Chapter 3

Zoogeography of felids

Chapter Outline
3.1 The zoogeographical regions 75 The great eruption of the Toba volcano in Sumatra 86
South-Eastern Asia: a biodiversity hot spot for safeguarding 3.2 The origins of the current distribution of felids around
some rare species of felids 83 the world 87
The Felids of the Indochinese Subregion and Sundaic 3.2.1 The most Ancient migratory wave 87
Subregion 84 3.2.2 The subsequent migratory waves and extinction
New species in the Sunda Archipelago 85 of the megafauna 89

The subject of zoogeography or biogeography attempts G The reconstruction of the ancient disposition of the
to give an answer to the current distribution of the emerged lands that continually varied with the sea
animal species, studying their diffusion and displace- level that increased or decreased as a result of climatic
ments of the past with the help of the following variations and caused the formation of intercontinental
parameters: bridges.
G The location of fossiliferous sites and records studied Today, genetics also helps in the study of zoogeogra-
by paleontology; phy: both by studying the DNA of current species and that
G The study of paleoclimatic variations (alternations of of fossils it is possible to understand that the presence of a
glacial and warm periods) that influenced migrations species in a particular place on Earth is the result of
or caused extinctions of whole taxa; genetic, ecological, and historical causes (phylogeography).

These two species of lion, now extinct, lived in Eurasia and the Americas, where there are no longer any lions. Both had woolly fur to protect them-
selves from the cold climate of the period in which they lived. Their manes were not as abundant as those of current lions, and their fur was probably
spotted. https://upload.wikimedia.org/wikipedia/commons/c/cb/Panthera_leo_atrox_Sergiodlarosa.jpg.

Felines of the World. DOI: https://doi.org/10.1016/B978-0-12-816503-4.00003-9

© 2020 Elsevier Inc. All rights reserved. 73
74 Felines of the World

Distribution area of the current lion and two species of extinct lion, according to the fossil record. Mitogenomics of the extinct cave lion, Panthera spe-
laea (Goldfuss, 1810), resolve its position within the Panthera cats
Barnett, R., Zepeda Mendoza, M.L., Rodrigues Soares, A.E., Ho, S.Y.W., Zazula, G., Yamaguchi, N., et al., 2016. Open Quat. 2, 4. https://upload.
wikimedia.org/wikipedia/commons/8/81/Cave_lion_range.png.

The distribution area of lions in late the Pleistocene

is indicated by both fossil finds and the cave paintings
left by our ancestors, which tell us that up to about
10,000 15,000 years ago some species of lion were dis-
tributed around almost the entire world, while their pre-
decessors (Panthera fossilis and Panthera shawi) were
already extinct.
About 1.8 Mya the separation between current
lions and the cave lions occurred; the latter are there-
fore to be considered as a separate species, not a sub-
species of the current lion. From Eurasia, the cave lion
and the Beringian cave lion (Panthera spelaea and
The current lion is now divided into just two subspecies, a northern one
Panthera vereshchagini) reached the northern part of (Panthera leo leo) that once lived in North Africa, the Middle East, and
North America through the Bering bridge. However, P. which today is also found in India (Gir forest where, however, perhaps it
fossilis had previously arrived in North America where was first imported?), and another (Panthera leo melanochaita) from East
it originated the American lion (Panthera atrox). This Africa to southern Africa (at one time it reached the Cape of Good Hope
species, known through many fossils, had first colo- but today it is extinct in almost all of South Africa). Evolved from the
lions of Laetoli in Tanzania (perhaps P. shawi, 3.6 Mya) they had spread
nized North America and then crossed the isthmus of throughout Africa, Turkey, the Middle East, India, and perhaps southern
Panama, according to the latest findings of fossil Europe. Clashes with the first Mediterranean civilizations and then the
remains, and reached Chilean and Argentine Patagonia massive anthropization of Africa greatly reduced their original range.
(Chimento and Agnolin, 2017). https://www.shutterstock.com/it/image-photo/lion-4-half-years-panthera-
leo-21140812?src 5 TXN1T681JqV2gpFmVcmL4Q-1-29.
 Zoogeography of felids Chapter | 3 75

