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REVIEW
published: 01 December 2020
doi: 10.3389/fvets.2020.570413
Frontiers in Veterinary Science | www.frontiersin.org 1December 2020 | Volume 7 | Article 570413
Edited by:
Alejandra Victoria Capozzo,
Consejo Nacional de Investigaciones
Científicas y Técnicas
(CONICET), Argentina
Reviewed by:
Saket Kumar Niranjan,
Indian Council of Agricultural Research
(ICAR), India
Rostam Abdollahi-Arpanahi,
University of Florida, United States
*Correspondence:
Antonio Humberto Hamad Minervino
ah.minervino@gmail.com
Specialty section:
This article was submitted to
Veterinary Epidemiology and
Economics,
a section of the journal
Frontiers in Veterinary Science
Received: 07 June 2020
Accepted: 27 October 2020
Published: 01 December 2020
Citation:
Minervino AHH, Zava M, Vecchio D
and Borghese A (2020) Bubalus
bubalis: A Short Story.
Front. Vet. Sci. 7:570413.
doi: 10.3389/fvets.2020.570413
Bubalus bubalis: A Short Story
Antonio Humberto Hamad Minervino 1
*, Marco Zava 2, Domenico Vecchio 3and
Antonio Borghese 4
1Laboratory of Animal Health, LARSANA, Federal University of Western Pará, UFOPA, Santarém, Brazil, 2Argentine Buffalo
Breeders Association, Buenos Aires, Argentina, 3Italian National Reference Centre on Water Buffalo Farming and
Productions Hygiene and Technologies (CReNBuf), Istituto Zooprofilattico Sperimentale del Mezzogiorno, Portici, Italy,
4International Buffalo Federation – IBF, FAO/ESCORENA Buffalo Network, Animal Production Research Institute, Rome, Italy
The domestic buffalo (Bubalus bubalis), also known as water buffalo or Asian buffalo
to prevent confusion with the American bison (Bison bison), wrongly named buffalo in
North America, comprises two subspecies: the river buffalo (B. bubalis bubalis) and the
swamp buffalo (B. bubalis kerebau). The swamp buffalo has a consistent phenotype
and is considered as one type, even if many breeds are recognized within it; conversely,
the river buffalo subspecies has many breeds. We found limited information available
regarding the worldwide distribution of buffaloes. The best estimate is that 208,098,759
buffalo head are distributed in 77 countries in five continents. In this review, we presented
the basic aspects of the water buffalo and unraveled the buffalo path followed from the
origin of the species to its current global distribution. We reviewed several data sources
to provide a better estimate of the world buffalo count and distribution.
Keywords: water buffalo, bubaline, breeds, dairy buffalo, buffalo population
THE Bubalus GENUS
Bovids are the largest family within the Artiodactyla order. The name bovines or Bovinae Gray,
1821 refers to a subfamily of the Bovidae family, which includes nine genera: Bos (cattle), Poephagus
(yak), Bison (bison), Syncerus (African buffalo), Boselaphus (nilgai), Pseudoryx (saola), Tetracerus
(four-horned antelope), Tragelaphus (kudu and relatives), and Bubalus (domestic buffalo) (1–3).
Figure 1 depicts the zoological classification of the Bovinae subfamily.
Bubalus, a genus of the Bovidae family, was distributed widely in Europe and Asia during
the Pleistocene. Wild animals of this genus include the wild Asian buffalo (Bubalus arnee) that
originates in northern India and lives in marshes and in the jungle. This is a very large animal,
reaching a height of up to 200 cm and a weight of up to 1,000 kg; it can be either gray-black,
dark gray, or dark brown and has large horns that are separated at the base (4). This species
was distributed across Asia but is now considered endangered (5). The anoa, which is endemic
to Indonesia, comprises two species: the lowland anoa (B. depressicornis) and the mountain anoa
(B. quarlesi); however, there is some debate on whether they are the same or distinct species (6).
The anoa inhabits only Indonesia, had 46 chromosomes, and is a small animal (100 cm tall) with
thin and straight horns (25 cm long). It is conserved in zoos and never used for draft work or
for food production and lives in mountains or in lowland forests (7). The anoa was divided in B.
depressicornis and B. quarlesi based on the environment it inhabited. Bubalus mindorensis is found
only on the island of Mindoro in the Philippines; therefore, it is also called tamaraw or Mindoro
buffalo. It is a small animal (100 cm tall) with short and strong horns. It is a critically endangered
species from the Philippines with only 220–300 mature animals remaining (8). Figure 2 shows the
phylogenetic tree based on the D-loop region of several species of the Bovini tribe, including the
Bubalus genus.
Minervino et al. Bubalus bubalis: A Short Story
FIGURE 2 | Phylogenetic tree of the genus Bubalus based on D-loop region complementing with IOC and cytochrome B.
THE WATER BUFFALO
The water buffalo or Asian buffalo (Bubalus bubalis) is a
domesticated species that can be confused by the general
population with the wild American bison (Bison bison),
incorrectly called Buffalo in North America, and occasionally,
with the African Buffalo (Syncerus caffer), which are two
unrelated species of wild animals. The water buffalo comprises
two subspecies: the river buffalo (B. bubalis bubalis) and the
swamp buffalo (B. bubalis kerebau), which are genetically
distinct with different chromosome numbers (50 and 48,
respectively), and distinct morphology, physiology, and
productive and reproductive performances (9). The two
subspecies are interfertile, with their progeny containing
49 chromosomes. Male crossbred progeny may sometimes
present fertility problems, while female progeny exhibits longer
calving intervals (10); however, this only applies in the case of
further backcrossing.
The morphology of the two subspecies differs considerably.
River buffaloes are usually larger than swamp buffaloes and
weigh between 450 and 1,000 kg. Most breeds have curled
horns and are mainly found in India, Pakistan, and some
European, western Asian, and American countries. The river
buffalo is primarily reared for dairy production, especially in
Asia and Europe, but is also used for meat production or as
a dual-purpose animal as well as for draft purposes (4,11).
