Lumpy Skin Disease
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Lumpy skin disease (LSD) is an infectious disease in cattle caused by Lumpy skin disease virus of the family ''
Poxviridae ''Poxviridae'' is a family of double-stranded DNA viruses. Vertebrates and arthropods serve as natural hosts. The family contains 22 genera that are assigned to two subfamilies: ''Chordopoxvirinae'' and ''Entomopoxvirinae''. ''Entomopoxvirinae'' ...
'', also known as Neethling virus. The disease is characterized by fever, enlarged superficial
lymph node A lymph node, or lymph gland, is a kidney-shaped organ of the lymphatic system and the adaptive immune system. A large number of lymph nodes are linked throughout the body by the lymphatic vessels. They are major sites of lymphocytes that includ ...
s, and multiple
nodules Nodule may refer to: *Nodule (geology), a small rock or mineral cluster *Manganese nodule, a metallic concretion found on the seafloor *Nodule (medicine), a small aggregation of cells *Root nodule Root nodules are found on the roots of plants, p ...
(measuring in diameter) on the skin and
mucous membrane A mucous membrane or mucosa is a membrane that lines various cavities in the body of an organism and covers the surface of internal organs. It consists of one or more layers of epithelial cells overlying a layer of loose connective tissue. It ...
s, including those of the
respiratory The respiratory system (also respiratory apparatus, ventilatory system) is a biological system consisting of specific organs and structures used for gas exchange in animals and plants. The anatomy and physiology that make this happen varies gr ...
and
gastrointestinal The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the digestive system that leads from the mouth to the anus. The tract is the largest of the body's systems, after the cardiovascular system. ...
tracts. Infected cattle may also develop
edema Edema (American English), also spelled oedema (British English), and also known as fluid retention, swelling, dropsy and hydropsy, is the build-up of fluid in the body's tissue (biology), tissue. Most commonly, the legs or arms are affected. S ...
tous swelling in their limbs and exhibit
lameness A limp is a type of asymmetric abnormality of the gait. Limping may be caused by pain, weakness, neuromuscular imbalance, or a skeletal deformity. The most common underlying cause of a painful limp is physical trauma; however, in the absence ...
. The virus has important economic implications since affected animals tend to have permanent damage to their skin, lowering the commercial value of their hide. Additionally, the disease often results in chronic debility, reduced milk production, poor growth, infertility, abortion, and sometimes death. Onset of fever occurs almost one week after infection by the virus. This initial fever may exceed and persist for one week. At this time, all of the superficial lymph nodes become enlarged. The nodules, which the disease is characterized by, appear seven to nineteen days after virus inoculation. Coinciding with the appearance of the nodules, discharge from the eyes and nose becomes mucopurulent. The nodular lesions involve the
dermis The dermis or corium is a layer of skin between the epidermis (skin), epidermis (with which it makes up the cutis (anatomy), cutis) and subcutaneous tissues, that primarily consists of dense irregular connective tissue and cushions the body from s ...
and the
epidermis The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and Subcutaneous tissue, hypodermis. The epidermal layer provides a barrier to infection from environmental pathogens and regulates the ...
, but may extend to the underlying
subcutis The subcutaneous tissue (), also called the hypodermis, hypoderm (), subcutis, or superficial fascia, is the lowermost layer of the integumentary system in vertebrates. The types of cells found in the layer are fibroblasts, adipose cells, and ma ...
or even to the muscle. These lesions, occurring all over the body (but particularly on the head, neck,
