25–48% of the biomass consumed by jackals across seasons, and consumption peaked during the lambing seasons, indicating sheep often were the main food resource for jackals. Another main food resource was wild ungulates <50 kg, primarily springbok (Antidorcas marsupialis) and steenbok (Raphicerus campestris), which comprised 8–47% of the biomass consumed. Other important food items were mammals 1–3 kg (4–16%), which included hares (Lepus spp.) and springhares (Pedetes capensis), and small rodents (10–14%). Compared to the biomass available, jackals selectively consumed mammals 1–3 kg over sheep across all seasons, whereas wild ungulates <50 kg were selectively consumed over sheep in most seasons. Our results showed that jackals selectively consumed different food items throughout the year and that wild prey were consistently selected over sheep.     

Late autumn trophic flexibility of the golden jackalCanis aureus

The feeding habits of the golden jackal Canis aureus (Linnaeus, 1758) were compared using scat analysis in Hungary (temperate climate agricultural area), Greece (Mediterranean marshland), and Israel (Mediterranean agricultural area). Samples (84, 70 and 64 scats, respectively) were collected during late autumn, a period with capital importance to the long term survival of young jackals, during which they become independent. Predation of wild-living prey species was highest in Hungary, consisting primarily of small mammals (biomass estimation: 51.5%, mainly rodents), contrary to Israel and Greece, where scavenging on domestic animals dominated the diet of jackals (74%, mainly poultry and 62.6%, mainly goats, respectively). The highest consumption of wild ungulates (mainly wild boar) was found in Greece (15.7%), and plants in Hungary (39%). Bird consumption was low in all three areas. Reptiles, amphibians and fish occurred only in the diet of jackals in Greece and Israel, whereas invertebrates were eaten more frequently in Hungary. Jackal dietary composition was extremely variable between regions, strongly associated with human presence. These results illustrate the golden jackal as having a very variable diet, resulting from opportunistic feeding habits.     

Resource partitioning among cape foxes,bat-eared foxes,and black-backed jackals in South Africa

Cape foxes (Vulpes chama) and bat-eared foxes (Otocyon megalotis) are sympatric with black-backed jackals (Canis mesomelas) over much of southern Africa, although competition with and/or predation by jackals may suppress local populations of both fox species. From 2005 to 2008, we captured, radio-collared, and monitored 11 cape foxes, 22 bat-eared foxes, and 15 black-backed jackals on a game ranch in South Africa to investigate their spatial, habitat, temporal, and dietary resource overlap. Mean annual home-range sizes were 27.7 km² for cape foxes, 5.0 km² for bat-eared foxes, and 17.8 km² for jackal family groups. Home ranges overlapped completely between species, although core areas overlapped less (<45%), with cape foxes and jackals overlapping the least (12%). When active, cape foxes, but not bat-eared foxes, used core areas of jackal groups less than expected. Additionally, both fox species used jackal core areas less than expected for their den sites, suggesting areas outside jackal core areas were used as refuges by foxes. Strong levels of habitat partitioning were not apparent at the study site or home-range levels, although habitat selection for den sites differed between jackals and cape foxes. Jackals were the most diurnal across seasons, whereas cape foxes were the most nocturnal. Diets overlapped little (R₀ = 0.20–0.34) among the canid species, with bat-eared foxes overlapping the least with the others. Jackals killed at least 5 collared bat-eared foxes and 1 collared cape fox, indicating potential interference competition, probably for exclusive use of territorial space rather than over shared resources. We conclude that bat-eared foxes coexisted with jackals primarily by their dietary specialization and group living. Cape foxes coexisted with jackals by exhibiting high levels of spatial, habitat, temporal, and dietary partitioning. Surprisingly, the fox species exhibited positive associations with each other. Our results show the mechanisms that may allow jackals to suppress fox populations, yet also show how foxes, in turn, use different mechanisms to coexist with a dominant canid. © 2012 The Wildlife Society.     