Swamp buffaloes are smaller and lighter than river buffaloes and
normally weigh between 325 and 450 kg. Swamp buffaloes are
reared mainly for draft purposes; however, they can also yield
high amounts of milk (up to 600 kg of milk per year) (11). Swamp
buffaloes are predominant in Southeast Asia and Australia (12).
A few animals can also be found in the north-eastern states of
India (13).
Buffaloes can be affected by the same diseases and parasites
as cattle; the extent to which they are affected varies
drastically depending on the country, region, and production
system. Buffalo breeds have varying degrees of tick resistance;
however, buffaloes are infested by the louse Haematopinus
tuberculatus that is specific to them (14). Brucellosis, tuberculosis,
leptospirosis, bovine viral diarrhea, fasciolosis, foot-and-mouth
disease, and protozoal infections have important economic
impacts on the water buffalo industry. Regarding public
health, water buffaloes are involved in the transmission
of schistosomiasis to humans (15,16). A recent report
showed that buffaloes may play some part in Q fever
epidemiology, as Coxiella burnetii was detected in buffalo
milk (17).
The buffalo has a higher capacity to convert feed with
poor nutritional value, and a high capacity for adaptation and
survival in different environments with distinct topography,
climate, and vegetation (18). The water buffalo is used as a
draft animal in Asia, owing to its strength; however, it is
also used as a riding animal on Marajó Island, Brazil. Water
buffaloes are highly adapted to hot and wet floodplains and
play an important role in the economy of many tropical and
subtropical countries, as they are used for meat and milk
production and as draft animals (19). Water buffaloes have
relatively lower heat tolerance than other livestock species
owing to their dark coat color and their inadequately dispersed
sweat glands, which result in a less efficient evaporative
cooling system (20). Recent molecular studies have shown
that buffalo breeds historically raised in a hot climate may
have developed higher heat tolerance (21). Buffaloes have
higher longevity than cattle (they can surpass 30 years of
age, while maintaining their reproductive capacity until they
are 18–25 years old). This docile, intelligent, and curious
animal (Supplementary Video 2) is farmed in several countries
worldwide, mainly for its good quality meat (which has better
nutritional composition compared to that of cattle) and its high-
fat milk (11,19,22,23).
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Minervino et al. Bubalus bubalis: A Short Story
WATER BUFFALO DOMESTICATION
The two distinct water buffalo subspecies (river and swamp)
descended from different populations of the wild Asian water
buffalo that diverged some 900,000 years before present (BP) and
evolved in separate geographic regions. The wild Asian buffalo
is considered the most probable ancestor of the water buffalo,
and the most accepted hypothesis supports two independent
domestication events from river and swamp buffalos. Recent
molecular studies support that domestication started from a wild
ancestor, Swamp-like, that was distributed across mainland Asia
and has long since diverged into the existing population of B.
arnee (9,24). Compared with Bos taurus (10,000 BP), the buffalo
domestication occurred recently, ∼3,000–7,000 years BP.
The domestication of river buffaloes probably occurred at
∼6,300 yr BP in north-western India, from where domesticated
river buffaloes migrated west, across south-western Asia, Egypt,
and Anatolia, and reached the Balkans and the Italian peninsula
(25). Phylogenetic analyses and archaeological evidence suggest
that the river buffalo was domesticated in an atypical and
complex manner, including multiple maternal lineages, with
the successive introgression of wild animals to domestic stocks
(26). River buffalo populations show a weaker phylogeographic
structure but higher phenotypic diversity, thereby resulting in
more breeds (27). After domestication, the water buffalo was
introduced in Egypt and Italy by the Arab invasion around the
eighth century. Buffaloes were also introduced in the Balkans and
Turkey by crusaders and by the Selgiukid invasion during the
expansion of the Ottoman Empire in the fifteenth century (7).
The swamp buffalo was most likely domesticated near the
border between China and Indochina at ∼3,000–7,000 years
BP, spreading across Southeast Asia. Swamp buffaloes have
strong phenotypic uniformity, but molecular studies show a
higher diversity of maternal and paternal lineages, strong genetic
differentiation affected by geography, and a lack of gene flow
(27,28). One large study with 46 buffalo populations in seven
Asian countries revealed an abundant genetic diversity and clear
phylogeographic structure in swamp buffalo populations (29).
One exception is considered a swamp buffalo lineage from China
that has a complex pattern of diversity, thereby suggesting the
presence of a long-term, strong gene flow, probably due to
extensive migrations of buffaloes alongside human movements
(30). A recent study with buffalo whole genome showed similar
genetic patterns in both cattle and buffalo breeds, an example of
convergent domestication down to the same mutation but which
occurred independently (31). Phylogenetic analysis revealed that
Chinese swamp buffaloes could be divided into two distinct
lineages, A and B. Of the two lineages, lineage A was predominant
across all populations. For predominant lineage A, Southwestern
buffalo populations possess the highest genetic diversity among
the three hypothesized domestication centers (Southeastern,
Central, and Southwestern China), suggesting that Southwestern
China is the most likely location for the domestication of lineage
A. However, a complex pattern of diversity is detected for lineage
B, preventing the unambiguous pinpointing of the exact location
of the domestication center and suggesting the presence of a
long-term, strong gene flow among swamp buffalo populations
caused by extensive migrations of buffaloes and frequent human
movements along the Yangtze River throughout history (31).
WATER BUFFALO BREEDS
As mentioned above, the water buffalo comprises two subspecies,
the swamp and the river buffalo. The swamp buffalo has
a consistent phenotype and is considered as one type;
however, separation in breeds still occurs, especially depending
on the geographic location. Conversely, the river buffalo
comprises several breeds. According to a Food and Agriculture
Organization (FAO) report based on data provided by countries,
there are 123 buffalo breeds, 90 in Asia alone, many of which
are local breeds, with only 15 breeds reported as transboundary
(32). There is a lot of variation in horn conformation. In many
cases, it is one of the indicators of the degree of purity in different
breeds. Figure 3 shows photos of swamp and river buffaloes,
while Table 1 presents a list of the most important breeds and
recognized populations of water buffalo.
