Ecology and predation

Ecology and predation

Chapter 6 Ecology and predation Chapter Outline Chapter 6.1. Population ecology The role of feline predation in the ecosystem Ecological or food pyra...

22MB Sizes 0 Downloads 40 Views

Chapter 6

Ecology and predation Chapter Outline Chapter 6.1. Population ecology The role of feline predation in the ecosystem Ecological or food pyramid (biomass) Predators maintain the plant herbivore balance Puma and mule deer in Kaibab’s highland (Arizona) Felines and the discovery of the dynamic ecological balance: numerical fluctuations The case of the Canadian lynx and the snowshoe hare Chapter 6.2. Predation: hunting methods and techniques Felids: the most skilled world’s predatory carnivores Relationship between dimensions of the predator and the prey The fatal bite Felines: not only solitary ambushes

310 311 311 311 313 313 315 315 315 317 321 322

6.2.1. The most common “hunting technique”: the ambush 6.2.2. The lion: hunting with the cooperation of members of the “pride” 6.2.3. The cheetah: long high-speed chase 6.2.4. The fishing cat (Prionailurus viverrinus) and flat-headed cat (Prionailurus planiceps): fishers, specialized hunters in wet habitats 6.2.5. Margay, marbled cat, Indochinese and Sunda clouded leopards: arboreal climbing hunters 6.2.3. Caracal and serval: masters of agility in the ambush final leap Serval (Leptailurus serval) Caracal (Caracal caracal)

323 325 327

330 334 338 339 340

Two cheetahs in front of a herd of wildebeest who have noticed their presence and remain vigilant fearing an attack. Photo courtesy of V. Martegani.

Felines of the World. DOI: https://doi.org/10.1016/B978-0-12-816503-4.00006-4 © 2020 Elsevier Inc. All rights reserved.

309

310

Felines of the World

This book does not address in depth the aspect of behavioral ecology of felines as this topic has many different results for each population of the various species, that it would be impossible to cover them all here, and the subject requires much further research. We confine ourselves to summarizing some basic points to highlight the general aspects of their ecology, the primary role played by felids in ecosystems, such as predators or superpredators, underlining the importance they have assumed over time, in maintaining ecosystem equilibrium. Predation for Felidae is an activity imposed by their very nature as “obligate carnivores” or Hypercarnivorous, as they require a series of nutrients contained in meat to live. These include taurine and arginine, two essential amino acids found only in animal tissues, and which are indispensable for the proper functioning of the heart, eyesight, and reproductive and excretory apparatus (kidneys). Other substances essential to the metabolism of felines are arachidonic and linoleic acid, contained in animal fats. Vitamins A, D, and the vitamin B complex must also be taken directly from animal tissues as felines are unable to obtain them using substances of vegetable origin or sunlight (as man can do, for example). Felines therefore have to exploit environmental resources, “consuming” through predation a part of the biomass of herbivores that live in the ecosystem; in turn herbivores and other biotic and abiotic components (soil type, characteristics of the territory, and climate) heavily influence the population structure of carnivores, their density, and also their social behavior. This means that, within the same species, populations distributed in different continents or even in different environments present particular eating habits; the various examples are innumerable. The leopard, widely distributed in Africa and Asia, has a range of different prey in the two continents not only due to the diversity of the herbivore species present. Being a highly adaptable species its diet

generally includes species weighing 8 17 kg, but in Gabon, according to Henschel et al. (2005), the range was 5.1 50 kg; and yet the range is even wider because its diet can include arthropods and small lagomorphs or large antelopes weighing more than 200 kg (kudu and young elands), young zebras, and young giraffes. The number of prey species is also noteworthy, ranging from African antelopes and gazelles to the deer of the Siberian regions (Capreolus pigargus, some Cervus sp., Moschus), mountain goats, Suidae (Phacochoerus, Potamochoerus, African and Asian wild boar), and many species of small and large primates, etc. The Asiatic wildcat (Felis lybica ornata) that feeds 74% on the Yarkan-Tarim hare (Lepus yarkandensis) in the cold steppes of the Chinese Xinijang (Abdukadir and Khan, 2013) has a completely different diet in the rest of its vast distribution area where it frequents very varied habitats (up to the arid and warm desert of the Indian Thar). Also, the phenomenon of “intraguild predation,” the predation of felids on other felids (or other carnivores) carried out by the larger species, influences the ecology of cats; all those species that frequent the same habitat as large cat species are subject to their predation or theft of their newly caught prey (see lions and cheetahs, etc.). In South America, is also the phenomenon called the “ocelot effect,” as the presence of ocelot, which is the largest species in the genus Leopardus, decreases the density of other species of the same genus, including margay and tiger cats, or oncillas, but also of the jaguarundi (Herpailurus yagouaroundi). Intraguild predation also influences the habits of some species: the marbled cat and Sunda clouded leopard of Sumatra have more arboreal habits than those of Borneo (Rabinowitz et al., 1987), where it is more terrestrial; this seems to be due to the fact that the tiger, marbled cat, and clouded leopard are sympatric in Sumatra (but the tiger is not present in Borneo).

Chapter 6.1 Population ecology Ecological pyramid referring to the use of biomass quantities at various trophic levels. https://savannabiomeac. weebly.com/pyramids.html; significantly modified by the author.

Ecology and predation Chapter | 6

The role of feline predation in the ecosystem It is as if nature had chosen the felids as guardians to safeguard the ecological balance of almost all terrestrial ecosystems. All species of big cats are superpredators at the tops of the food pyramids of many of the environments in which they live. Meanwhile the species of small wildcats, even the smallest, are still indispensable parts of the food web of most terrestrial ecosystems. Every natural environment, to be able to maintain the balance between the climatic characteristics, the soil, the vegetation, and the living beings that inhabit it, is subject to some natural mechanisms that have evolved over time and have made the ecosystem reach the state of “climax.” When an ecosystem reaches the climax state this means that the ecological succession that sustains it is capable of remaining in place over time, as long as the environmental conditions remain stable.