3.1 The zoogeographical regions As regards the mammals that characterize these zoo-
geographic regions, the longer the isolation of a region
The various terrestrial continents have been divided into has been, the more numerous its endemisms are; as an
zoogeographic regions and subregions, based on the species example, the Australian region, being an island and
(especially of vertebrates) that inhabit them. Not all experts having almost never had any wildlife exchanges with
agree on the number of regions and their exact boundaries, other continents, has evolved a group of mammals, the
many considering some areas of faunal transition between marsupials, which are almost all endemic.
one region and another, while others consider these transition The Afrotropical or Ethiopian region has some endemic
zones as true zoogeographic subregions. Each region is char- mammals such as the Giraffidae, the Hippopotamidae, and
acterized and defined by a number of orders, families, gen- some prosimians: the Galagonidae family, the monkeys of
era, or species that are exclusive and typical and which are the genera Cercopithecus, Cercocebus, Mandrillus, the
called “endemisms” (from the Greek: indigenous, native). Colobidae and many others, and among the great apes
The Felidae family has colonized almost all zoogeo- (Antropoidea) the gorillas and chimpanzees. Typically of
graphic regions: Afrotropical origin (although some doubts have recently
1. Palearctic region (Eurasia, including most of China, arisen about the exact origin of some Families, not exclu-
North Africa, Middle East). The central part of this sively African) is the Supraordinal mammalian Clade
region is considered the area of origin of the Felidae, Afrotheria which includes the Families: Tubulidentata
and certainly of the Panterinae, whose oldest fossil so (Orycteropus afer), Tenrecidae (Tenreks), Chrysochloridae
far was found in Tibet; today it is frequented by a (Golden moles), Macroscelidae (Sengis or elephant-
greater number of species (16) than any other region. shrews), Hyracoidea (hyraxes), Proboscidea (elephants),
2. Nearctic region (North America and part of Central Sirenia (dugongs and Manatees).
America). The Palearctic and Nearctic regions, for The Neotropical region has many endemisms such as
many valid ecological and wildlife reasons, are often the Xenarthra (sloths, anteaters, and armadillos) and the
brought together in the Holarctic super region. In monkeys Platyrrhini (Cebidae, Atelidae, etc.), and the only
North America there are only three species of felidae: marsupials that live outside the Australasian region, the
the puma, the bobcat, and the Canadian lynx; how- opossum (Didelphidae), of which one species is the North
ever, further south toward Central America, the num- American or Virginia opossum (Didelphis virginiana),
ber increases with the addition of some South which has colonized part of the Nearctic region.
American or Neotropical species (ocelot, etc.). The Palearctic and Nearctic regions have very few
3. Ethiopic or Afrotropical region (Africa south of the endemisms because for a long time, in the past, during
Sahara and part of the southwestern Arabian peninsula). the great Pleistocene glaciations, they were very often
The large island of Madagascar, which the felids have united by a bridge of emerged lands, where today there is
never colonized, is sometimes associated with this region a sea passage, called the Bering Strait, between Alaska
and sometimes separated into the Malagasy region. and Eastern Siberia. Therefore previously, horses, lions,
4. Indo-Malaysian region (India, south of the Himalayas, jaguars, saber-toothed cats, mammoths, and mastodons
Indochina, the Sunda Islands up to Bali, the Philippines). were widespread in both continents. Some disappeared
In the tropical forests and the big islands of the Sunda during the period of the great extinction of the
live a large number of species of felids of which eight Pleistocene megafauna, which hit the boreal areas bring-
are endemic; therefore this region has a higher number ing about climatic and ecological upheavals which many
of endemic species than any other region. species were unable to adapt to or escape from.
Bali island and Lombok are separated by a sea pas- It is almost certain, however, that the low temperatures
sage, not wide but very deep, which has never allowed due to glaciations, however rigid, took place gradually and
the two islands to unite. Here passes the border line certainly over long timescales, so they were not the cause
that separates the Indo-Malaysian region from a wild- of great extinctions. In reality, in the late Pleistocene, a
life province that also includes the island of Sulawesi series of very severe climatic episodes were recorded with
(formerly Celebes), which reaches up to New Guinea— sudden temperature falls of more than 15 C, which have
this is called Wallacea—in these islands and in the been confirmed as the cause of the extinction of many
Australasian region, there have never been felids. large mammals (Megafauna) in many regions, since such
5. Neotropical region (South America and parts of Central wide and sudden variations did not leave time either to
America, Cuba, and Caribbean islands). This region adapt or to migrate. Climate change could affect the ecol-
presents a large number of endemic species of the ogy of entire regions; for example, the disappearance of a
genus Leopardus, which is exclusive to the region certain type of vegetation could mean the extinction of
except for the ocelot which extends to the north some populations of herbivores, and consequently of the
encroaching into the Nearctic region with the jaguar. carnivores which usually preyed on them.
76 Felines of the World

(Top) Map by Serban Proches and Syd Ramdhani (2012) https://www.researchgate.net/figure/Vertebrate-zoogeographical-regions-and-subregions-as-

derived-from-the-four-analyses_fig1_234563430.

These were the main causes of the extinction of many These maps show two different proposals for the sub-
felids that were once present in Eurasia and today are no division of terrestrial continents, in zoogeographic regions
longer there; among the great predators of the past, a and subregions. Not all authors accept the Andean and
saber-toothed cat (Homotherium) disappeared 28,000 Arctic regions; others such as the Caribbean, the
years before the present (BP), the cave lion (and the North Wallacean, the New Guinean, and the Madagascar regions
American lion) 11,000 12,000 years BP, and even the are often considered to be subregions or transitional zones
current leopard lived in Europe up to 27,000 years BP. between a region and the neighboring region, such as the
 Zoogeography of felids Chapter | 3 77

Saharo-Arabian subregion, often considered a transition (China), also known as the Chinese transition zone,
faunistic zone between Africa and Eurasia. and the Saharo-Arabian subregion, also known as the
The Super Oloarctic region with boreal distribution is Saharo-Sindic transition zone, North Africa, the north-
composed of three regions with many zoological and ern Sahara, and the Middle East. Here live some spe-
ecological-environmental affinities, including: cies or subspecies of very interesting felids, in
populations that are isolated and separated from their
1. The Arctic region, containing Alaska, northern Canada,
sub-Saharan Africa congeners. The famous lions of the
Greenland, and northern Siberia, which not accepted by
Atlas or Barbary lion were hunted almost to extinction
all experts; it has only one terrestrial habitat, the Arctic
at the start of the 20th century. The caracal and the ser-
tundra and the circumpolar glaciers; probably the only
val they survive in Algeria and Tunisia and in some
species of felines whose distribution areas touch the
small areas of the Moroccan mountains, maybe with
merional limits in this area are the Eurasian lynx and
the leopard. In some particularly favorable small areas,
the Canadian lynx in North America.
there is a population of very rare Sahara cheetahs
2. The Nearctic region of North America.
(Acinonyx jubatus heckii?) which some genetically
3. The Palearctic region of continental Eurasia, to
identify more closely with Asian cheetahs.
which must be added the Sino-Himalayan subregion