The swamp buffalo, despite having a consistent phenotype,
had somatic and physiological differences that can be noted
depending on the climate and the land, owing to the total
isolation of herds. The swamp buffaloes of Sulawesi that live in
the jungle are completely different from the Sumatran swamp
breed that lives in lagoons. The Sulawesi breed is clearly a
meat animal, strong and muscular with a big head, and very
appreciated in ceremonies, particularly if it is a spotted animal
(Figure 3A), a characteristic that adds value to the animal.
Sumatran buffaloes are lighter animals, with thin heads, and are
mostly used for milk purposes (7).
The Carabao breed that is native to the Philippines is a swamp-
type buffalo (Figure 3B) used as a draft animal in small-hold,
land-based agriculture. Carabao buffaloes have low, wide, and
heavy built bodies. Their body coloration varies from light gray
to slate gray. Their horns are sickle-shaped or curved backward
toward the neck (11). On the island of Marajó, Pará, Brazil, these
animals are used for meat production. They are brown-gray, have
white spots on their legs, are developed for work, and are suitable
for meat production, while their milk production capacity is
usually poor (33).
The Chinese buffalo is a swamp-type buffalo, used mostly for
draft work, and, depending on the region, comprises a diverse
array of breeds, which amount to 18 local breeds. The most
important of these breeds are the Fuling breed (Figure 3C) living
in the Sichuan mountains, and the Binhu breed, living in Hunan
province; these are the breeds with the highest populations, which
amount to more than 400,000 head each. Other breeds include
the Fuzhong and Xilin breeds from Guanxi province, Enshi
breed from Hubei, Dongliu breed from Anhui province, and the
Shanghai breed. Swamp breeds in China are mainly used for draft
purposes as they are very efficient in the marshlands, particularly
in rice fields, and have limited milk production capacity (10,28).
Swamp subspecies are also dominant in Southeast Asian
countries such as Malaysia, Laos, Cambodia, Vietnam, Myanmar,
and Thailand. In these countries, the buffalo is an important part
of the crop production system, as it is the main draft power
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Minervino et al. Bubalus bubalis: A Short Story
FIGURE 3 | Buffalo breeds worldwide. (A) Sulawesi breed spotted animal (Indonesia); (B) the Carabao breed from Philippines; (C) Fuling breed in China; (D) Murrah
buffaloes herd from São Paulo, Brazil; (E) Nili-Ravi breed at Pakistan; (F) Mediterranean Italy breed raised in Mozambique; (G) Jaffarabadi bull in Brazil; (H) Buffalypso
breed from Trinidad and Tobago. Source: Castro, S.R., Borghese (7).
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Minervino et al. Bubalus bubalis: A Short Story
TABLE 1 | Breeds or recognized population of water buffalo worldwide.
Breed or recognized
population
Subspecies Local Purpose
Anatolian River Turkey Milk and meat
Azeri or Caucasian River Azerbaijan, Georgia Armenia, Iran Milk and meat
Azikheli River Khwazakhela, Swat, Pakistan Milk
Bangladeshi breed River West Bangladesh Milk and draft
Bangladeshi Surti River Central Bangladesh Milk
Bhadawari River Uttar Pradesh, India Milk and draft
Binhu Swamp Hunan, China Draft
Buffalypso River Trinidad and Tobago, Cuba Selected for meat and draft
Carpathian River Transylvania, Romania Milk and meat
Dongliu Swamp Anhui, China Draft
Egyptian River Egypt Milk and meat
Enshi Swamp Hubei, China Draft
Fuling Swamp Sichuan, China Draft
Fuzhong Swamp Guanxi, China Draft
Jafarabadi River Gujarat, India and Brazil Milk and meat
Jerangi River Andhra Pradesh, India Draft
Kalahandi River Orissa, India Draft and milk
Kerbau Papangan Swamp South Sumatra marshlands Milk and meat
Khuzestani River Iraq and Iran Milk and meat
Kundi River Sindh, South Pakistan Milk
Lime River Nepal mountains and river valleys Triple aptitude
Manda River Andhra Pradesh, India Milk and draft
Mazandarani River Iran Milk and meat
Mediterranean River East Europe, Macedonia, Greece, Serbia, Albania Triple aptitude
Mediterranean Italian River Italy and export (live animals and semen) Selected for mozzarella and processing industry
Meshana River Gujarat, India Milk
Mesopotamian River Iraqi marshlands Milk and meat
Murrah River India and export (live animals and semen) Selected for milk
Murrah, Bulgarian River Bulgaria Milk
Nagpuri River Madhya Pradesh, India Milk and meat
Nili-Ravi River Pakistan and Punjab, India Selected for milk
Original Far East Buffalo Swamp Thailand, Vietnam, Laos, Cambodia, Myanmar, Malaysia Triple aptitude, family animal, rice fields
Pandharpuri River Maharashtra, India Milk
Parkote River Nepal mountains and river valleys Triple aptitude
Philippines Carabao Swamp Philippines rice fields Triple aptitude
Primitive Bangladeshi Swamp East Bangladesh Draft and meat
Sambalpuri River Madhya Pradesh, India Milk
Shanghai Swamp Shanghai, China Draft
South Kanara River Mangalore west coast Draft
Sulawesi spotted Swamp TanaToraja Sulawesi, Indonesia Sacrificed during ceremonies
Surti River Gujarat and Rajasthan, India Milk
Tarai River Uttar and Madhya Pradesh, India Milk and meat
Toda River Madras, India Milk and for ceremonies
source for land preparation, supplies fresh organic manure for
cultivation, and is highly adapted to living in different habitats.
Australia imported swamp animals from Asia as the management
systems in the country are extensive (11,34).
The river subspecies breeds have been named according to
Mason (35). There are three international river breeds that
are widely distributed worldwide, owing to their high genetic
potential for milk production: Murrah, Nili-Ravi, and the Italian
Mediterranean (36).