Erratum to: Using multistate capture–mark–recapture models to quantify effects of predation on age-specific survival and population growth in black-tailed deer

Effective species management and conservation relies on accurate estimates of vital rates and an understanding of their link to environmental variables. We used multistate capture–mark–recapture models to directly quantify effects of predation on age-specific survival of black-tailed deer Odocoileus hemionus columbianus in California, USA. Survival probabilities were derived from individual encounter histories of 136 fawns and 57 adults monitored over 4 years. Based on results from our survival analysis we parameterized a Lefkovitch matrix and used elasticity analyses to investigate contributions of mortality due to predation to changes in population growth. We found strong evidence for age-specific survival including senescence. Survival of females >1 year old was consistently low (0.56 ± 0.18 for yearlings, 0.77 ± 0.13 for prime-aged females, and 0.55 ± 0.08 for senescent individuals), primarily due to high puma Puma concolor predation during summer. Predation from black bears Ursus americanus and coyotes Canis latrans was the primary cause for low annual survival of fawns (0.24 ± 0.16). Resulting estimates of population growth rates were indicative of a strongly declining population (λ = 0.82 ± 0.13). Despite high sensitivity to changes in adult survival, results from a lower-level elasticity analysis suggested that predation on fawns was the most significant individual mortality component affecting population decline. Our results provide a rare, direct link between predation, age-specific survival and the predicted population decline of a common ungulate species. The magnitude of predation was unexpected and suggests that ungulates in multi-predator systems struggle to cope with simultaneous reductions in survival probabilities from predators targeting different age classes.     

Landowners’ perspectives of black‐backed jackals (Canis mesomelas) on farmlands in KwaZulu‐Natal,South Africa

Despite continued efforts to eradicate black‐backed jackals (Canis mesomelas), they are considered an abundant mesopredator on agricultural land across South Africa, resulting in ongoing human–wildlife conflict and concern for farmers and wildlife managers. We conducted a questionnaire survey and semi‐formal interviews with farmers throughout KwaZulu‐Natal, examining farmers’ livestock husbandry, land‐use changes and perspectives towards jackals as a perceived threat to livestock. Many (75%) respondents acknowledged expanding agricultural activities on their farmlands since the onset of their farming careers. However, the perception was that these changes placed little pressure on mesopredators as farmers reported frequent daily (25%) and weekly (31%) sightings of jackal, and regular predation on livestock (72%). Some landowners (31%) reported between one and five livestock losses annually and suggest that mitigation strategies to prevent livestock losses are in place. Farmers suggested the increasing intensity in agricultural practices provided a greater food source for jackals allowing them to thrive in expanding agricultural conditions and, in some circumstances, farmers admitted to possibly being a cause through poor disposal techniques for dead animals. Feedback from farmers emphasized the importance of having collaboration between farmers to control jackal predation and reduce human–wildlife conflict.     

Social organization,home ranges,and extraterritorial forays of black-backed jackals

We radio-tracked 15 black-backed jackals (Canis mesomelas) from 8 adjacent family groups on Benfontein Game Farm (i.e., Benfontein) in South Africa to investigate their movement patterns and social organization. Jackal family groups consisted of mated pairs (alphas), 0–3 nonbreeding adults (betas), and pups, depending on the season. Mean (±SE) home-range size of alphas (9.4 ± 1.2 km², n = 6) did not differ (P = 0.766) from betas (9.8 ± 0.7 km², n = 8). Most beta jackals (8 of 10) remained philopatric on Benfontein, apparently because of the high density of springbok (Antidorcas marsupialis), their preferred prey. Three of 5 alphas and all 8 betas went on extraterritorial forays (i.e., forays). Generally, betas spent more of their active time on forays (2–20% of time) than alphas (0–3%; P = 0.048), and betas went farther on forays (2–8 km) than alphas (2–3 km; P = 0.003). The number of forays differed (P < 0.001) among seasons; most forays occurred during summer (64%) when jackals visited neighboring livestock farms, apparently to predate on domestic sheep. Overall, our results indicate forays by jackals are affected by social status, seasonal availability of preferred prey, and the reproductive cycle of jackals. To reduce jackal predation on livestock farms near reserves, we recommend that preventative measures (e.g., use of herders, jackal control activities) be increased during summer when jackals are most likely to travel outside reserves. © 2019 The Wildlife Society.     