Ecological or food pyramid (biomass) Solar energy is the basis of every ecosystem, as described next. 1. Producer level (or autotroph plants) Solar energy is used by autotroph plants or producers, together with inorganic soil substances (nitrogen and other elements) and water and carbon dioxide in the air, to produce organic matter such as cellulose. The amount of plant matter produced in a given environment is called the plant biomass; this is expressed in weight or mass (e.g., kg) per unit of surface (e.g., m2 ) produced in a unit of time (e.g., 1 year), therefore plant biomass is expressed in, for example, kg/m2 /year. 2. Level of consumers of the first order or primary consumers Plant biomass is used as food by herbivorous animals, or consumers of the first order; in every environment, herbivores consume only about 10% of the available plant biomass. 3. Level of consumers of the second order or secondary consumers (also IIIrd, IVth order, etc.). The biomass of herbivorous animals of an environment, in turn, is preyed on and consumed only to a level of about 10% by the carnivores or secondary consumers.

311

4. Decomposing organisms The quantities of biomass and animal biomass plants that are not used by consumers in the food chain are decomposed by decomposing organisms (fungi, necrophagous invertebrates, bacteria), which transform the organic substances so that this returns to enrich and fertilize the soil; this favors the growth of plants, and the ecological cycle begins again. Solar energy-Autotroph plants or producers -Herbivorous animals or primary consumers-Carnivores or secondary consumers ----mm Decomposing organisms mm Moving from the base to the apex of the food pyramid, the number of species and the number of individuals of any terrestrial environment decrease. Often, at the top there is only one species of superpredator.

Predators maintain the plant herbivore balance It is now clear that where the big felids have been eliminated from an ecosystem, in a short time there has been an uncontrolled increase in the number of herbivores, with the consequent impoverishment of food resources provided by vegetation, which are exploited to complete exhaustion, causing the onset of famines, epidemics, and mass extinctions. The situation is not always reversible and many original ecosystems with their own species of vegetables, herbivores, and carnivores can no longer be reestablished due to the fact that the bare ground goes through a process of laterization (hardens, loses nutrients, and becomes dry) due to exposure to sun etc., to the point that suitable flora are no longer able to take root for environmental restoration. This happened, for example, on many of the Mediterranean coasts both in Europe and North Africa. North Africa, for example, was much less arid than now, and was once covered with woods of Mediterranean origins and also with tall trees such as holm oak (Quercus ilex), thorny oak (Quercus coccifera), cork oak (Quercus suber) and also cypresses, pines of Aleppo, etc. However, starting at the time of Roman domination, the species of felines that lived in that territory (lion, leopard, cheetah, caracal, serval) were almost completely exterminated at the same time as the destruction of their prey and the forest habitat in which they lived.

312

Felines of the World

The mosaics of an ancient Roman villa, Villa del Casale in Piazza Armerina in Sicily, document the hunts that were organized, especially in North Africa (but also in the Middle East), during the Roman domination. You can see many species of gazelles and large antelopes and also of felids that no longer live in northern Africa, such as the Barbary lion (in the photo is a lioness) and the leopard. Other animals that are represented there, but which today no longer live in North Africa are the elephant, the hartebeest (Alceplaphus buselaphus), the bear, the aurochs, etc. The big cats were captured and transported to Italy for shows in which they fought in the big arenas such as the Colosseum. The ecosystems of these regions were completely destroyed and turned into dry areas. https://upload.wikimedia.org/wikipedia/commons/thumb/d/d7/Villa_Romana_del_CasaleAmbulacre_de_la_Grande_Chasse-Lionne_transperc%C3%A9e_d%27une_fl%C3%A8che.jpg/1280px-Villa_Romana_del_CasaleAmbulacre_de_la_Grande_Chasse-Lionne_transperc%C3%A9e_d%27une_fl%C3%A8che.jpg

The puma (Puma concolor concolor), after the jaguar, is the largest feline in the Americas. Photo courtesy of A. Nun˜ez-Lemos.

Ecology and predation Chapter | 6

Puma and mule deer in Kaibab’s highland (Arizona) What happens when predators are missing from an ecosystem? A community of 5000 6000 mule deer (Odocoileus hemionus) protected in the forests of the Kaibab plateau in an area of 30,000 hectares in Arizona is exemplary. To allow an increase of deer, for 25 years (from 1906 to 1931) almost all their natural predators were destroyed, including more than 600 pumas (Puma concolor cougar). As shown in the graph, the result obtained was that, toward 1925, the population of deer

313

grew dramatically, reaching 100,000; the relationship between animal and plant resources became disastrous and over 60,000 specimens of mule deer died from hunger and disease, with the total number, after more than 30 years, returning to the original amount. The ecological role of predators is to limit the number of herbivores within optimal reproductive levels for a correct and rational exploitation of vegetation; this allows the vegetation to thrive and at the same time to nourish an adequate community of herbivores that allows the survival of the predators themselves.

By V. Ziswiler.

Felines and the discovery of the dynamic ecological balance: numerical fluctuations The ecological balance between the plant, herbivorous, and carnivorous species of an ecosystem with a number of species that is not too high, is not constantly stable, but

subject to regular cyclical variations called “fluctuations.” Applying a mathematical model (called the prey predator model) based on the equation of Lotka-Volterra (1925 26), the multiannual fluctuations between the number of prey and that of the predator have been demonstrated in a theoretical way.

314

Felines of the World

From the graph obtained by applying the equation, it is clearly seen that there are regular increases and decreases in the number of individuals of the prey species, followed equally regularly by increases and decreases in the number of individuals of the predator species. It was only by studying the populations of the Canadian lynx (Lynx canadensis) and those of its natural

prey, the snowshoe hare (Lepus americanus), that it was possible to prove for the first time that it was possible to apply in nature the mathematical model that expresses the predator/prey ratio, inserting in the theoretical graph the actual numerical variations shown by lynx and hare populations during the relevant years.

Graph of the predator prey mathematical model based on Lotka-Volterra’s equation.