(A) The canadian lynx (Lynx canadensis) lives in the Tundra and in the forests of the Taiga, in the Nearctic zoogeographic region.
(B) The siberian tiger (Panthera tigris tigris ex altaica) lives in the mixed birch and coniferous threes forests of eastern Siberia in the Palearctic zoogeo-
graphical region. https://www.shutterstock.com/it/image-photo/running-tiger-snowy-furr-wild-winter-625445456?src 5 9J7BV8gnkuCHGAUM03IYVw-
5-78; https://www.shutterstock.com/it/image-photo/canadian-lynx-canadensis-walks-right-through-630052376?src 5 pynzE6piBhns4VpHscLTTQ-3-94.
78 Felines of the World

The Holarctic super region with boreal distribution is cheetahs. In North Africa the Libyan desert cat (Felis
composed of three regions with many zoological and lybica lybica) and the sand cat (Felis margarita) can also
ecological-environmental affinities, including: be found, while in the delta of the Nile in Egypt the only
African population of the jungle cat lives (Felis chaus).
1. The Arctic region, containing Alaska, northern Canada,
Greenland, and northern Siberia, which not accepted by In North America and Eurasia, the same latitudinal
all experts; it has only one terrestrial habitat, the Arctic bands are matched by the same climatic-vegetation charac-
tundra and the circumpolar glaciers; probably the only teristics. From north to south in succession there are: the
species of felines whose distribution areas touch the tundra, the coniferous taiga, the mixed coniferous
merional limits in this area are the Eurasian lynx and and deciduous forests, and broad-leaved forests, grassy
the Canadian lynx in North America. areas (the North American prairies and the Eurasian
2. The Nearctic region of North America. steppes), and the xerophilous scrub vegetation (the
3. The Palearctic region of continental Eurasia, to which Mediterranean maquis on the coasts of the Mediterranean in
must be added the Sino-Himalayan subregion (China), Europe and North Africa), the chaparral in North America
also known as the Chinese transition zone, and the and, finally, further South, a dry predesert and desert belt.
Saharo-Arabian subregion, also known as the Saharo- Eurasia and North America, especially in the northern
Sindic transition zone, North Africa, the northern areas, still share many species, genera, or entire families
Sahara, and the Middle East. Here live some species or that in the Pleistocene were able to cross the Bering bridge
subspecies of very interesting felids, in populations that between the two continents. For example, among the
are isolated and separated from their sub-Saharan Africa Bovidae they have in common: the bison (American and
congeners. The famous lions of the Atlas or Berbers European), the musk ox, the wild sheep (Ovis); among the
were hunted almost to extinction at the start of the 20th Cervidae the deer wapiti, the moose, the reindeer, and the
century. They survive in some small areas of the caribou; and among the carnivores the brown and polar
Moroccan mountains, along with the leopard, and in bear, the gray wolf, the red fox, many mustelids (wolver-
Algeria and Tunisia, the caracal and the serval. In some ine, martens, stoat, weasels), and the Lynx genus. Even
particularly favorable small areas, there is a population among the rodents there are many shared species:
of very rare Sahara cheetahs (Acinonyx jubatus heckii?) marmots, squirrels, flying squirrels, ground squirrels
which some genetically identify more closely with Asian (Citellus, etc.), and chipmunks (Tamias, etc.)

The current distribution of the terrestrial ecoregions (ecosystems) in the various continents is influenced by the characteristics of the climate
(especially by temperature, rainfall, and seasonality) that vary with latitude and altitude. With the climate the characteristics of the vegetation change,
which in turn affects the type of fauna. This map shows the layout of the various habitats and highlights that in the northern hemisphere, from the
north the same succession of ecosystems is present in both North America and Eurasia.
In the intertropical zone (between the two tropics) we notice the presence of savannas and large tropical and equatorial forests. https://upload.wikime-
dia.org/wikipedia/commons/e/e4/Vegetation.png.
The distribution of terrestrial ecosystems has changed many times in the past, both under the influence of land provision, which changed based on
continental drift and ocean level, and on slow but continuous climate changes. One of the causes that influenced the current distribution of animals
was the succession of four glaciations (cold periods), alternating with four interglacial periods (warmer periods), which occurred in the Pleistocene
period. The map shows the distribution of the terrestrial ecoregions, during the last great glaciation of the Pleistocene or Last Glacial Maximum
(LGM, called Würm in Eurasia and Winsconsin in North America) that occurred about 20,000 year BP. Under the influence of the lowest tempera-
tures, polar ice had invaded the most northern parts of the continents and the transformation of large masses of water into ice had decreased sea levels
by as much as 120 m. This meant that Eurasia and North America were united by the Bering bridge, the platform of the Sunda had completely
emerged, India and Ceylon were united, and Japan was in contact with Siberia.
The cooling of the climate had upset the arrangement and extension of many terrestrial ecoregions, increasing the spread of the cold-climate ecosys-
tems (tundra, taiga, etc.) that pushed more and more southward, and decreased the temperate-warm climate woods and the equatorial forests, which
were replaced by grasslands and tropical savannahs.