Diet and prey profiles of three sympatric large carnivores in Bandipur Tiger Reserve, India

We conducted a field study of diets of three sympatric large carnivores, the tiger Panthera tigris , the leopard Panthera pardus and the dhole Cuon alpinus in Bandipur Tiger Reserve, India, based on analyses of 381, 111 and 181 scats, respectively. The frequency of occurrence of prey items in scats was converted to relative biomass and number of prey consumed using regression equations based on earlier feeding trials. The results showed that although these predators kill ∼11–15 species of vertebrate prey, relatively abundant ungulate species provide 88–97% of biomass consumed by them. Although the dietary niche overlap among the three species was high (Pianka's index of 0.75–0.93), some specialized predation was observed. The largest ungulates, gaur Bos gaurus and sambar Cervus unicolor , provided 73% of biomass consumed by tigers, whereas medium-sized chital Axis axis and wild pig Sus scrofa formed 65 and 83% of the biomass intake of leopards and dholes, respectively. In terms of the relative numbers of prey animals killed by the three predators, chital, which is the most abundant prey species, dominated their diets (tiger=33%, leopard=39% and dhole=73%). The results of the study, in conjunction with earlier work, support the prediction that abundance of ungulate prey species, as well as their availability in different size classes, are both critical factors that facilitate sympatry among the three predators.     

Black-backed jackal diet at Mokolodi Nature Reserve, Botswana

We examined the feeding habits of black‐backed jackals at Mokolodi Nature Reserve, Botswana, by analysing 237 scats collected between November 1995 and February 1997. Jackal dietary habits reflected the availability of a wide variety of food items and the differential vulnerability of prey. Potential animal and plant food available to jackals varies throughout the year because of its seasonal character. Seasonality of prey occurrence in scats was pronounced for small mammals, miscellaneous fruits and invertebrates. Across all seasons, mammals were the most common food resource (32.4%, n = 168), followed by anthropogenic items (14.8%), fruits (12.9%), invertebrates (10.8%), birds (8.5%), unidentified items (3.5%) and reptiles (1.4%). The presence of domestic mammals and poultry remains in scats reveals their importance in the diet of jackals and the tendency of jackals to frequent human settlements in search of food. Some ecological implications of jackal dietary habits are also explored.     

Postpartum Group Cohesion of Sympatric Deer in Texas

ABSTRACT Postpartum behavior of maternal deer may be specific to species of deer and predators. We captured sympatric white-tailed deer (Odocoileus virginianus) and mule deer (O. hemionus eremicus) fawns from radiocollared adult females in 2004–2006 on rangelands of west central Texas, USA, where predators larger than bobcats (Lynx rufus) were absent. Our objective was to determine whether differences in postpartum antipredator behavior existed between deer species, and if so, examine efficacy of those strategies. We collected postpartum group cohesion data in 2004 and 2005 by using radiotelemetry and examined dead fawns for cause of mortality. During fawns' hider phase, <3 weeks postpartum, mule deer females kept fawns closer to themselves (95% CI = 39−66 m) and twins closer to each other (95% CI = 25–49 m) than did white-tailed deer females (95% CIs = 152–234 m and 163–255 m, respectively). After 30 days postpartum, familial group cohesion was similarly tight for both species. During hider phases from 2004 to 2006, predated carcasses of white-tailed deer fawns (11 of 11) were dismembered or consumed more than mule deer fawns (7 of 13, P = 0.016), which was one line of evidence for maternal defense by mule deer adults. During hider phases in 2004 and 2005, predation rate of mule deer fawns was lower than that for white-tailed deer fawns. In 2006, predation rate increased for mule deer but was similar for white-tailed deer fawns compared with previous years. The tight cohesion strategy of mule deer exhibited in 2004 and 2005 seemed successful at thwarting small predators. Without large predators, the loose cohesion strategy of white-tailed deer females was maladaptive. When meso-predators are abundant due to extermination of larger predators, predation on fawns could increase if a deer species has relatively fixed postpartum maternal antipredator behavior.     