Numerical fluctuations in the populations of Canadian lynx (Lynx canadensis) and snowshoe hare (Lepus americanus) alternate regularly every 7 9 years. This natural mechanism allows the vegetation to renew itself and support the population of hares. which in turn supports the population of lynxes (MacLulich, 1937). https://www.google.it/search?q 5 animal 1 population 1 over 1 time&tbs 5 simg:CAQSlwEJrCeZhdLP0VwaiwELEKjU2A QaBAgVCAoMCxCwjKcIGmIKYAgDEiiKCI0InRT3ApsUggjNEpQKoQeaFKM-7z_1wP9c_1pD6lK6E-oj6JK-IzGjAjuIXbDjvo0wpL4C-qdCq1MepbGB iIluLoEEBan8zHVbiNA_1W9C0_1ohimOKx5KUjQgBAwLEI6u_1ggaCgoICAESBAdH1eQM,isz:m,sur:fc&tbm 5 isch&source 5 lnt&sa 5 X&ved 5 0 ahUKEwifr_fD7rDeAhWPzoUKHZDGDmcQpwUIHg&biw 5 1024&bih 5 649&dpr 5 1#imgrc 5 kyo466WhoMOvoM:

Ecology and predation Chapter | 6

The case of the Canadian lynx and the snowshoe hare By studying the ecological relationships of a natural population in which the predatory species was the Canadian lynx and the preferred predated species was the snowshoe hare, we were able to demonstrate with real statistical calculations the exactness of the prey predator mathematical model. The number of lynx and hare skins caught and then purchased by the Hudson Bay Company in Canada from 1855 to 1935 for the American fur market was counted. From the graph that resulted using the data collected in all these years it was found that the population of hares and of lynx in a period of 80 years had shown constant multiannual numerical fluctuations about every 7 9 years, according to this mechanism: G

G

The years of greater abundance of hares are followed by years with a gradual increase in the number of lynxes; The maximum number of hares allowed by the conditions of overexploited vegetation is reached, followed

G

G

315

by a period of 7 9 years, during which hares gradually decrease, causing a consequent decrease in the lynxes; The decrease in hares restores the good condition of the vegetation; This is why in 7 9 years the number of hares increases again and the cycle of fluctuations continues.

Chapter 6.2 Predation: hunting methods and techniques The big cats and their hunts to catch prey are one of the most evocative and exciting natural shows that safaris in the great national parks of the world have to offer. Much of the tourist attraction to these parks is based on the opportunity to witness these moments of true wilderness. Countries such as Kenya, Tanzania, India, and South Africa know this well, and they base part of their economic-productive system on those parks where hunting parties of big cats are the primary tourist attraction.

Royal tiger photo safari in India. https://upload.wikimedia.org/wikipedia/commons/d/d1/Royal_Bengal_Tiger_Kanha.JPG; https://upload.wikimedia. org/wikipedia/commons/4/47/Tiger_couple_of_kanha.jpg

A lion and lionesses cross the track in front of a jeep. Lionesses: https://upload.wikimedia.org/wikipedia/commons/0/0b/Serengeti%2C_Tanzania_% 282336929627%29.jpg. Lion: https://upload.wikimedia.org/wikipedia/commons/5/56/Game_Tracker_on_Safari.jpg

316

Felines of the World

Lions and lionesses of the same pride collaborate to try to overcome one of Africa’s biggest and most dangerous prey: the Cape buffalo. https:// upload.wikimedia.org/wikipedia/commons/1/14/Lions_hunting_Africa.jpg

A lioness that has just killed a zebra; sometimes lionesses hunt alone, but more often the lionesses of the same pride collaborate in the capture of prey. Photo courtesy V. Martegani.

Felids: the most skilled world’s predatory carnivores Their powerful muscles and their formidable hunting weapons, with sharp claws and jaws with long teeth (see

Chapter 3: Zoogeography of felids) are just some of the anatomical features typical of the harmonious bodies and supple movements of felids. As large as tigers or as small as cats, felids enchant observers, but they are lethal carnivores for their prey.

Ecology and predation Chapter | 6

Throughout the terrestrial animal world, only the lion and the tiger are able to take down prey the size of a buffalo or a young elephant. Meanwhile other cats, such as the leopard and the puma, even if smaller, can still kill prey whose weight is two or three times higher than their own and also have the strength to hoist them into the trees or drag them to safe places sheltered from other larger predators (other large felids, bears, etc.) or those who, by hunting in packs (wolves and spotted hyena), may seize them. It has been documented, for example, that quite often the cheetah must escape, leaving their fresh prey to lions, in order not to become prey themselves (intraguild predation is quite common among carnivores, and in felids in particular).

Relationship between dimensions of the predator and the prey The anatomy of the felids has remained remarkably unchanged over the last 20 million years, illustrating a well-known proverb: “the winning model does not change.” The basic anatomical model of felids is in fact similar for all species, even if the difference in size sees a range of body weight that can vary up to almost 200 times: from the small rusty spotted cat (Prionailurus rubiginosus) the smallest existing feline, with a weight of about 1.6 kg, up to the mighty tigers of the Siberian region of Amur, whose most majestic males can reach 300 kg. This conformance of proportions and anatomical resemblance (also due to the genetic affinity present throughout the family) contrasts with the great variability in size, and is probably due to the selective pressure that allowed the felids to occupy, always as hypercarnivorous predators or “obligate carnivores,” a large variety of ecological niches especially characterized by the size and variability of prey. The variety in the dimensions determines the type of prey on which the different species feed: big cats (more than 40 kg), medium cats (from 10 to 40 kg), or small wildcats (less than 10 kg) (sec Mattern and McLennan, 2000) in general

317

predate large, medium, or small prey, however this rule is not always respected, and exceptions are quite common. It is clear that in relation to size and mass, the energy needs of the three categories of cats are different, but even here there is not a linear relationship. We know that the smallest species have high metabolic needs (proportionally higher than the larger ones), but some species of small cats (e.g., the black-footed cat) have an even higher metabolism than one might expect. However, this parameter probably does not affect the mass of the prey but rather its quantity. As for large cats, it has been shown that the large captured prey provide a greater supply of energy but are not always convenient because the energy used to capture and defend the carcass from other carnivores is doubled (Macdonald and Loveridge, 2010). We have already seen how many species of felines are opportunistic and hunt what they find and/or manage to capture based on the availability of prey, their predatory skills, and the defensive abilities of their prey. Often the prey is chosen according to their quantity and presence in the territory (available biomass), which can vary seasonally (such as the wildebeests and zebras in the Ngorogoro-Serengeti-Mara ecosystem or reindeer and caribou in the tundra borealis, or the herbivores of the seasonally flooded environments such as the Llanos and the Pantanal of South America). The abundance of a certain prey species in a poor prey environment is adapted to the nourishment needs of the predator, and determines close ecological relationships between predator and “favorite prey,” as happens for example: G

G

G

between the Canadian lynx and snowshoe hare in the tundra and taiga of North America; between the Iberian lynx and wild rabbit, in the Mediterranean woods and the xerophyla bush of the Iberian peninsula; between the Chinese steppe cat and pika in the highaltitude steppe of central China. The variability of weight (and size) between the species of the Felidae family is very wide; the weight of the Siberian tiger that reaches 260 325 kg is about 180 200 times that of the smallest species among the Felidae, with the rusty spotted cat weighing only 1.5 kg.