African leopard Photo by V. Martegani

Snow leopard

(Continued )
 (Continued)
Among the felids there are species that have anatomical, In contrast, other species (stenobionte species from the Greek
physiological, and behavioral characteristics which enable them steno 5 narrow, limited, and bionte 5 that can live) have less
to live in different natural environments. These are euribionte adaptability, are very specialized, and live only in certain
species (from the Greek euri 5 large, great, and bionte 5 that can habitats, as they need particular climatic-ecological conditions.
live) that have a great potential for adaptation to both different Among the felid stenobionte species is the snow leopard which,
temperatures and types of prey. Euribionte species among the with its thick fur, lives only in high mountainous environments or
felines include the leopard, the tiger, and the Asian leopard cat. in cold climates; this big cat has specialized in hunting mountain
These species do not have extreme specializations and live in animals such as goats and marmots. Even the Iberian lynx, due to
warm humid tropical habitats and in the cool temperate forests of its specialized diet based on European wild rabbits (Oryctolagus
Manchuria, feeding on a wide range of prey. The leopard cuniculus), lives only in the Iberian peninsula (Spain and Portugal)
especially has a vast distribution area that includes the in Mediterranean scrublands.
Afrotropical, Palearctic, and Indo-Malaysian regions. The puma
also has this ability and lives in many habitats of the Nearctic and
Neotropical regions; among the felids, the puma is the species
with the widest north south distribution area: from the Yukon
(north Canada) to southern Chilean Patagonia.
Zoogeographic distribution of the species and the main subspecies of Felidae.

Regions Endemic species Species/subspecies

Six endemic species
European wildcat (Felis silvestris) Amur leopard (Panthera pardus
Chinese desert cat (Felis bieti) orientalis)
Iberian Lynx (Lynx pardinus) Persian leopard (P. pardus tulliana
Eurasian lynx (Lynx lynx) ex saxicolor)
Snow leopard (Panthera uncia) Arabian leopard (P. pardus nimr)
Manul or Pallas cat (Otocolobus North African Lion (Panthera leo leo)
manul) Siberian and Caspian tiger (Panthera
tigris tigris)
Asian African wildcat (Felis libyca
libyca and Felis libyca ornata)
Sand cat (Felis margarita)
Caracal (Caracal caracal)
16 total species
Asian cheetah (Acinonyx jubatus
venaticus)
Saharan cheetah (A. jubatus hecki)
Jungle cat (Felis chaus)
Asian leopard cat (Prionailurus
bengalensis euptilurus)
Temminck’s or Asiatic golden cat
(Catopuma temminckii moormensis)
One endemic species
Canadian lynx (Lynx canadensis) Bobcat (Lynx rufus)
Ocelot (Leopardus pardalis)
Northern puma (Puma concolor
cougar)
Jaguarundi (Puma (Herpailurus)
yagouaroundi)
Jaguar (Panthera onca)

6 total species
Three endemic species
Black-footed cat (Felis nigripes) Leopard (P. pardus parduss)
African golden cat (Caracal aurata) Cheetah (A. jubatus)
Serval (Leptailurus serval) West and Central African Lion (P. leo
leo)
East and South African Lion (P. leo
melanochaitus)
Jungle cat (F. chaus)
Caracal (C. caracal)
African wildcat (F. libyca cafra)
Sand cat (F. margarita)

10 total species
Eight endemic species
Indochinese clouded leopard (Neofelis Asiatic lion (P. leo leo ex persica)
nebulosa) Leopard (P. pardus)
Sunda clouded leopard (Neofelis Tiger (P. tigris)
diardii) Leopard cat (P. bengalensis)
Sunda leopard cat Temminck’s or Asiatic golden cat
(Prionailurus javanensis) (C. temminckii)
Rusty spotted cat (Prionailurus Jungle cat (F. chaus)
rubiginosus)
Marbled cat (Pardofelis marmorata)
Bornean bay cat (Catopuma badia)
14 total species Flat-headed cat (Prionailurus
planiceps)
Fishing cat (Prionailurus viverrinus)

(Continued )
 (Continued)

Zoogeographic distribution of the species and the main subspecies of Felidae.

Regions Endemic species Species/subspecies

Seven endemic species
Pampas cat (Leopardus colocola) Southern puma (Puma concolor)
Geoffroy’s cat (Leopardus geoffroyi) Bobcat (L. rufus)
Guigna or Kod kod (Leopardus guigna) Ocelot (L. pardalis)
Andean cat (Leopardus jacobita) Jaguarundi (P. (Herpailurus)
Northern oncilla (Leopardus tigrinus) yagouaroundi)
Southern oncilla (Leopardus guttulus) Jaguar (Panthera onca)
Margay (Leopardus wiedii)

12 total species

The Borneo bay cat (Pardofelis badia) and the Iberian lynx (Lynx pardinus) are two of the rarest feline species with the smallest distribution areas. The
first is an endemism of the Island of Borneo. The second is an endemism of the Iberian peninsula (Spain and Portugal). https://www.lynxexsitu.es/.