Golden jackal (Canis aureus L.) diet in Southern Greece

The diet of jackals was studied in the Mediterranean lowlands of Fokida and Samos island, Greece, by analyzing 127 scats collected between January 2002 and May 2003. Across all seasons frequencies of food items show that the most common items were mammals (frequency 42.7%, biomass 69.8%) and birds (12.0%, biomass 27.7%). Although the frequencies of plant material (27.3%) and insects (18.0%) were quite high, their biomass contribution was low (1.7%, 0.8% respectively). Most of the biomass consumed composed of mammals of domestic livestock origin (55.9%) which were presumably scavenged. This reveals the importance of this food item to the opportunistic jackals in wildlife-poor ecosystems like the anthropogenic Mediterranean lowlands. The occurrence of small mammals in the scats was very low while very few traces of grass and human refuse (such as leftovers of meals, plastic, pieces of paper etc.) were found in the diet of jackals. Furthermore, the findings support the opportunistic nature of a species capable to exploit any easily available food source.     

Ungulate herbivory reduces abundance and fluctuations of herbivorous insects in a boreal old-growth forest

Ecological theory predicts the strongest ecosystem effects of herbivory when dominant and ecologically important species are consumed. Bilberry, Vaccinium myrtillus, is such a key plant species, attractive to many other species in the boreal forests, for example ungulate and invertebrate herbivores. Large herbivores may remove substantial biomass and alter plant quality and therefore affect abundance and populations of invertebrate animals sharing the same food plant. We combined experimental exclusion of ungulates with a browsing intensity gradient to investigate the 15-year effect of ungulate (Cervus elaphus and Ovis aries) browsing on bilberry plant size and on bilberry-feeding herbivorous larvae (Lepidoptera and Symphyta), in a Norwegian old growth boreal forest ecosystem. Bilberry ramets in exclosure plots had nearly nine times higher dry mass and three times higher abundance of invertebrates feeding on them than in ungulate-access plots. Sweep-netting data verified these findings as larval numbers were twice as high in exclosure plots. The pattern in the large herbivore effects on bilberry size and abundance of herbivorous larvae were identical along the browsing gradient. Differences in larval abundance between treatments, as indicated by leaf-chewing, increased during the 15-year study period, and the community fluctuations were larger when ungulate herbivores were excluded. The browsing effect was moderated by plant quality as larval densities were lowest on both heavily-browsed and non-browsed plants, and highest on ramets that had 50–74% of annual shoots browsed. Our study supports previous findings in that bilberry is relatively disturbance tolerant and may recover quickly, but that ungulates may compete with herbivorous larvae for food biomass. Additionally, our results strongly indicates that population insect community peaks and fluctuations are dampened by ungulate consumption. Our findings add to the understanding on how ungulates may structure forest ecosystems directly and indirectly.     