318

Felines of the World

The variability of weight among the Panthera lineage species is among the highest; the weight of the Siberian tiger reaches 260 325 kg and is about 12 13 times that of the smaller species of the Panterinae, the two species of clouded leopards (25 kg).

The variability of weight among the species of the lineage Puma is probably the highest among all the various lineage of the felines; the weight of a puma reaches 80 120 kg, which is 13 15 times that of the smallest species, the jaguarundi (7.6 kg).

Ecology and predation Chapter | 6

319

The jaguarundi (Puma yagouaroundi) is the smallest species of the puma lineage. It presents various color phases from dark gray, almost black, to red. In this photo is a specimen of the gray phase. This species feeds on prey weighing about 1 kg but often catches prey of 2 3 kg. Photo by M. Korinek.

Modified by Macdonald, D.W., Loveridge, A.J., 2010. Biology and Conservation of Wild Felids. Oxford University Press.

320

Felines of the World

As we can appreciate from the graph, it can be seen that the relationship between the size of the predator and the prey is not always in a linear relationship of direct proportionality. 1. Small wildcats such as the black-footed cat and flatheaded cat, with sizes between 1.5 and 2.5 kg, feed on small prey; but the rusty spotted cat that has similar size, feeds on larger prey, almost the same size as those of the serval, which has a much higher weight, around 15 kg (reaching 18 kg), but it feeds mainly on small mice (Arvicanthis, Meriones, Lemniscomys, etc.). The manul or Pallas cat feeds on hares, pika. and rodents, prey larger than those of the serval which is three times heavier, but the biomass of the small rodents that the serval feeds on is enormous, and for this feline is an excellent source of food to draw on easily, and with a minimal presence of other dangerous competitors. Its anatomy and hunting behavior have evolved to fit into this ecological niche. 2. The fishing cat is a striking example: although it can exceed 15 kg, its prey (fish, amphibians, and crustaceans) are the same size as those of the flat-headed cat that has the same type of diet but weighs at most 2.5 kg. 3. Also the ocelot, leopard cat, and jaguarundi feed on prey of the same size, but the ocelot is much larger than the other two (18 kg against the other two at about 7 kg).

4. The jungle cat (12 13 kg) and the African golden cat (15 16 kg) are about the same size, but the second feeds on much larger prey, which weighs almost 10 times the prey of the jungle cat. 5. The cheetah, Asian leopard, and puma, while being about the same size, feed on prey of different sizes. 6. In the genus Panthera, the jaguar, although very large and able to break down large animals the size of domestic cattle, prefers to feed on smaller species than those of the lion and tiger. 7. The tigers of the island of Sumatra, perhaps because they have only large prey available (Malayan tapir, wild boar, and big deer like the sambar), are the feline, that on average, feed on larger prey, greater than those of the Indian tiger and the lion. The tiger and lion can feed on large prey such as the Indian and African buffalo (but the lion does it preferably by hunting in a pride) but both the tiger and the lion do so with lower frequency, having also available a large amount of prey that is smaller and more numerous, and which make up the majority of the total biomass of their respective ecosystems (mediumsized deer such as axis in Asia or zebras and antelopes such as wildebeest, hartebeest, and gazelle in Africa). The capture of these species also requires an average energy expenditure lower than that required for larger prey.

The jaguar (Panthera onca) has a powerful skull with jaws and teeth that give it a very powerful bite. It is a very skillful hunter and manages to kill dangerous prey such as caimans or large constrictor snakes like boas and anacondas. However, analyzing its food ecology it has been noted that it feeds largely on smaller prey than predators that are of a size similar. https://upload.wikimedia.org/wikipedia/commons/0/0c/ Panthera_onca_at_the_Toronto_Zoo_2.jpg

Ecology and predation Chapter | 6

The fatal bite The variations in size and weight are closely related to the size of the skull and the size of the latter determines the power of the bite, which is essential in taking prey. This power is given above all by the complex of temporal muscles, however the dimensions of the skull do not increase proportionally with those of the body; this is why small cats have proportionately larger skulls that are suitable for attachment of these muscles. On the other hand, large cats, which have proportionately smaller skulls, have had to develop bone crests on the brain case, which serve to increase the attachment surface of large temporal muscles (see Chapter 2: Family Felidae). The force of the bite is therefore mainly due to the size of the skull (of the jaw and of the maxilla) that allow the attachment of the temporal and masseter muscles, of varying sizes, and the strength of the neck muscles. Furthermore, the conformation of the mandible (lower jaw) is very important, with a shape suitable for attaching the masseter so as to maximize the bite mechanism. The bite is facilitated by contraction of the temporal muscles and the masseter, connected to the cranial case and the jaw. By contracting (shortening) these muscles the jaw clamps by rotating around the condyloid process embedded in the glenoid fossa of the temporal bone. This operation is like that of a lever, with the arm (jaw) that rotates around the fulcrum. The power of the bite of felids is also enhanced by the shape of the jaw which optimizes the physical laws that regulate the strength developed by this lever. Comparing the power of the bite according to the proportions of the skull, a greater bite force (proportionally to the size) is evident in the species that have a conformation of the cranial case, the zygomatic arches, and the jaw, allowing the attaching of muscles, temporal and masseter, that are large and well functioning. A wide parietal fossa implies a large temporal muscle and a wide zygomatic arch allows a large and powerful masseter to be contained, but these must also have a suitable space in the right position for conjoining the mandible.