Murrah is the most represented and well-known buffalo breed
in the world. It is selected for milk production in northwest India
and is known for its curled horns (Figure 3D). Its name is a
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Minervino et al. Bubalus bubalis: A Short Story
Hindu word that means spiral and derives from the shape of
its horns. Its skin and hair color is jet black. White spots are
not accepted, except at the end of the tail. These are massive
and stocky animals with heavy bones. Their horns are short and
tightly curled. This breed originated in the center of Haryana
and spread across India. It has been exported to several countries
worldwide, such as Brazil, Bulgaria, and East Asian countries (10,
37). In Bulgaria, the only buffalo breed is the Bulgarian Murrah,
which resulted from crossbreeding the Indian Murrah and the
local Mediterranean breed. Many animals have been exported
from Bulgaria to neighboring countries, such as Romania and
Germany, and to distant countries, such as Brazil and Venezuela.
Murrah buffaloes were exported to China and Indonesia to
increase milk availability, as the local breeds are of the swamp
type with low milk production capacities; the same applies for
many Asian countries, in which the original buffalo population is
of the swamp type (19,37).
The second most represented breed in the world is the Nili-
Ravi (Figure 3E). This is the most important livestock breed in
Pakistan with a head count higher than 10,000,000 in Punjab,
while it is also present in India (38). This breed resembles the
Murrah breed but has white markings on its extremities and
walled eyes; its horns are less curled than those of the Murrah
breed. The skin color of the breed is generally black, but there are
some albino animals, brown, spotted, or with clear eyes (38,39).
Nili-Ravi is the most important dairy buffalo breed in Asia and
is selected for this purpose, with its milk yield reaching 3,050 kg
(40). The Kundi breed is widespread in the Sindh region in south
Pakistan (10). The Azikheli buffalo is an undocumented buffalo
breed from Khwazakhela, Swat, Pakistan. It has a small body size,
brown-colored coat, and horns lying flat, bending laterally, and
directed backward and slightly upward without twisting (41).
The Mediterranean buffalo descended from the river buffalo
introduced to Europe from India during the Arab occupation
of the eighth century in Sicily and in the south of Italy. The
Italian breed is now classified as Italian Mediterranean breed
(Figure 3F) as it was selected for 60 years for milk purposes
and is clearly a dairy breed. This breed was later exported to
several countries in America and Asia, with a large export of
Mediterranean semen from Italy. Buffaloes were also introduced
in east Europe by crusaders in the twelfth century and later with
the Turkish invasion of the Ottoman Empire (fifteenth century).
These breeds are different from the Italian Mediterranean breed.
Balkan breeds such as the Carpathian or the Macedonian are draft
animals, used in carriages, and they are smaller in size and with
low milk yield capacity (36).
European Mediterranean animals have black, brown, and dark
gray coats, while Italian Mediterranean animals are exclusively
black. Their horns are flat at the bottom and face backward, and
their points face upward and inward. The Mediterranean is a
compact breed with a deep and wide chest, but short back and
rump. The udder is medium-sized with squarely placed quarters
and halves and with cylindrical teats (10,36). The breed’s size,
weight, and productivity vary greatly depending on the genetics,
environment, and management system. The average daily milk
yield varies widely and depends on factors such as genetics and
the feeding system. It can range from 3 to 4 kg milk/day for
poorly fed animals up to 15 kg milk/day in intensive production
systems (10). The Italian Mediterranean breed has high milk
yield and quality, as it was selected for the creation of a dairy
breed specialized for the cheese industry, with a genealogical
book established in 1980, and the National Association of Buffalo
Breeders (ANASB) established in 1979. The use of the milk
data recorded for females registered on the genealogical book,
the application of several cycles of progeny testing, and the
spread of artificial insemination with the use of semen of proven
high genetic value created an improved dairy breed, with many
females producing more than 5,000 kg milk/270 days of lactation,
to a maximum of 5,600 kg with 8.3 fat and 4.6 protein, as
the selection goal was not the total milk yield but mozzarella
production (7).
The Jafarabadi breed (Figure 3G) is originally from Gujarat,
India. It is a large animal with a massive and long-barreled
conformation and black coat. Its horns are long, heavy, begin
at the base of the head, face downward, and then curve. The
Jararabadi breed is of particular importance in India, with
animals being exported to Brazil. In Brazil, the Jafarabadi breed
has two varieties: the Gir, which is of medium size and the
most widespread variety, and the Palitana, a large breed used
in crossing.
The milk yield of the Jafarabadi is good; however, this is
also a large animal with high muscularity. Therefore, this breed
was selected by many countries in the Americas as a meat-
purpose breed. It is possible to find pure Jafarabadi animals
or their descendants in Brazil, Colombia, and other South
American countries, where this breed showed great adaptability
to marshlands and other environments, and a high affinity for
meat production. Additionally, the Jafarabadi was the basis for
the creation of the Buffalypso breed in Trinidad and Tobago and
in Cuba (7).
The Buffalypso is the typical buffalo breed of Trinidad and
Tobago (Figure 3H) derived from the Jafarabadi breed imported
in Trinidad in 1905. In 1948, Steve Bennet crossed Bhadwari
bulls with Jafarabadi cows to create the Buffalypso breed (Buffalo
×Calypso, a form of popular Caribbean music), with the
purpose of obtaining a meat breed, which is also useful as
a draft animal. Subsequently, the Buffalypso was crossed with
the Murrah, Surti, Nili, and Nagpuri breeds. It is now also
used for milk production (10,33). The highest Buffalypso head
count is in Cuba and Trinidad and Tobago; however, the breed
can also be found in several countries such as Venezuela,
Costa Rica, Guatemala, the Republic of Honduras, Nicaragua,
Brazil, Panama, Mexico, Colombia, USA, and even Taiwan.
Historically, Buffalypso is the only breed selected exclusively
for meat, as it is a very muscular animal with developed back
and rump, brown-colored, and sold in the American market
as beef.