The current felids are distributed in all the zoogeo- found as far as the Manchuria area in the Palearctic
graphic regions except the Australian or Australasian region.
region and the Antarctic continent. The Puma lives in North, Central, and South America,
Many species of felids live in more than one region; while the jaguar and ocelot are found in Central and
for example, the lion lives in the Ethiopic region and, in South America and also in some restricted areas of North
historical times, lived in southern Europe, while in north America (Texas).
Africa it was present until about 1920 (or possibly as late Some species, like many felines of the genus
as 1940). Today it also survives in some small reserves in Leopardus (L. colocolo, L. geoffroyi) are endemic to
western India, with the best known being the semiarid for- the Neotropic region (South and Central America),
est of Gir in Gujarat; the caracal lives in Africa, the some are endemic to the Andean region or subregion, such
Middle East, and India; the cheetah lives in Africa and in as the Andean cat (L. jacobita) and guigna (L. guigna).
the Middle East (now only in Iran); and the Libyan wild- In the Indo-Malaysian region there are many ende-
cat, with three subspecies, and the leopard with eight or misms such as the clouded leopard (Neofelis), the marbled
nine subspecies, have wide distributions in Africa, cat (Pardofelis marmorata), and Temminck’s cat
Eurasia, and the Indo-Malaysian region. (Pardofelis temminckii), whereas the Borneo bay cat
The Asian leopard cat (Prionailurus bengalensis) and (Pardofelis badia), the Sunda leopard cat (Prionailurus
the tiger are found in the Indo-Malaysian region, in the javanensis), and the flat-headed cat, are endemic to the
south to the island of Bali, while in the north they are Sunda island and Sundaic subregion.
 Zoogeography of felids Chapter | 3 83

The European wildcat, the Chinese desert cat, the climate changes that have caused repeated transformations
snow leopard, the Eurasian lynx, and the Iberian lynx are of the various ecosystems.
endemic to the Palearctic region. This complex of phenomena has favored speciation
The Canadian lynx lives only in North America, while phenomena and the present and past presence of many
the red lynx or bobcat goes further south and reaches forest environments rich in ecological niches has allowed
Mexico and Central America. many species of felids to evolve and adapt by cohabiting
without significant competition; today we have arboreal
(marbled cat and the two species of clouded leopards),
South-Eastern Asia: a biodiversity hot spot moderately arboreal species and terrestrial (leopard,
for safeguarding some rare species of felids Temmink’s cat), terricolous (tiger and Borneo bay cat),
and aquatic species (flat-headed cat, fishing cat), etc.
The Eastern or Indo-Malaysian zoological region includes Unfortunately, today most of the forests of Southeast
India, the countries of Southeast Asia, and the archipela- Asia and the Sunda Islands have been destroyed to obtain
gos of the Philippines and the Sunda Islands. timber or to make way for huge oil palm plantations that
The various types of forest habitats of Southeast have made the ecological landscape more uniform and
Asia (tropical evergreen forests, plains, mountains, man- monotonous, while anthropization has continued to restrict
groves, deciduous forests, monsoon forests, bamboo, the original natural areas (original wildlife). For this reason
Dipterocarpacee, etc.) are a complex mosaic of ecosys- many species and subspecies or insular populations of Felidi
tems that are ideal for hosting a great biodiversity includ- have become extinct, such as those of the tigers of the
ing a large number (14) of Felid species. islands of Bali and Java; many others have become very rare
The complex geological vicissitudes of the Malacca or are on the verge of extinction, given the small island areas
peninsula and the Sundaic Archipelago between the end to which they had adapted. Today, the Sundaic or Sumatran
of the Pliocene and throughout the Pleistocene, have tiger and Malay and Java leopards and the various species of
favored phenomena of species expansion and subsequent small cats (fishing cat, marbled cat, Borneo bay cat,
isolation, due to the continuous variations in sea level and flat-headed cat, Sunda leopard cat) all risk becoming extinct.

https://www.shutterstock.com/it/image-vector/tiger-681343525?src 5 mWFVeMaycln1rZoNCWvl4g-1-22.
84 Felines of the World

The Felids of the Indochinese Subregion and favored the diversification of various species of small
Sundaic Subregion and medium-sized felids; one of these, the Borneo bay
cat is endemic to Borneo, while the Sunda leopard cat
Southeast Asia is divided into subregions, two of (P. javanensis) is endemic to the Sunda and some small
which are: islands of the Philippines. In addition to the tiger and
G
leopard and the two species of clouded leopards (line-
The Indochinese subregion which includes parts
age Panthera), many small cats live in these regions.
of south China, Vietnam, Laos, Myanmar, and
The small cats of Indochina and the Sunda archipelago
Thailand;
G
belong to the ancient lineage of the Borneo bay cat and
The Sundaic subregion or Sundaland, with Malaysia
to the more recent lineage of the Asian leopard cat.
and the large and small Sunda islands (Sumatra,
Borneo, Java, Bali, etc.). It is probable that long periods In Southeast Asia, more species and subspecies of
of geographic isolation on the various islands have Felids live than in any other part of the world.