udder An udder is an organ formed of two or four mammary glands on the females of dairy animals and ruminants such as cattle, goats, and sheep. An udder is equivalent to the breast in primates, elephantine pachyderms and other mammals. The udder is ...
,
scrotum In most terrestrial mammals, the scrotum (: scrotums or scrota; possibly from Latin ''scortum'', meaning "hide" or "skin") or scrotal sac is a part of the external male genitalia located at the base of the penis. It consists of a sac of skin ...
,
vulva In mammals, the vulva (: vulvas or vulvae) comprises mostly external, visible structures of the female sex organ, genitalia leading into the interior of the female reproductive tract. For humans, it includes the mons pubis, labia majora, lab ...
, and
perineum The perineum (: perineums or perinea) in placentalia, placental mammals is the space between the anus and the genitals. The human perineum is between the anus and scrotum in the male or between the anus and vulva in the female. The perineum is ...
), may be either well-circumscribed or they may coalesce. Cutaneous lesions may be resolved rapidly or they may persist as hard lumps. The lesions can also become sequestrated, leaving deep
ulcer An ulcer is a discontinuity or break in a bodily membrane that impedes normal function of the affected organ. According to Robbins's pathology, "ulcer is the breach of the continuity of skin, epithelium or mucous membrane caused by sloughin ...
s filled with
granulation tissue Granulation tissue is new connective tissue and microscopic blood vessels that form on the surfaces of a wound during the healing process. Granulation tissue typically grows from the base of a wound and is able to fill wounds of almost any size ...
and often suppurating (forming
pus Pus is an exudate, typically white-yellow, yellow, or yellow-brown, formed at the site of inflammation during infections, regardless of cause. An accumulation of pus in an enclosed tissue space is known as an abscess, whereas a visible collect ...
). At the initial onset of the nodules, they have a creamy grey to white color upon cut section, and may exude serum. After about two weeks, a cone-shaped central core of
necrotic Necrosis () is a form of cell injury which results in the premature death of cells in living tissue by autolysis. The term "necrosis" came about in the mid-19th century and is commonly attributed to German pathologist Rudolf Virchow, who is ...
material may appear within the nodules. Additionally, the nodules on the mucous membranes of the eyes, nose, mouth,
rectum The rectum (: rectums or recta) is the final straight portion of the large intestine in humans and some other mammals, and the gut in others. Before expulsion through the anus or cloaca, the rectum stores the feces temporarily. The adult ...
, udder and genitalia quickly ulcerate, aiding in transmission of the virus. In mild cases of LSD, the clinical symptoms and lesions are often confused with Bovine Herpesvirus 2 (BHV-2), which is, in turn, referred to as pseudo-lumpy skin disease. However, the lesions associated with BHV-2 infections are more superficial. BHV-2 also has a shorter course and is more mild than LSD.
Electron microscopy An electron microscope is a microscope that uses a beam of electrons as a source of illumination. It uses electron optics that are analogous to the glass lenses of an optical light microscope to control the electron beam, for instance focusing i ...
can be used to differentiate between the two infections. BHV-2 is characterized by intranuclear
inclusion bodies Inclusion bodies are aggregates of specific types of protein found in neurons, and a number of tissue (biology), tissue cells including red blood cells, bacteria, viruses, and plants. Inclusion bodies of aggregations of multiple proteins are also ...
, as opposed to the intracytoplasmic inclusions characteristic of LSD. Isolation of BHV-2, or its detection in negatively-stained
biopsy A biopsy is a medical test commonly performed by a surgeon, interventional radiologist, an interventional radiologist, or an interventional cardiology, interventional cardiologist. The process involves the extraction of sampling (medicine), sample ...
specimens, is only possible approximately one week after the development of skin lesions.