Ungulates,rodents, shrubs: interactions in a diverse Mediterranean ecosystem

Ungulate abundance has increased dramatically worldwide, having strong impacts on ecosystem functioning. High ungulate densities can reduce the abundance, diversity and/or body condition of small mammals, which has been attributed to reductions in cover shelter and food availability by ungulates. The densities of wild ungulates have increased recently in high-diversity Mediterranean oak ecosystems, where acorn-dispersing small rodents are keystone species. We analysed experimentally ungulate effects on seed-dispersing rodents in two types of oak woodland: a forest with dense shrub layer and in dehesas lacking shrubs. Ungulates had no significant effects on vegetation structure or rodent body mass, but they reduced dramatically rodent abundance in the lacking-shrub dehesas. In the forest, ungulates modified the spatial distribution and space use of rodents, which were more concentrated under shrubs in the presence than in the absence of ungulates. Our results point to the importance of shrubs in mediating ungulate–rodent interactions in Mediterranean areas, suggesting that shrubs serve as shelter for rodents against ungulate physical disturbances such as soil compaction, trampling or rooting. Holm oak seedling density was reduced by ungulates in dehesa plots, but not in forests. Acorn consumption by ungulates may reduce oak recruitment to a great extent. Additionally, we suggest that ungulates may have a negative effect on oak regeneration processes by reducing the abundance of acorn-dispersing rodents. Given that shrubs seem to mediate ungulate effects on acorn dispersers, controlled shrub encroachment could be an effective alternative to ungulate population control or ungulate exclusion for the sustainability of the high-diversity Mediterranean oak ecosystems.     

Predator–prey relationships in a middle Asian Montane steppe: Persian leopard versus urial wild sheep in Northeastern Iran

Management controversies arise when both of the prey and predator in an ecosystem are species of conservation concern. We investigated trophic interactions between the endangered Persian leopard (Panthera pardus saxicolor) and a declining mountain ungulate, urial wild sheep (Ovis vignei), on a high-altitude steppe of Iran. During two consecutive photo-trapping seasons of 1,300 nights in total, a minimum population of four adult leopards (one female and three males) was documented. Scat analysis indicated that urial wild sheep was the staple of the leopard diet with 48.44 % of total biomass consumed. Remains of domestic livestock in leopard scats were negligible yet alarming (14.53 % biomass consumed), followed by wild pigs (8.13 %) and wild goat (1.26 %). Financial costs of leopard depredation to livestock breeders during our study period were comparatively lower than livestock–leopard conflict hotspots across Iran. Using distance sampling, urial density was 15.8 individuals km^?2 (±SE 6.2), and a total biomass of 47,621.5 kg for wild ungulates in the study area was estimated. We estimated that the annual removal rate of urial by leopards during our study period was 9.4 % of the total urial population. We suggest that continuous monitoring of the leopard and prey populations to assess predation impact should be considered, particularly in areas where a single species comprises a remarkable proportion of the leopard diet. In the meantime, assessing probable conflicts with local communities is recommended as a parallel management action to ensure long-term human–leopard coexistence. Our findings will aid wildlife managers in prey-depleted arid environments of western Asia to identify susceptible wild prey populations to predation by large carnivores; hence, significantly contribute in development and implementation of effective conservation measures to mitigate management conflicts.     

Winter diet composition of the golden jackal (Canis aureus L., 1758) in Serbia

The winter diet composition of golden jackals was determined by analysing the stomach contents of 248 specimens collected between December and February 2005–2009 at six localities in Serbia. The average weight of stomach contents was 189.9 ± 137.3 g. At all localities, livestock carcasses were the primary food category (frequency 56.1%, biomass 77.7%). The secondary food category consisted of small mammals taken as live prey (frequency 20.7%, biomass 5.2%). Other food categories were present less frequently (roe deer, wild boar, hare, and birds), and rarely (plant material, dogs, carnivores, lizards, and inedible inorganic material). No statistically significant differences were found in diet between jackals from different localities. The only difference was found between yearlings and adults with regard to the consumed biomass (%B). The analysis of the winter diet of golden jackals in Serbia indicates that the species has opportunistic feeding habits consisting primarily of easily accessible food sources.     

Oscillations in large mammal populations: are they related to predation or rainfall?