321

In the big cats, for example, even if the shape of the skull is different, that of the jaw is little different. According to Safe and Oliveira (2011), only the size of the skull and the strength of the bite show a correlation with the weight ratio prey/predator. Despite some exceptions, the direct association “big cats stronger bites less constraints in the selection of prey” is the main model followed by the evolution of felids over time, even so, the relationship between the dimensions and the anatomic diversity is not necessarily a linear function. It is only the dimensions that make the difference, while the system used for the bite does not constitute a major element in the capture of prey, which is suitable for the size of the predator. The differences that are found between the various groups of species in the bite mechanics do not show a greater or lesser advantage in the capture of prey and do not represent a particular evolutionary adaptation. For example, among neotropical small wildcats such as ocelot, margay, oncillas (Leopardus), and African and Eurasian ones (Felis) there is a marked difference in the shape of the skull and in the different development and use of the temporal muscle complex or masseters (the latter is more developed in the genus Felis), but this did not determine a different evolutionary success—both groups feed on rodents and other vertebrates of medium or small size, and their different conformation of the skull and the bite mechanism does not correspond to particular adaptations to a specific type of prey. The bite of felids is efficient in catching prey even if the mechanism used changes a little due to the anatomical differences of the different species. The fishing cat (Prionailurus viverrinus) and the flatheaded cat (Prionailurus planiceps) have a very specific diet, based on fish, amphibians, and crustaceans, slimy and slippery prey that need to be captured in water; yet the anatomy of the jaw, the mandible, and the bite mechanism have not changed. The adaptation to this type of prey has only been at the level of the teeth which have undergone changes to make them more suitable for a sure grip, as they are more pointed and have incisors and canines facing forward more than in the other species. Hayward, M.W., Kamler, J.F., Montgomery, R.A., Newlove, A., Rostro-Garcı´a, S., Sales, L. P., et al., 2016. Prey preferences of the jaguar Panthera onca reflect the post-pleistocene demise of large prey. Front. Ecol. Evol., 25 January 2016.

322

Felines of the World

The jaguar (Panthera onca) has a large size, in order of weight and dimensions, placing it third largest of the felids, after the tiger and the lion. It also has a powerful skull that allows it a bite among the most powerful; yet comparing the size of its prey with that of the prey of other species of solitary hunter (the lion that can hunt in a group is excluded) it is noted that this species prefers smaller prey, that is, in relation to the size of the predator the ratio size predator/size prey, is in favor of the first ( . 0.5); in the tiger it is almost identical (1), while the snow leopard (1.3) often preys on species that exceed its weight. This illustrates that the diets of large Felidae are mainly determined by behavioral and ecological factors and in this case by the abundance of a particular type of prey. The jaguar that lives in high biodiversity habitats is attracted to the abundance of prey and prefers those living in herds, regardless of size. The snow leopard that lives in mountain environments, not rich in prey, easily finds only large ungulates like wild sheep and ibexes that are often heavier than itself.

Felines: not only solitary ambushes Almost all felids practice solitary ambush hunting, following sequences of similar behaviors for each species. For this reason, within the family, adaptive radiation concerning hunting is very minimal, and the evolutionary specializations of anatomy and behavior, which deviate a

little from this traditional technique, are very few, however there are other methods and other hunting techniques, that can be summarized in four groups: 1. A single species is a true “runner hunter” who pursues prey on medium long distances, 300 500 m, too far for other felids: this is the Cheetah (Acinonyx jubatus). 2. A single species has adopted “cooperation in group hunting” and usually lives in packs called “prides”: the lion (Panthera leo). Also at the base of the hunting techniques of the lion there is always the ambush, but in this social species there is coordinated hunting among the various members of the pride. 3. There are four species of real “expert climbers,” arboreal with remarkable scansorial ability. They often hunt in the branches of large trees and capture arboreal prey such as monkeys, squirrels, and birds. These include the margay (Leopardus wiedii), with unique hunting techniques, even based on the imitation of prey recalls, the marbled cat (Pardofelis marmorata) and the two species of clouded leopard (Neofelis nebulosa and Neofelis diardi). 4. Two species use “fishing with a largely piscivorous diet” and ambushing close to water and, mostly hunting, they “fish” for mainly fishes, amphibians, and crustaceans: these are the fishing cat (P. viverrinus) and the flat-headed cat (P. planiceps); the latter, like the otter, chases fish, plunging and swimming under water. (A) The ocelot is not a specialized feline that adopts particular hunting techniques, it tends to ambush both on the ground and from trees. It can catch monkeys, squirrels, coati (Nasua sp.), opossum of various species and arboreal anteaters (Tamandua sp., Cyclopes sp.) among the branches, as it has, like most small and medium-sized cats, good agility even among branches (medium scansorial ability); however it uses the lower branches more often to launch ambushes and to hunt prey on the ground. (B) Ocelots also live much of the time on the forest floor where they ambush their usual prey, such as young or little cervids like Mazama, rodents like agoutis (Dasyprocta and Myoprocta sp.), paca (Cuniculus 5 Agouti paca), and armadillos (Dasypodidae), which are all terrestrial species. (A) Photo courtesy of V. Martegani.

Ecology and predation Chapter | 6

323

A leopard and a domestic cat in ambush. From large to small cats the most common hunting technique is the ambush, with which felines are able to sneak up to their prey and, with a quick dash catch them, sometimes after a short chase. Leopard: https://upload.wikimedia.org/wikipedia/commons/4/ 47/Leopard_%28Panthera_pardus_pardus%29_stalking.jpg Cat: https://upload.wikimedia.org/wikipedia/commons/6/6a/Catstalkprey.jpg

6.2.1 The most common technique”: the ambush

“hunting

Therefore, in addition to their anatomy, the methods of hunting techniques also do not vary much within the Felidae family. All are focused on detection (sighting), approaching, and ambush of prey; arriving at the right distance, allows the pursuit, capture and killing and, finally, the fulfillment of the purpose of this entire sequence of behaviors: feeding on the prey. This sequence represents the hunting-base method for felids of any size and the duration of the chase in terms of distance varies according to the size of the species, from a

few meters for species of small wildcats up to a few tens of meters in the larger ones; while many canines have physiological adaptations that allow long pursuits in terms of space and time, the muscles of felines allow performance with very fast sprints but which last a short time because of the enormous “energy needs” to put them into action. Lions and tigers reach 80 km/h in a few seconds but abandon the pursuit if, after a few tens of meters, the prey is not reached. This is why most cats have adopted special behaviors, flattening themselves on the ground, hiding in vegetation, and approaching, slowly, as close as possible to the prey without being seen, before unleashing their lightning attack.

Caracal in ambush in open habitat: the muscles are ready to sprint, the senses are alert, and the eyes are focused on the prey. https://www.shutterstock. com/it/image-photo/super-predator-caracal-moves-through-african-33734230?src 5 ar7ApYYlDqzkH7Eh-JYDUQ-1-62

324

Felines of the World

The sequences of predatory behavior used by felids are common to most species.

This diagram shows how in the felids the morphological adaptations that have evolved to favor hunting are used for different purposes in the hunting and feeding sequences. Modified by Macdonald and Loveridge.