Other river breeds in India include the Surti breed in
north Gujarat, one of the most important breeds in the
Rajasthan and Gujarat regions, with a population higher than
500,000 buffalo head. These animals have two white chevrons
on their chest, and their horns are flat and sickle-shaped,
directed downward and backward, and then turn upward at
the tip to form a hook (13,42). The Tarai breed amounts
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Minervino et al. Bubalus bubalis: A Short Story
to ∼1,000,000 buffalo head. These animals have long and flat
horns with coils bending backward and upward. This breed is
raised in Uttar Pradesh and Madhya Pradesh (4,10). Other
breeds of ∼300,000–400,000 buffalo head are the Mehsana (its
characteristics are intermediate between the Surti and Murrah
breeds), reared between the Mahi and Sabarmati rivers in
Gujarat, and Nagpuri, a local breed improved by selection
from Indian breeds that have long (50–65 cm) and flat-cur ved
horns (4,10).
Other breeds with smaller populations are the Bhadawari
raised in Uttar Pradesh and Madhya Pradesh, the Jerangi (a
small animal) and Manda (each amounting to 100,000 buffalo
head), which are breeds raised along the border of Orissa with
Andhra Pradesh, Sambalpuri from Bilaspur in Madhya Pradesh,
and Toda, which is a large breed raised in the Nilgiri hills of
Madras (7). Additionally, there are Pandharpuri, Kalahandi, and
South Kanara River buffalo breeds in India.
In Nepal, the Lime breed is believed to have descended from
the ancestral wild buffalo, possibly with river-type introgression,
and became domesticated in the country (43). The Lime buffalo
is found in the mountains, high hills, and hill river valleys in
Nepal with a herd population of ∼700,000 head, which accounts
for ∼35% of the total indigenous buffalo population. This is a
small-sized breed, with small sickle-shaped horns that are curved
toward the neck, and light brown in color with chevrons of
gray/white hair below the jaw and around the brisket (7).
The Parkote buffalo is a typical animal of the mid-hills
and river valleys in Nepal. However, owing to the traditional
crossbreeding with the Lime and, more recently, the Murrah
breeds, their pure breed population estimated at 500,000 head is
now declining. The Parkote have dark-colored coats, black skin,
black muzzles, and no markings on their legs. These animals
are medium-sized, with sword-shaped horns directed laterally or
toward the back (10,35).
The native buffaloes in the western part of Bangladesh are of
the river type, known as the Bangladeshi breed, and account for
∼45% of the total population. The coat color of these animals is
normally black, while they have curly horns. Few individuals have
spots in their tail switch, and some are brown-colored or have
hanging horns. Their average milk yield is ∼620 kg in 270 days of
lactation. Male buffaloes are used for draft and females for dairy
purposes (44).
The Anatolian buffalo that descended from the migratory
Indian buffalo (seventh century) has been raised in Turkey for
centuries. These animals are black-colored and have long hair,
with variations in their tail lengths and frequent white switches.
The Anatolian breed has an estimated population of 122,000
and is raised predominately in the Black Sea region, north of
middle Anatolia, Thrace, Hatay, Mus, Kars, Diyarbakir, Afyon,
and Sivas (45).
The Azeri or Caucasian breed originated in the Indo valley
and descended from the Indian buffalo. Archeological evidence
suggests that buffaloes were raised in Iran (Lorestan region),
around the ninth century B.C., as buffalo heads have been found
engraved on a bronze stick from the same period (11,46). This is
a black-colored breed, with short horns growing backward and a
population size of ∼600,000 head. It is found in Iran, Azerbaijan,
and along the Caspian Sea; it was widespread in Georgia and
Armenia until 1940, but then its numbers declined (47).
The Khuzestan buffalo (sometimes called Khuzestani) is a
breed amounting to hundreds of thousands head raised in Iraq
and Iran. They are large animals, probably the biggest buffalo
breed in the Near East, with short horns pointing upward and
forming a ring at the end (7,46). Raised mostly for milk purposes,
the average milk production of these animals is ∼1,950 kg of
milk within 210–240 days of lactation, with a reported average
of 8.6 kg/head/day for Khuzestan buffaloes (48). The Azeri
and Khuzestan buffaloes, the most common indigenous breeds
of Iran, present genomic regions associated with adaptation
to different environments resulting from divergent selection in
different regions (21,49). In Iran, there is also the Mazandarani
breed raised mainly in the Mazandran province (in the north
region of the county) with 4,000 individuals (47).
The Egyptian breed was introduced to Egypt from India,
Iran, and Iraq, probably around the seventh century B.C. The
differences between the Egyptian buffalo types depend only on
the environment. This breed is very important in the dairy
industry in Egypt, as it produces 45% of the milk consumed in
the country (50). These animals are blackish gray in color, with
long and narrow heads, and horns varying in format (from lyre
to sword-shaped) (7).
WATER BUFFALO HERD
The most robust information regarding the buffalo population
is available by the FAO and is collected exclusively from
official governmental institution records (51). When there is
an absence of official records, the FAO statistics do not count
the buffalo herds in that country, which produces a lack of
consolidated information regarding the number of countries
with buffaloes and the number of buffalo herds worldwide. For
instance, FAO statistics indicate that only three countries in
the Americas have buffalo populations (Brazil, Trinidad and
Tobago, and Suriname), excluding two large buffalo herds in
America, Colombia, and Venezuela. Therefore, we conducted
an extensive literature review to provide a more accurate count
of buffalo herds and to establish the buffalo distribution in the
different continents.
The worldwide water buffalo population reported by the
FAO in 2017 was ∼201 million head with an expansion to
206.6 million in 2018, showing an increase of 2.8% in 1 year.
The FAO data includes 43 countries with buffalo populations
but did not present an actual herd number in two of those
listed countries, Australia and Singapore. There are some reports
of buffaloes distributed in 129 countries, first mentioned by
Iamartino et al. (52) and replicated by Rahmaninia et al. (47),
but this information is inaccurate. The number 129 represented
the total number of countries involved in the FAO report
and does not necessarily mean that buffaloes exist in each
of those countries. Thus, taking as a starting point these
FAO data from 2018 and researching recent literature, we
retrieved a list of 77 countries that have buffalo herds, with an
estimation of 208,098,759 head. Table 2 presents our aggregate
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Minervino et al. Bubalus bubalis: A Short Story
TABLE 2 | Current data on buffalo population worldwide.