Cyclic population dynamics is relatively common among populations of small mammals in high latitudes but is not yet established among African savanna ungulates. However, oscillations may be expected in large mammal populations subject to quasi‐periodic oscillations in regional rainfall. We evaluated evidence for environmentally entrained oscillations in a large‐mammal predator–prey system in Kruger National Park (KNP), South Africa, where rainfall exhibits quasi‐periodic oscillations. The evaluation is based on analysis of comparative changes in the abundance of twelve ungulate species throughout South Africa's KNP using population counts over the period 1965–1996. We present evidence suggesting that (i) twelve ungulate populations display cyclic variability with half‐periods ranging between 10 and 18 years, (ii) this variability was associated with lagged rainfall between 3 and 10 years back in the past for different ungulate species, and (iii) the ungulate species respond in contrasting ways to rainfall, with some reaching highest abundance during periods of low rainfall and others under conditions of high rainfall. These findings are not consistent with the response pattern we would expect if the population oscillations were driven directly by the rainfall influence on food availability. Instead they seem to be an outcome of predator–prey interactions, which are entrained by the effect of rainfall on habitat conditions affecting the relative susceptibility of the different ungulate species to predation.     

Comparing patterns of human harvest and predation by Eurasian lynx Lynx lynx on European roe deer Capreolus capreolus in a temperate forest

Human harvest is the most important mortality factor for wild ungulates in Europe and can affect several aspects of ungulate biology. There is a growing concern about possible negative side effects of human harvest. To better understand the differences between human and natural mortality, we compared the extent, age and sex structure, nutritional condition, spatial and temporal distribution of human harvest, and natural predation by the Eurasian lynx Lynx lynx on the European roe deer Capreolus capreolus, the most abundant wild ungulate in Europe. Compared to the human harvest, lynx were less likely to kill fawns and yearlings than adults, and among adult deer, lynx were more likely to kill females. The proportion of roe deer with fat-depleted bone marrow was higher among lynx prey than among harvested animals. Average lynx kill rate was estimated to 47.8 roe deer per year, and lynx predation was considerably lower than the human harvest in the same area. While human harvest increased with higher roe deer density, lynx predation was similar across the gradient of roe deer densities. Comparison with other countries indicated that differences between human harvest and natural mortality of ungulates vary considerably in different parts of Europe. Variation in hunting practices and, even more importantly, carnivore predation may have an important role in buffering unwanted side effects of harvest of wild ungulates.     

Short‐term foraging responses of a generalist predator to management‐driven resource pulses

Generalist predators characteristically switch prey in response to resource pulses. Ungulate population reinforcement through translocation provides stressed and vulnerable prey, and apex predator reintroduction provides carrion, both resources that can be exploited by generalist predators. We investigated the dietary response of black‐backed jackal to two management interventions in the Karoo National Park, South Africa, namely the reinforcement of the springbok population, and then the reintroduction of lions. By analysing jackal diets from scats collected before and after each management intervention, we show that the relative per cent occurrence and biomass consumed of springbok increased in the diet following the population reinforcement of springbok. In contrast, large ungulates were more prevalent in the diet following apex predator reintroduction. These results suggest that jackals took advantage of a potentially vulnerable abundant springbok population following their population reinforcement, and then switched to foraging on carrion provided by reintroduced lions. These results provide insights into the dietary response of a generalist mesopredator to management interventions. A key lesson from the study is that the diet of generalist predators is context‐dependent and should be interpreted in that light.     

Leopard food habits in the Lopé National Park, Gabon, Central Africa

To determine leopard Panthera pardus (Linnaeus) food habits in the Lopé National Park in Gabon, Central Africa, 196 leopard scats were collected and analysed. Prey items were determined using undigested matter isolated from the scats, and a reference collection of hairs and bones from potential prey species. A minimum of 30 different prey species were identified, 27 of which were mammalian. Leopards preyed mainly on ungulates, which made up 59% of the biomass consumed. Diurnal primates (18%) and large rodents (17%) were also heavily preyed upon. The mean prey weight estimated from scats was 29.2 kg. The most important single prey species was found to be red river hog Potamochoerus porcus (Linnaeus), making up 20% of the biomass consumed, followed by forest buffalo Syncerus caffer nanus (Boddaert) and cane rat Thryonomys swinderianus (Temminck), each comprising 13% of biomass consumed.     

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.