Ecology and predation Chapter | 6

6.2.2 The lion: hunting with the cooperation of members of the “pride” Savannahs are environments with a rather high biodiversity, and their herbaceous vegetation supports a community of very diverse herbivores; speaking only of African savannahs we have some species of gazelles, many of antelopes, buffaloes, elephants, zebras, giraffes, warthogs, and almost all these species have social habits and live in more or less numerous herds. The lion, a typical species of the savannah, has adapted to this habitat and is part of it as a superpredator. It is a skilled hunter that exploits the rich animal biomass of this ecosystem, and has evolved its behavior, unique among all felids, because it has

325

adopted social habits and therefore lives in herds (clans or “prides”), which are composed of up to 20 30 members. As G. Shaller wrote (1973) “The entire social system of lions is undoubtedly an adaptation to hunt large prey in open terrain.” The members of the pride are lionesses all from the same families and some male lions, coming from other prides and living together. After going through long traineeships the young learn to communicate, and after many months of experience also develop the ability to coordinate in predatory activities. These skills represent an important evolutionary success, with lions hunting alone having about a 17% chance of success, while when hunting in a pride the odds increase to at least 30%.

When hungry, the lionesses, also followed by the males, begin “rituals of hunting involvement.” The members of the pride begin rubbing each other head to head and side by side, roaring, coughing, licking each other’s mouth and their own mouth (picture on the left), before setting off through the savannah in search of prey (photo on the right).

The lion is a fast hunter over short distances, sprinting very fast and after a few seconds reaching 80 km/h but it can only maintain this speed for 20 30 m, as his heart, which must work hard for such a powerful and conspicuous muscle mass, cannot withstand overly prolonged exertions. Many of its prey, including zebras, gazelle, and other antelopes, can easily outrun it over longer distances. This is why the lion is forced to carry out an ambush that brings it closer to its prey. Cooperation in hunting has at least two purposes: 1. If the prey is solitary and as large as a Cape buffalo, the lions advance in a group divided into three prongs (center, left and right) surreptitiously surrounding

the prey and then assailing it simultaneously from several parts. 2. If instead the prey is a herd of fast herbivores like gazelles and medium-sized antelopes, some members crouch in the tall grass or behind shelters; others circumvent the herd, overcome it, and feign an attack by chasing the gazelles toward lions lurking in ambush that, at the right moment, leap out and take down the prey that have come within range. When applying this technique lions are also able to assess the wind direction, and arrange themselves in such a way that the lurking members are leeward, so that their smell is not perceived by the excellent olfactory abilities of the herbivores.

326

Felines of the World

(A) The lions beginning the ambush approach. (B) Eventually the prey is surrounded. (C) When the prey is surrounded by members of the pride, and has no escape routes, it is approached and a coordinated attack is launched so that the lions surround and bite the prey on the rump and hind legs, trying to spread it on the ground, while one of the lions is in front, often the strongest and most experienced male, and attempts to bite the throat (trachea) or nose (in a grip called the kiss of death) and hence suffocating the prey. (B) By Stander, P.E., 1992. Cooperative hunting in lions: the role of the individual. Behav. Ecol. Sociobiol. 29.

This photo captures a very significant moment in the social life of lions. A wildebeest has been surrounded by lionesses and some young pride members who cooperate in hunting; this is how females teach the young communication and collaboration in hunting activities; aided by a certainly innate predatory instinct, the young lions learn and, more importantly, start building their indispensable experience. Photo courtesy of F. Veronesi.

Ecology and predation Chapter | 6

327

Diagram illustrating the technique of cooperation in the ambush and hunting by a group of four lionesses of the same pride, of a herd of gazelles, according to a study by Georg Schaller in the savannahs of the Serengeti Park (Tanzania). Modified from Schaller, G.B., 1973. Golden Shadows, Flying Hooves. Alfred A Knopf, New York.

6.2.3 The cheetah: long high-speed chase

Between Kenya and Tanzania, in the ecological complex of the Ngorogoro-Serengeti-Mara (named after the three most famous parks of this ecosystem), a cheetah chases a large male Grant’s gazelle (Gazella 5 Nanger granti) and a Thomson’s gazelle (Gazella 5 Eudorcas thomsonii), under the gaze of some white-bearded gnu or wildebeest (Connochaetes taurinus albojubatus) on high alert.

328

Felines of the World

Among the felines, the cheetah is the only species that is truly suitable for daytime hunting based on pursuit over fairly long distances; this species has evolved a specialized anatomy that allows exceptional performance in a fast race: in just 3 seconds it reaches an acceleration of 90 km/h and after 20 seconds it can reach and exceed 110/115 km/h (about 30 m/s), performing a very fast series of leaps of 6.9 m. This makes this species the fastest land animal in the world. This feline has a somewhat singular appearance due to the length and subtlety of its limbs, its small head and very wide chest, which are determined by extraordinary adaptations to running, which particularly involves the skeletal, respiratory, and circulatory (heart) systems. In the skeleton there is considerable lengthening of the bones of the limbs that allow the cheetah a wide stride and very long leaps; in the legs the nonretractable claws (in adults) improve the grip on the ground. The vertebral column is extremely flexible and, during while in motion, it flexes alternately, giving a highly concave curvature when the legs are gathered and a convex curvature with extended legs; in this way it works like a spring that pushes the body forward.

In the muscles of the musculoskeletal system, among the muscle fibers there is a majority of those with rapid contraction that allow lightning reactions and very rapid movements. In the respiratory system the large lungs can contain a lot of air (oxygen) inhaled in large quantities with each breath through the very wide nasal choana. Oxygen is indispensable for the muscles in order to produce energy. In the circulatory system the heart has enormous dimensions and quickly pumps oxygen-rich blood to the muscles engaged in the effort of the pursuit. After a long chase, the consumption of oxygen and energy is very high, so that the heart beats greatly increase their frequency and the breaths go from 15 per minute, in normal resting conditions, up to 150. For this reason after the capture of the prey the cheetah is forced to pant and remain still to regain normal breathing. In addition, the long, strong, and muscular tail is used to balance its weight and maintain equilibrium in the sudden changes of direction during the pursuit of agile prey (gazelle) that run in a zigzag to confuse the predator. Even the pursuit of prey by the cheetah, however, never continues beyond 400 600 m and is more often concluded if the prey is not reached in 100 200 m. https://www.shutterstock.com/it/image-photo/cheetah-acinonyxjubatus-running-south-africa-99403178

In the two pictures of cheetahs running, one can see how the spine alternately assumes curved and concave positions; moreover, you can see the length of the stride due to the lengthening of the spine in a concave position and the distension of the very long front and back limbs. In this position the cheetah at high speed can make leaps of 6 m. https://www.shutterstock.com/it/ image-photo/cheetahs-focusing-436208863