Southeast Asia is divided into four subregions; the Indochina or Indochinese subregion and
The Indo-Malaysian the Sundaic (or Sundaland) subregion have as a point of passage from one to the other, both
today and in the past, the Isthmus of Kra. Instead, in the south, between the islands of Bali
zoogeographic region and Lombok it crosses the border with the subregion Wallacea (Sulawesi Timor and other
islands of New Guinea). In the northeast there is the subregion of the Philippines, often united
in the Sundaica

https://en.wikipedia.org/wiki/Indomalayan_realm#/media/File:Ecozone_Indomalaya.svg, Map by David S. Woodruff, D.S., 2010. Biogeography and

conservation in Southeast Asia: how 2.7 million years of repeated environmental fluctuations affect today’s patterns and the future of the remaining refugial-
phase biodiversity. Biodivers. Conserv. 19, 919 941.
 Zoogeography of felids Chapter | 3 85

Starting from the late Pliocene and during the Pleistocene sea levels have risen and lowered several times under the influence of episodes of sudden
variations in temperature that have occurred, especially in the late Pliocene. 10 rapid level increases of more than 80 m and 48 increases of 40 80 m
have been documented in the last 5 million years (Woodruff and Turner, 2009). This has caused continual expansions and decreases of land in the
Sundaic region, with the Isthmus of Kra being a geographical barrier to the expansion of the Sundaic species in Indochina. The lowering of the sea
level allowed the emergence of land bridges between the continent and the Sunda Islands. (A) In the Pleistocene period of the Great Glaciations, dur-
ing the LGM which occurred about 20,000 year BP, the sea level lowered to a level never seen before, 120 m, so that the Sunda platform emerged
and the archipelago became a single block of land, a continuation of the Indochinese peninsula. Because of these phenomena, many species, including
the Felids, were able to colonize lands that they could not reach previously. (B) (C) When the sea levels increased, the various species remained iso-
lated on the various islands of the archipelago and had no further contact with the Indochinese species. Map by Woodruff, D.S., 2010 Biogeography
and conservation in Southeast Asia: how 2.7 million years of repeated environmental fluctuations affect today’s patterns and the future of the remain-
ing refugial-phase biodiversity. Biodivers. Conserv. 19, 919 941.

New species in the Sunda Archipelago researchers (Shu-Jin Luo et al.) published a scientific article
in 2014 where they analyzed many biological samples.
Three species of small felidae are endemic to the Sundaic They proved that, what were considered the subspecies of
subregion: the Borneo bay cat (Bay cat lineage), the flat- the leopard cat of the Sundaic islands and the Philippines
headed cat, and the Sunda leopard cat (leopard cat lineage), (P. bengalensis borneoensis, P. bengalensis javenensis,
in addition to the Sunda clouded leopard (lineage P. bengalensis sumatranus, P. bengalensis heaneyi, P. ben-
Panthera). Separate species live in the Indochinese subre- galensis rabori) and of the marbled cat (P. marmorata mar-
gion and in the islands of the archipelagos: the fishing cat, morata), have been isolated for an extremely long time (up
Asiatic leopard cat, marbled cat, Temminck’s golden cat, to 2 million years) and have no had no further genetic
and the tiger and leopard. exchanges and so are phylogenetically distinct from those
These species took advantage of the repeated lowering living in the rest of Indochina and the Indo-Malayan region.
and rising of the sea level, which united and divided the These researchers thus proposed to group the insular subspe-
islands of the Sundaic subregion with the Malacca penin- cies of the leopard cat by separating them into a new species
sula. By crossing bridges of emerged lands (like the Isthmus called the Sunda leopard cat (P. javanensis) splitting the
of Kra), they became distributed throughout almost the leopard cat of India from that of China and Indochina
whole territory. With the definitive increase in the level of (P. bengalensis). The family Felidae has therefore been
the seas, islands were formed where the cats remained “iso- enriched by a new species, while studies on the marbled cat
lated” and over time diversified into a number of subspecies will soon give results for or against separation into a further
distinct from those of the Indochinese peninsula. By study- new species.
ing the insular subspecies from a genetic point of view,
86 Felines of the World

Map courtesy by Peter Gerngross (2012) www.biogeomaps.eu. Modified by Bellani.

Observing this map with the distribution of felids in the corresponding species in Indochina: the Sunda
the Sundaic or Sundaland subregion, we can see some clouded leopard (Neofelis diardi) and the Sunda leop-
peculiarities in the distribution of some species: ard cat (Prionailurus javanensis). It is possible that a
G
different species of marbled cat (from Sumatra and
The fishing cat today does not reach Bali, where it
Borneo) will soon be found;
seems to have never lived; G
G
Temmink’s cat from the Malay Peninsula reached
The leopard is not present in Sumatra and currently
Sumatra but not Borneo;
we have no evidence that it lived there in the past; G
G
The flat-headed cat has only populated Sumatra,
It is established that although there are no longer tigers
Borneo, and the Malacca peninsula but has gone no
in Borneo, a large island that could sustain a good pop-
further north of the Strait of Kra;
ulation of this big cat, they were present on the island G The position of the Toba volcano in Sumatra (today a
until the beginning of the Holocene, as reported by
volcanic lake) is also indicated, the eruption of which
Piper et al. (2007); moreover, tiger fossils dating back
caused climate changes that greatly influenced the dis-
to less than 20,000 years ago have also been found in
tribution of the species of many mammals in
Sri Lanka and Palawan where it no longer lives, proba-
Southeast Asia and particularly in the Sundaic region.
bly due to the eruption of the Toba volcano;
G Tigers were present until the end of the 20th century
in Bali and Java, the islands of the sub-Sundaic region The great eruption of the Toba volcano in
more distant from Indochina and where they were
extinguished by excessive anthropic pressure (hunting
Sumatra
and destruction of forests); The current distribution of felids (and probably other spe-
G The clouded leopard (Neofelis nebulosa) and the cies of mammals) in the Sunda archipelago has also been
Asian leopard cat (Prionailurus bengalensis) in the greatly influenced by the large volcanic eruption of
Sunda Islands have diversified into species other than Mount Toba, north of Sumatra Island. This is a
 Zoogeography of felids Chapter | 3 87