Region/country Water buffalo population†
Worldwide 208,098,759
Africa (%) 3,509,256 (1.69%)
Egypt 3,506,061
Mauritius 25
Mozambique 1,000
South Africa 170
Tanzania 2,000
Americas (%) 2,562,711 (1.23%)
Argentina 121,276
Belize 632
Bolivia 35,000
Brazil 1,390,066
Canada 1,120
Chile 100
Colombia 336,417
Costa Rica 20,000
Cuba 60,000
Ecuador 10,000
El Salvador 250
Guatemala 5,000
Honduras 1,500
Mexico 45,000
Nicaragua 800
Panama 4,000
Paraguay 15,000
Peru 1,500
Suriname 905
Trinidad and Tobago 6,145
United States of America 7,000
Uruguay 1,000
Venezuela 500,000
Asia (%) 201,428,230 (96.79%)
Armenia 717
Azerbaijan 176,195
Bangladesh 1,485,000
Bhutan 531
Brunei 2,319
Cambodia 651,945
China 27,116,250
East Timor 125,760
Georgia 18,361
Hong Kong 329
India 114,151,770
Indonesia 894,278
Iran 199,000
Iraq 300,000
Jordan 95
Kazakhstan 10,414
Laos 1,200,040
Malaysia 117,707
Myanmar 3,790,031
(Continued)
TABLE 2 | Continued
Region/country Water buffalo population†
Nepal 5,277,819
Pakistan 38,848,000
Philippines 2,882,655
Sri Lanka 308,790
Syria 8,000
Taiwan 2,057
Tajikistan 15,351
Thailand 1,258,272
Turkey 161,439
Vietnam 2,425,105
Europe (%) 460,795 (0.22%)
Albania 95
Bulgaria 12,809
Germany 9,613
Greece 9,239
Hungary 1,000
Italy 402,796
Kosovo 400
North Macedonia 643
Poland 69
Romania 14,000
Russia 5,311
Serbia 1,000
Switzerland 1,200
United Kingdom 2,500
Ukraine 120
Oceania (%) 137,767 (0.07%)
Australia 133,000
Guam 94
Micronesia 173
New Zealand 1,000
Papua New Guinea 3,500
†The majority of data was obtained at FAOSTAT (51). Additional sources: (7,27,33,47,
53–77). Detailed methodology is presented in the Supplementary Table 1.
data from buffalo herds worldwide. Supplementary Table 1
provides detailed information regarding the source of the buffalo
head count presented.
Notably, the absence of official data regarding buffalo herds in
most countries mandates the retrieval of estimates from buffalo
studies. Even developed countries (high-income economies),
such as Canada or New Zealand, do not have any data
available regarding their buffalo populations, according to their
respective ministries of agriculture. Despite the absence of official
information, it was possible to retrieve data on the global
distribution and head count of buffaloes with some degree
of confidence, mainly owing to our contacts with breeders’
associations and with fellow scientists around the globe. Our
data may be unreliable for two major countries, Australia and
Venezuela. Australia, in particular, has two distinct herds, one
feral water buffalo herd and one domesticated herd, which have
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Minervino et al. Bubalus bubalis: A Short Story
FIGURE 4 | Water buffalo population worldwide. Legend indicate number of head in each country; t, thousand; m, million. Data source available at
Supplementary Table 1.
been increasing in numbers, owing to the use of animals for
dairy purposes and the increase in animal imports in recent years
(78). Recent reports estimate the population in the Northern
Territory of Australia to be ∼70,000 animals; however, the
herd sizes of Western Australia and Queensland are not known
(79). The number of feral buffaloes in Australia was recently
estimated at 120,000 animals by Lemcke (2019) apud Zhang (27).
The Australian Buffalo Industry Council estimates that there
are ∼15,000–20,000 domestic water buffaloes in the country;
however, a recent report projected 13,000 farmed buffaloes (27).
Therefore, the best estimate is for 133,000 buffalo head in
Australia (27).
Venezuela has an increasing buffalo herd that is mostly raised
in the wetland of the Llanos region, representing the second
largest herd in the Americas, after Brazil (80). The herd evolved
from 180,000 (81) to 350,000 animals (33). A recent report
of the Venezuelan Buffalo Breeders Association reported the
existence of 500,000 animals belonging to more than 190 farmers,
with unverified reports suggesting that there are more than
1.8 million buffalo head in Venezuela (53). Considering the
absence of official data, we considered the data reported in the
aforementioned recent report. Considering the current economic
situation in Venezuela, it is uncertain if this population still
stands. Based on our aggregate data, we created a world map with
the estimated distribution of water buffalo herds (Figure 4).
Buffalo herds have been subjected to major changes in recent
history. Supplementary Video 1 presents the evolution of buffalo
herds in different countries from 1962 to 2017 using FAO
data (51). Regarding the buffalo population trends, Figure 5A
presents the water buffalo herd evolution in the past 50 years
using data from 1968 to 2018. During these 50 years, buffalo
herds worldwide almost doubled, with a population increase
of 97.9%. Compared with cattle, buffalo herds had a superior
trend of increase (considering the percentages), as cattle herds
increased by ∼40% from 1968 to 2018. Although the buffalo
herd had a greater proportional increase, the cattle population
is far more numerous. The cattle herd in 1968 was around
1.06 billion reaching 1.49 billion in 2018, with an increase
in the past 50 years of 425 million head (51). An evaluation
of each 10-year period from 1968 to 2018 showed that the
global buffalo count increased steadily at a mean rate of 14.7
±5.0% per 10 years. The 10-year increase was ∼11% for
the periods that ended in 1978, 1998, and 2018, with sharper
increases in 1988 (23%) and 2008 (15.6%). An evaluation of
the buffalo population trends from the past 50 years, depending
on the continent (Africa, the Americas, Asia, and Europe) is
shown in Figure 5B. It can be noted that the buffalo population
of Asia has a constantly increasing trend, while the buffalo
population of Europe decreased until 1998, at which point it
started to increase, but without reaching the original herd count
of 1968. This phenomenon could be attributed to the fact that
in Europe, only Italy constantly increased its buffalo population
because of its dairy breed, cheese industry, and a market with
a strong economy; on the other hand, in Eastern European
countries, where buffaloes were used as draft animals for land
work or carriages and had low milk yield, they were replaced
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Minervino et al. Bubalus bubalis: A Short Story
FIGURE 5 | Worldwide buffalo population trends: (A) worldwide buffalo
population from the past 50 years (1968–2018); (B) worldwide buffalo
population from the past 50 years (1968–2018) separated by continent. The
right red axis indicates the count for Asia. America, Africa, and Europe are
represented in the left axis. Source FAOSTAT (51).