Ecology and predation Chapter | 6

329

https://www.shutterstock.com/it/image-vector/set-vector-drawings-on-theme-predators-724882663?src 5 YEwwve8CO7Jhiqc--_H9XQ-1-21

The wide stride of the running cheetah is enabled by the great mobility of the scapula, shoulders, and pelvic girdles that are very mobile, and with the help of the flexible spine. During the chase the breathing is also facilitated by the movements of the race: (Top) The organs are compressed and the lungs expel air (expiration) ---; (Bottom) The organs are stretched and the lungs dilate by inhaling air (inspiration) ’’’. https://en.wikipedia.org/wiki/Giant_cheetah#/media/File: Acinonyxrunningrespiration.png

330

Felines of the World

A cheetah about to launch its paw at a young wildebeest. https://www.shutterstock.com/it/image-photo/cheetahs-focusing-436208839? src 5 1rXivLvrWJumdBRN8gEQOw-1-33

To take down prey the size of a gazelle, the cheetah uses a particular technique. When chasing the prey and closing to the right distance, it launches a strong paw, hitting the prey on the thighs or buttocks and wounding it using the large hooked nail (dew claw) present on the first toe of the front legs. In this way the victim loses its balance, falls, and is held by the throat and suffocated within a few minutes. Modified from De La Fuente and Lalanda.

6.2.4 The fishing cat (Prionailurus viverrinus) and flat-headed cat (Prionailurus planiceps): fishers, specialized hunters in wet habitats

These two species of cats with piscivorous food habits are excellent swimmers and have partially palmate front legs and long nails and hooks that, even if they are retracted, always stick out with their tips from their casing. They tend to ambush primarily fish (76% of their

diet), but also amphibians and small rodents, on the shores of water basins in the wet environments they frequent: marshes, reed beds, water courses in wet or flooded forests, and mangrove formations, but also rice paddies typical of the regions of Asia where they live.

Fishing cat in mangroves habitat of Sundarbans, West Bengal, India. https://upload.wikimedia.org/wikipedia/commons/0/02/Fishing_cat_amidst_ mangroves.jpg

Flat-headed cat. Photo by J. Pfleiderer.

Even other cats, however, without being so specialized, enjoy hunting along the shores of aquatic environments. Serval (Leptailurus serval) are known to catch fish and amphibians; and very often the jaguar, and also the tiger, can capture aquatic prey (big fish, crocodiles, and

big snakes) because they are good swimmers; besides hunting near the water, or tending to ambush where prey drink, many cats chase them even where the water is deep.

332

Felines of the World

The narrow and elongated skull of these two species shows a dentition with pointed incisors and canines protruding forward; the row of mandibular teeth is well developed with premolars and molars with very pointed cusps. This specialized toothing serves to grasp and retain fish, prey that can slip away as they are often slimy.

The fishing cat in ambush on the high margins of a river. https://upload.wikimedia.org/wikipedia/commons/5/58/Fishing_cat_in_San_Diego_Zoo. jpgu06-01-9780128165034

Ecology and predation Chapter | 6

333

Where they live Fishing and flat-headed cats live in southern Asia; both species are very elusive and live in fairly inhospitable environments. For this reason they are little known, and even less studied. Sometimes even the people living in the areas they frequent do not know of their existence. They have a very discontinuous distribution, divided into restricted areas which therefore contain a very fragmented populations. Fishing cat (P. viverrinus), South India, Sri Lanka, Pakistan, Nepal, Indochina, Malacca Peninsula, Java island (but intermittently and in restricted areas) (see Chapter 5.6.3)

https://www.shutterstock.com/it/image-photo/fishing-cat-prionailurus-viverrinus-wildlife-animal-529473250?src 5 mxkvT9WIn-lFCc0CmCy2tA-1-49

Flat-headed cat (P. planiceps)Malacca Peninsula, Sumatra islands and Borneo (but intermittently and in restricted areas) (see Chapter 5.6.4)

Photo by J. Pfleiderer.

A sequence in which a fishing cat dives into the water to catch the fish it feeds on. The hunting method of the fishing cat is a little less tied to the water than that of the flat-headed cat. This species carries out its ambush near water which is not too deep and, with a partially webbed paw, catches the fish that get close; it often dives into the water but rarely dives in completely. On occasion, it also captures small animals up to the size of a mouse-deer (Tragulus) or a chital (Axis) or muntjak (Muntjacus) fawn. Sometimes it found on farms where it may kill poultry or young goats. https:// www.shutterstock.com/it/image-photo/fishing-cat-prionailurus-viverrinus-hunting-swimming-623208596?src 5 mxkvT9WIn-lFCc0CmCy2tA-1-69; https://www.shutterstock.com/it/image-photo/fishing-cat-prionailurus-viverrinus-hunting-watching-1240259224?src 5 mxkvT9WIn-lFCc0CmCy2tA-1-

Even more than the fishing cat, the flat-headed cat dives completely in and swims underwater to catch fish, as do other Carnivora species such as otters, some mongooses, the aquatic genet (Genetta piscivora), and perhaps the otter civet (Cynogale bennettii).

6.2.5 Margay, marbled cat, Indochinese and Sunda clouded leopards: arboreal climbing hunters This group of specialized arboreal hunters includes a species from central and southern America: the margay (L. wiedii), and three Asian species: the marbled cat (P. marmorata), the Indochinese clouded cat (N. nebulosa), and the Sunda clouded leopard (N. diardi). These are four species of forest felines, true expert climbers that have excellent scansorial abilities, thanks to their particular anatomy; they all have a very long and muscular tail that they use to balance and maintain themselves in equilibrium in the acrobatics they perform while running on branches. They demonstrate an extraordinary

sense of balance and can walk on thin branches without falling, like a tightrope walker. All have skeletal adaptations that make the ankles of the hind legs flexible enough to be able to rotate 180 degrees; moreover the margay and the clouded leopards have modified and lengthened phalanges and metacarpals, in order to make the legs wider and larger, this allows them to grasp with the nails of their hind legs to the bark and to come down from almost smooth trunks upside down, or to remain hanging in the void, clinging to a branch with only one leg. These species exhibit an extremely mimetic fur, variously spotted with disruptive colors and patterns, which imitate the light/dark of the play of light between leaves. They hunt, mostly on trees, arboreal prey such as monkeys, squirrels, birds, and lizards.