phenomenon that has only been studied in depth recently, Consequently, with passing time and growth of the popula-
and has given truly surprising results from the point of tion, the territory occupied by a group of felines becomes
view of climate change at the time. About 73,000 years ever wider. Felines are also very skilled predators,
ago the volcano exploded emitting 2800 km3 of ash and and they are able to quickly explore and conquer new
lava, perhaps the largest known volcanic explosion; it is areas as soon as the opportunity arises, and so it is not sur-
very probable that the fumes emitted obscured the sky for prising that they have successfully colonized unexplored
a vast area, causing temperatures to fall by almost regions.
3 C 5 C for several years after the event. It seems that These migratory movements and territorial dispersions
in the first year the temperatures may have decreased by were favored by the emergence of land bridges
as much as 10 C, having a tremendous influence on the created by lowering sea levels, and which allowed the col-
ecology of the entire planet; this icy wave seems to have onization of continents that had never previously been
also caused the extinction of many species of mammals reachable.
and much of the human population at the time. This also favored many species of felids that, moving
This phenomenon could be due to the singular absence easily from one region to another and moving away from
of the leopard from Sumatra Island, as it is certain that their original populations, often evolved into new species.
about 73,000 years ago a large part of the population of
tigers from Southeast Asia was almost wiped out and that
the current population shows a low genetic variability of 3.2.1 The most Ancient migratory wave
the species which indicates that it has undergone a genetic
bottleneck, and which has repopulated today’s areas start- From the Early-Middle Miocene at the
ing from a small contingent of survivors after some envi- beginning of the Pliocene, from about 15 12
ronmental catastrophe. to 5 4 Mya
We do not know exactly how many millions of years ago
the first specimen of carnivore appeared that would later
3.2 The origins of the current distribution of become the progenitor of all modern felids. Various
hypotheses have been formulated ranging from about 9 to
felids around the world about 15 Mya, in the period between the Early and Middle
The current distribution of all the species of Felidae in the Miocene. That the place of origin of the current felids was
various continents of the world and in the related zoogeo- Central Asia has been hypothesized and also calculated
graphic regions is the result of migratory waves (at least through genetic analysis based on molecular biochemistry,
two) that took place over a few million years, and which but this has never really been proven. The discovery of the
were caused by climatic-ecological changes that alternated fossil Panthera blyteae, on the Tibetan plateau, dating
in the last geological periods, from the Miocene to today. back to 4 6 Mya, gives us almost irrefutable proof. It is
Some of these changes, such as the great glacial and inter- therefore Central Asia that has been identified as the place
glacial periods of the Pleistocene, up to the LGM, have where the first ancestors of the Felids in the Miocene
influenced sea levels and alternately created and destroyed began and evolved and then spread, first across the conti-
land bridges between continents, such as the Bering bridge nent and then, thanks to their success as predators in the
which united Asia and North America, the isthmus of various natural environments, they began to diversify into
Panama between the Americas, and the lowering of some various species and, under the influence of climate-
inland seas such as the Mediterranean and the Red Sea. ecological changes, to migrate from their place of origin,
These appearing and disappearing land bridges also which may have become too cold, to Africa (as precursors
occurred between continents and islands, such as the isth- of the caracal lineage) and North America (as precursors of
mus of Kra between Indochina and the Sunda Islands, the lynx, puma/jaguarundi, and ocelot lineages) and, only
which for a limited period of time were all united by the much later, also to South America (ocelot lineage).
emergence of the Sondaic Platform of which they repre- During this period, migrations were facilitated by par-
sent the highest lands. During the LGM sea levels fell by ticularly low sea levels, as the glaciation of many water
approximately 120 meters, and the entire Sunda Shelf was resources caused levels to decrease by about 60 m. For
exposed. Some groups of felids adapted to climate change, this reason they formed bridges of emerged lands such as
others became extinct, but many started migrations from the isthmus of Bering, a Panama bridge even larger than
one continent to another, perhaps following their prey that the current one in Central America, a well-traveled route
often had migratory habits; behavioral instinct would also between the Middle East and Africa (through the Red
encourage many species to expand their distribution area Sea), and between the Indochina peninsula and Sunda
as new generations were able to move away from the Islands. By means of these bridges the felids began to
places they were born to conquer new territories. conquer new territories.
88 Felines of the World

G From about 15 to 12 Mya, Asia was also inhabited by the precursors of the Pantherinae and about 6 Mya the genus Neofelis began its separation
from the precursors of the genus Panthera, and migrated to Southeast Asia and the Sundaic Islands.
G The ancestors that gave rise to the caracal, the serval, and the African golden cat (caracal lineage), reached Africa about 12 8 Mya.
G Many felines crossed the Bering bridge and arrived in North America in different eras: 10 Mya the ancestors of the ocelot lineage began. This
was followed about 6 Mya by lynxes and, almost 5 Mya, by the precursors that gave rise to the genera of Puma and Jaguarundi (while their rela-
tives, the cheetahs, remained in Eurasia and later headed to Africa and the Middle East as the current species).
G In the same period, about 6 Mya, an ancestor of the golden cat of Borneo was separated by dividing from the lynx and began a migration to the
south that led it to Indochina and the Sunda Islands; and here, at the beginning of the Pliocene (5 Mya) it gave rise to the present species, first the
marbled cat and then the two species of golden cat.
G Afterwards (about 3.8 Mya), probably due to climate cooling, the ocelot lineage moved toward the south looking for more favorable climatic-
ecological conditions and reached South America. Here, the most favorable climate, a tropical environment and with very diverse geographical
conditions (from wetlands and forests to semiarid plains up to the Andes) favored the diversification of the current species of the genus Leopardus
(ocelot, kod kod, margay, Geoffroy’s cat, Pampas, and Andean cats, etc.).
After this wave of migration the climate changed and sea levels increased, again separating the continents, so that many groups of felids remained iso-
lated and continued their evolution separately from their predecessors.