by Friesian cows and mechanization. Similarly, in Asia, India,
and Pakistan, the dairy breed populations increased rapidly,
while in southeastern countries, the swamp buffalo populations
decreased. Countries in the Americas and Africa experienced
increasing trends; however, both areas experienced a 10-year
period when their buffalo populations decreased (1988–1998 in
the Americas and 2008–2018 in Africa).
BUFFALO MILK PRODUCTION
In many countries, buffalo breeding has a family character in
which the herds are represented by a few animals used for
subsistence. In many societies, buffalo milk is acquiring more and
more visibility linked to its intrinsic value as a dairy product, as it
can not only be consumed directly, as is the case in the Asian
continent, but can also be transformed into dairy products of
high commercial demand (36,40). According to FAO data, the
buffalo milk production accounted for ∼15.14% of the global
fresh milk production in 2018, while cattle milk accounted for
more than 80% (Table 3). Supplementary Video 3 presents the
buffalo milk production (in tons of milk per year) in different
countries from 1962 to 2017 using FAO data (51).
The comparison of these percentages within each continent
produces interesting scenarios. For example, in Asia where
buffalo milk accounts for 35.30% of the total milk production,
buffaloes are the major milk-producing animal in India and
Pakistan; additionally, there are higher populations of dairy
buffaloes than dairy cows in Egypt and Nepal (51). The top 10
largest producers of buffalo milk worldwide (India, Pakistan,
China, Egypt, Nepal, Italy, Myanmar, Iran, Colombia, and Brazil)
are responsible for 97.59% of the global production (Table 4). It
should be considered that, given the lack of production data for
over 50 of the 77 countries in which buffaloes are bred, these
values represent an estimate of the real production potential of
the species; in fact, of 208 million buffaloes, 7.48 million are
TABLE 3 | Buffalo, camel, cow, goat, and sheep milk whole fresh produced in 2018 for each region.
Milk, whole fresh†Buffalo Camel Cow Goat Sheep Total
World Tons 127,658,734 3,137,071 683,217,056 18,712,088 10,631,058 843,356,007
% 15.14% 0.37% 81.01% 2.22% 1.26% 100.00%
Asia Tons 124,958,493 262,233 213,201,098 10,627,509 4,924,398 353,973,731
% 35.30% 0.07% 60.23% 3.00% 1.39% 100.00%
America Tons 189,739 NA 184,304,156 779,806 90,871 185,364,572
% 0.10% NA 99.43% 0.42% 0.05% 100.00%
Europe Tons 390,137 76 220,377,066 2,722,332 3,168,166 226,657,777
% 0.17% 0.00% 97.23% 1.20% 1.40% 100.00%
Oceania Tons NA NA 30,706,258 39 NA 30,706,297
% – – 100.00% 0.00% – 100.00%
Africa Tons 2,120,365 2,874,762 34,628,478 4,582,402 2,447,623 46,653,630
4.54% 6.16% 74.22% 9.82% 5.25% 100.00%
†Data from 2018 (51). The percentage (%) of animal species milk was calculated according to each geographical area. NA, not available.
Frontiers in Veterinary Science | www.frontiersin.org 11 December 2020 | Volume 7 | Article 570413
Minervino et al. Bubalus bubalis: A Short Story
bred in countries in which there are no reported data for buffalo
milk production.
To date, only Italy has established an official system for the
traceability of milk production and the entire buffalo supply
chain since 2014 (83), which provides real-time information on
milk production. The establishment of this system was favored
by the economic value that the supply chain represents for the
country. In fact, Italy, which ranks fifth globally in terms of its
milk production, produces 97% of the buffalo milk produced in
Europe (Table 4); only the buffalo mozzarella that is protected
under the EU’s Protected Designation of Origin (PDO) scheme,
is estimated at a sales value of 766 million euros (84).
According to FAO data, the global buffalo milk production
increased by 32.57% from 2011 to 2018 (from 96 to 127 million
tons); on the contrary, cattle milk increased by 10.67% (from
617 to 683 million tons). Many factors that characterize the
various breeding activities worldwide make it very complex to
estimate the actual production of fresh buffalo milk. The breed,
production purpose (milk, meat, work), breeding type (extensive
and intensive), feeding techniques, seasonality at different
latitudes, age at first birth, weaning techniques, reproductive
techniques applied, lactation duration, milking of the subjects,
genetic selection applied, and the longevity owing to economic
sustainability linked to the commercial value of the milk are some
of these factors (42,85).
There are many factors that affect the milk yield in different
species; breed, genetic background, season and period of calving,
health status, and environmental factors such as feeding, climate
conditions, and welfare are some of these factors. These factors
are also important in buffalo species; however, the genetic
potential is the most important factor, as exemplified in the
Italian Mediterranean, Murrah, and Nili-Ravi breeds. A lot
of work has underwent in selecting the most appropriate
female lines by recording animals and creating genetic centers
with selected bulls for the spread of artificial insemination.
Milk production during lactation is a continuous physiological
function. In buffalo species, the lactation curve, which is a
graphical representation of milk yield during lactation, shows
a rapid increase, peaks, reaches a plateau phase, and declines
gradually until the end of lactation (86). In economic terms,
the most important phase is the persistence of lactation, which
means the ability to maintain milk production at high levels after
the peak of lactation (87). Genetic and environmental influences
on the persistence of lactation were studied in Indian Murrah
buffaloes by Geetha et al. (88) and in Nili-Ravi buffaloes by
Chaudhry et al. (89), who showed the effect of parity and of
lactation length on persistency. The effects of parity, age, and
calving season were also studied in the Bulgarian (90) and the
Italian Mediterranean breeds (91,92).