Ecology and predation Chapter | 6

335

A young Margay specimen showing its distinctly arboreal habits. https://upload.wikimedia.org/wikipedia/commons/3/32/Margaykat_Leopardus_ wiedii.jpg

Margay in the forest of French Guyana (Leopardus wiedii vigens) climbs on a tree trunk. https://www.shutterstock.com/it/image-photo/wild-margaycat-leopardus-wiedii-natural-1180042297?src 5 gWMlnedkH81pgJY_HqthXw-1-12

336

Felines of the World

The Indochinese clouded leopard spends a lot of time in trees where it often hunts monkeys, squirrels, birds, reptiles, etc. The pattern and colors of its beautiful fur are an example of disruptive mimicry between the light and dark of the fronds.

A marbled cat (Pardofelis marmorata) descends upside down from a tree trunk. The particular shape of the ankles and legs of arboreal felines represent well the phenomenon of evolutionary convergence due to adaptations to the same type of habitat, that of tropical forests, and to the same type of food ecology, hunting for arboreal prey. These adaptations allow them to perform with extreme agility on the branches and trunks of large trees. https://upload.wikimedia.org/wikipedia/commons/3/3f/Marbled_cat_ borneo.jpg

The margay is perhaps the most skillful arboreal feline. It presents adaptations of joints and bones of the legs so that is can pause and dangle in the void. With these anatomical adaptations and with other behavioral systems they manage to prey on even the most agile primates of the South American forests. According to de Oliveira Calleia et al. (2009) the margay is able to lurk in branches and emit recalls similar to those of the wild pied tamarin (Saguinus bicolor) young, a small monkey, when in trouble. These sounds attract adults who come to the rescue and are captured by the feline.

Ecology and predation Chapter | 6

337

338

Felines of the World

It is commonly known that many species of Felidae are quite skilled at tree climbing, think of the leopard which often tends to lurk in trees and swoop on prey from above and, after it has killed it, hoists it onto the branches to keep it from other carnivores. The evolutionary history of the felids is linked to an arboreal life, since their first appearance, there were also arboreal ancestral species, such as Pseudaelurus and the first true fossil cat known, Proailurus lemanensis, and another more recent ancestor, Styriofelis lorteti. Reconstructions of Proailurus (left) and Styriofelis lorteti (right). Courtesy of M Anto`n.

6.2.3 Caracal and serval: masters of agility in the ambush final leap

(Left) Serval (Leptailurus serval), (right) caracal (Caracal caracal). Both have large ears and excellent hearing. https://upload.wikimedia.org/wikipedia/commons/4/43/Caracal_hunting_in_the_serengeti.jpg

Ecology and predation Chapter | 6

The caracal (Caracal caracal) and serval (L. serval) are close relatives as they belong to the same evolutionary line (caracal lineage). Both are excellent hunters of grassy savannahs, but the caracal also frequents semidesert areas, while the serval prefers more humid areas, including forests. Their anatomy reveals a strong back train and elongated limbs; the hind legs have a powerful musculature that allows them to take sprint quickly and make very high jumps. They are solitary hunters that especially catch rodents, hares, and small antelopes both by day and night; the final jump is their specialty and reveals a particular

339

ability to catch birds even after they have managed to rise in flight. The serval, equipped with huge ears, captures the rustle of small rodents and stands out for being able to jump precisely on hidden prey. The caracal and serval have very large and mobile earbuds that help their very fine hearing, allowing them to hear even the slightest rustle of a small animal among the tall grasses of the savannah.

Serval (Leptailurus serval)

The typical jump of a serval. This species, after the black-footed cat, is probably the most effective hunter among all felids, it is successful in capturing prey in 50% of attempts. In comparison, the leopard is successful in 38% of cases and the lion is successful in only 20% 30% of attacks. https:// www.shutterstock.com/it/image-photo/serval-leaping-serengeti-tanzania-africa-57612052?src 5 2jUbvysnmyN_UjSslZoD9A-1-0

340

Felines of the World

In this drawing is illustrated the typical jump of the serval when capturing a zebra mouse (Muridae) of the genus Lemniscomys. The serval hunts various prey (from frogs to small antelopes) but is specialized in catching small rodents and generally prey that does not exceed 200 g, which make up a large part of its diet. The savannahs have a rich biomass of small rodents (Meriones, Achomys, Lemniscomys, Arvicanthis, and many other genera). The small prey are almost always detected with the hearing and captured with this typical jump (up to 4 m long by 2 of height) with which the feline lands on the front legs accurately on the prey. They also stand out for making 3 m high jumps to catch birds in flight.

Caracal (Caracal caracal)

The caracal has a very rapid reaction speed, and perhaps making the quickest acceleration (over a short distance) among all the felids; moreover it has hooked claws that can catch prey and not allow it to escape. These skills, combined with its agility, qualify it as an excellent hunter. It can jump three meters in height and brings down large birds while flying. It has a diet composed of prey more varied and larger than those predated by the serval, most however with a weight of less than 5 kg, but sometimes managing to kill also antelopes the size of a gazelle ( , 50 kg). Modified from de La Fuente and J. Lalanda.

Ecology and predation Chapter | 6

341

The steenbok (Raphicerus campestris) (right) is one of the most common small species of antelopes (7 15 kg) preyed on by the caracal, which prefers young or small females. In its vast African area, prey also include dik-dik (Madoqua sp.), oribi (Ourebia), klipspringer (Oreotragus), young gazelles, etc. The caracal has a very varied range of prey in both size and species, so it is a very adaptable species and this has enabled it to colonize many habitats and a large distribution area that, in addition to much of Africa, includes the whole of the Middle East to India; in Asia prey include some species of gazelles, but mostly hyraxes, rodents, and hares. https://upload.wikimedia.org/wikipedia/commons/d/d4/Raphicerus_campestris_male_%28Etosha%2C_2012%29.jpg

Cheetah and caracal taming for hunting. https://upload.wikimedia.org/wikipedia/commons/e/e6/Kurt_boeck_indien_nepal_144B.jpg

342

Felines of the World

Sometimes in India, until the beginning of the 1900s, not only the caracal and the cheetah were domesticated for hunting but, as can be seen from the photo, even the leopard https://upload.wikimedia.org/wikipedia/commons/d/da/Cheetah_on_cart%2C_used_for_hunting_in_India_%28probably_early_20th_century%29.jpg?uselang=it