(A) Ocelot (Leopardus pardalis). The ancestors of the ocelot lineage crossed the Bering bridge to North America about 10 Mya; 3.8 Mya they arrived
in South America and differentiated into various species, including the ocelot.
(B) Serval (Leptailurus serval). The ancestors of the lineage caracal, which also includes the serval, reached Africa very early between 12 and 8 Mya.
Only the caracal then reached the Middle East and western India.
 Zoogeography of felids Chapter | 3 89

3.2.2 The subsequent migratory waves and

extinction of the megafauna

A beautiful image of the Spanish paleontologist-artist Mauricio Anton. Megafauna is represented in the north of Spain during the Pleistocene: woolly
mammoths (Mammuthus primigenius), woolly rhinoceros (Coelodonta antiquitatis), a herd of Equidae and European cave lions (Panthera leo spelaea)
with a reindeer carcass. https://upload.wikimedia.org/wikipedia/commons/e/e6/Ice_age_fauna_of_northern_Spain_-_Mauricio_Ant%C3%B3n.jpg

From the Pliocene to the Pleistocene: 5 4 Mya or less warm periods, drastically changed the ecological
to up to 8000 10,000 years BP landscapes of the continents and their geography, continu-
In a period ranging from four million up to a few thou- ally connecting them with each other and then dividing
sand years BP, many other migrations occurred, always in them again, due to changing sea levels. Under the influ-
correspondence with the lowering of sea levels and the ence of these recurring climatic changes and with the pos-
consequent emergence of new lands; these served sibility of migrating from one continent to another, the
as passages or areas to expand the territories to be felids diversified into new species that were increasingly
colonized. The last migration of the felids was that of the adapted to the new conditions. In fact, between the
puma which occurred between 8000 and 10,000 years Pliocene and the Pleistocene the diversification and birth
ago. This species returned from South America to repopu- of the largest number of new species occurred. These are
late North America where it had become extinct during the same species that still populate almost all the conti-
the LGM which had caused a great extinction of the nents of the Earth today.
megafauna in the Northern Hemisphere. Naturally, as fossil finds show us, many other species
were unable to adapt and survive; for example, during the
Pleistocene the so-called “great extinction of the mega-
Species and climatic-ecological changes fauna” took place which destroyed more than half (per-
Between the Pliocene and the Pleistocene there was a haps as much as 70%) of the large species of herbivores
continuous succession of different climatic phases, up to and carnivores that had represented the typical fauna of
the four great glaciations of the Pleistocene (Gunz, the Quaternary. Especially in North America the phenom-
Mindel, Riss, and Wurm) that, with the continuous alter- enon was very drastic because with the melting of the last
nations between cold and interglacial periods with more glaciation ice (Wurm, about 15,000 12,000 years ago)
90 Felines of the World

the vegetation did not recover as in the past, and the her- disappeared with them. These included many species of
bivores (horses, camels, mastodons, woolly rhinos, giant saber-toothed cats (Machairodontinae), American pumas,
ground sloths, etc.) became extinct, and their predators and the puma-cheetah-like Miracinonyx.

G About 7 Mya two related lineages, the ancestors of the domestic cat lineage (Felis), and the Asian leopard cat lineage, separated and began to col-
onize Asia and Africa (the former) and Southeast Asia (the latter).
The lineage of the domestic cat (genus Felis) expanded and first reached Africa as the black-footed cat (Felis nigripes), an ancient species that
dates back to almost 3 Mya and reached South Africa where it is still widespread today. The sand cat and Felis libyca followed.
Meanwhile the Manul or Pallas cat, an ancient species attributed (with some doubt) to the lineage of the Asian leopard cat, dates back to 6 Mya
and has not undergone large variations to the present.
G Less than 2 Mya some lynxes returned to Eurasia, while others stayed in North America, where they remain today.
G The cheetahs populated Eurasia (Acinonyx pardinensis, etc.), while the current species expanded to reach Africa and the Middle East.
G About 300,000 400,000 years ago, the cave lion (Panthera spelaea) and perhaps other species populated Eurasia, while others reached the
Americas and became Panthera atrox. These cold-climate lions died out around 10,000 11,000 years BP.
The current lions survived from Asia and migrated to Africa and the Middle East at the same time as the leopard.
G The jaguar, with a Eurasian origin (such as Panthera gombazgoegensis 5 toscana), migrated to North America 3 Mya and then reached South America.
G The puma and the jaguar colonized South America.
Only recently did the Puma migrate to South America from North America and then return to recolonize almost all of North America.
G About 2.5 Mya the tiger began its evolution and colonized southern Asia, pushing toward the Sunda islands. About 73,000 years BP the terrific eruption
of the Toba volcano in Sumatra destroyed almost the entire population that was also once widespread in Borneo, Sri Lanka, and Palawan.