Compared to cow’s milk, buffalo milk is approximately twice
as high in fat content and ∼30% higher in total solids; therefore,
if the calculation was made on a dry matter basis, the importance
of buffalo milk worldwide would be even greater (93). In terms
of energy, 1 kg of buffalo milk equals 5.10 Mj, which is much
higher than the 2.90 Mj/kg of 1 kg of cow’s milk (94). The water
buffalo is of the most productive domestic animals, has a longer
productive life when compared to cattle, and is economically
TABLE 4 | Current data on buffalo milk production worldwide.
Region/country Milk productionα%βReferences
Worldwide 127,658,734 100.00%
Africa 2,120,365 1.66% †
Egypt 2,120,365 100.00% †
Americas 189,739 0.15%
Brazil 87,472 46.10% Data estimated (82)
Colombia 102,267 53.90% Data estimated (76)
Asia 124,958,493 97.88%
Bangladesh 35,691 0.03% †
Bhutan 282 0.00% †
Brunei 184 0.00% †
China 3,003,323 2.40% †
Georgia 6,186 0.00% †
India 91,817,140 73.48% Official dataγ
Indonesia 71,166 0.06% †
Iran 129,904 0.10% †
Iraq 49,893 0.04% †
Malaysia 8,190 0.01% †
Myanmar 193,841 0.16% †
Nepal 1,338,277 1.07% Official dataγ
Pakistan 28,109,000 22.49% Official dataγ
Sri Lanka 85,914 0.07% Official dataγ
Syrian 6,300 0.01% Official dataγ
Turkey 75,742 0.06% Official dataγ
Viet Nam 27,460 0.02% †
Europe 390,137 0.31%
Albania 12 0.00% †
Bulgaria 11,753 3.01% Official dataγ
Greece 402 0.10% †
Italy 377,970 96.88% Official data (65)
αWater buffalo milk production worldwide in tons (2018 data). βPercentage is related to the
contribution of each continent to the global buffalo milk production and from each country
to their continent total milk production. †FAO data based on imputation methodology (51).
γOfficial data retrieved from Faostat 2019 (51).
important, especially for small-scale producers in developing
countries. Given its characteristics of resilience and adaptability
to tropical climates (95), the buffalo represents an important food
source of animal origin.
Buffalo milk production in several countries is driven for
processing and for the cheese industry. The mozzarella cheese is
recognized and largely consumed worldwide as an adulteration
of the original Mozzarella cheese, which is made exclusively
from buffalo milk. Only in 1993, was mozzarella from buffalo
recognized as “Mozzarella di Bufala Campana PDO.” The PDO
certification pertains to buffaloes bred in the Campania and Lazio
regions of Italy that produce the milk used for the production
of the famous mozzarella cheese; what this essentially means is
that this cheese has to be produced in the defined areas only
from fresh buffalo milk from the Italian Mediterranean breed,
registered in the Buffalo Genealogical Book (36). The mozzarella
industry in Italy resulted from 34,990 recorded females of the
Italian Mediterranean breed, which account for ∼30% of the total
dairy buffalo population (this percentage does not exist in any
Frontiers in Veterinary Science | www.frontiersin.org 12 December 2020 | Volume 7 | Article 570413
Minervino et al. Bubalus bubalis: A Short Story
other country) and have a mean production of 2,356 kg milk in
270 days of lactation, with 8% fat and 4.63% protein (96).
CONCLUSION
There is a surprising lack of data regarding buffalo herds
worldwide. In this study, we reviewed the literature and
established a more appropriate buffalo distribution and
population count. Our results showed that buffaloes are present
in 77 countries in five continents with a population of more
than 208 million head. Considering buffalo characteristics such
as rusticity and productivity, we believe that buffalo production
should be expanded worldwide, especially in developing
countries with adequate natural conditions. This amazing and
subutilized animal can become more productive, and should be
promoted as a target species to be used in smallholder production
systems. Governments worldwide must look carefully to this
amazing animal and start developing projects to introduce
this species in regions with favorable conditions, in order to
substitute cattle herds.
AUTHOR CONTRIBUTIONS
All the authors conceived, research, write, and revised
this manuscript.
FUNDING
This work was supported by Ricerca Corrente IZS ME 8/18 RC:
Study of animal welfare and sustainability of the dairy buffalo
production chain through a multidisciplinary approach CUP
C75B18000630001. Financed by the Italian Ministry of Health.
ACKNOWLEDGMENTS
This work is dedicated to all buffalo farmers around the
world, with a special honor to Victor Makol Julien “o reizinho
do búfalo” (in memorian). Those passionate and dedicated
workers are the main responsible for the buffalo expansion
and production. AM is grateful to CNPq for his research
productivity fellowship.
SUPPLEMENTARY MATERIAL
The Supplementary Material for this article can be found
online at: https://www.frontiersin.org/articles/10.3389/fvets.
2020.570413/full#supplementary-material
Supplementary Table 1 | Sources and methodology for the aggregate data of
the world population of water buffalo (Bubalus bubalis) per country.
Supplementary Video 1 | This video presents the buffalo herd evolution (in
number of animals) in the different countries from 1962 to 2017 using the data
from FAO (51). Video produced through FlourishTM platform (Kiln Enterprises Ltd,
London, UK). Available online at: https://flourish.studio/.
Supplementary Video 2 | Herd of docile and curious mixed-breed buffaloes in
Santarém, Amazon, Brazil.
Supplementary Video 3 | This video presents the buffalo milk production (in tons
of milk per year) in the different countries from 1962 to 2017 using the data from
FAO (51). Video produced through FlourishTM platform (Kiln Enterprises Ltd,
London, UK). Available online at: https://flourish.studio/.
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Conflict of Interest: The authors declare that the research was conducted in the
absence of any commercial or financial relationships that could be construed as a
potential conflict of interest.
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