Lumpy skin disease virus


Classification

Lumpy skin disease virus (LSDV) is double-stranded
DNA virus A DNA virus is a virus that has a genome made of deoxyribonucleic acid (DNA) that is replicated by a DNA polymerase. They can be divided between those that have two strands of DNA in their genome, called double-stranded DNA (dsDNA) viruses, and t ...
. It is a member of the capripoxvirus genus of ''
Poxviridae ''Poxviridae'' is a family of double-stranded DNA viruses. Vertebrates and arthropods serve as natural hosts. The family contains 22 genera that are assigned to two subfamilies: ''Chordopoxvirinae'' and ''Entomopoxvirinae''. ''Entomopoxvirinae'' ...
''. Capripoxviruses (CaPVs) represent one of eight genera within the Chordopoxvirus (ChPV) subfamily. The capripoxvirus genus consists of LSDV, as well as sheeppox virus, and goatpox virus. CaPV infections are usually host-specific within specific geographic distributions even though they are serologically indistinguishable from one another.


Structure

Like other viruses in the ''Poxviridae'' family, capripoxviruses are brick-shaped. Capripoxvirus virions are different than
orthopoxvirus ''Orthopoxvirus'' is a genus of viruses in the family ''Poxviridae'' and subfamily ''Chordopoxvirinae''. Vertebrates, including mammals and humans, and arthropods serve as natural hosts. There are 12 species in this genus. Diseases associated wi ...
virions in that they have a more oval profile, as well as larger lateral bodies. The average size of capripoxvirions is 320 nm by 260 nm.


Genome

The virus has a 151-kbp
genome A genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding genes, other functional regions of the genome such as ...
, consisting of a central coding region which is bounded by identical 2.4 kbp-inverted terminal repeats and contains 156 genes. There are 146 conserved genes when comparing LSDV with chordopoxviruses of other genera. These genes encode proteins which are involved in transcription and
mRNA In molecular biology, messenger ribonucleic acid (mRNA) is a single-stranded molecule of RNA that corresponds to the genetic sequence of a gene, and is read by a ribosome in the process of Protein biosynthesis, synthesizing a protein. mRNA is ...
biogenesis,
nucleotide Nucleotides are Organic compound, organic molecules composed of a nitrogenous base, a pentose sugar and a phosphate. They serve as monomeric units of the nucleic acid polymers – deoxyribonucleic acid (DNA) and ribonucleic acid (RNA), both o ...
metabolism,
DNA replication In molecular biology, DNA replication is the biological process of producing two identical replicas of DNA from one original DNA molecule. DNA replication occurs in all life, living organisms, acting as the most essential part of heredity, biolog ...
, protein processing, virion structure and assembly, and viral
virulence Virulence is a pathogen's or microorganism's ability to cause damage to a host. In most cases, especially in animal systems, virulence refers to the degree of damage caused by a microbe to its host. The pathogenicity of an organism—its abili ...
and host range. Within the central genomic region, LSDV genes share a high degree of
collinearity In geometry, collinearity of a set of points is the property of their lying on a single line. A set of points with this property is said to be collinear (sometimes spelled as colinear). In greater generality, the term has been used for aligned ...
and amino acid identity with the genes of other mammalian poxviruses. Examples of viruses with similar amino acid identity include suipoxvirus,
yatapoxvirus ''Yatapoxvirus'' is a genus of viruses, in the family ''Poxviridae'', in the subfamily ''Chordopoxvirinae''. Monkeys and Baboon, baboons serve as natural hosts. There are two species in this genus. Diseases associated with this genus can cause H ...
, and leporipoxvirus. In terminal regions, however, collinearity is interrupted. In these regions, poxvirus homologues are either absent or share a lower percentage of amino acid identity. Most of these differences involve genes that are likely associated with viral virulence and host range. Unique to Chordopoxviridae, LSDV contains homologues of interleukin-10 (IL-10), IL-1 binding proteins, G protein-coupled CC
chemokine receptor Chemokine receptors are cytokine receptors found on the surface of certain cells that interact with a type of cytokine called a chemokine. There have been 20 distinct chemokine receptors discovered in humans. Each has a rhodopsin-like 7-tran ...
, and epidermal growth factor-like protein, which are found in other poxvirus genera.


Epidemiology

LSDV mainly affects
cattle Cattle (''Bos taurus'') are large, domesticated, bovid ungulates widely kept as livestock. They are prominent modern members of the subfamily Bovinae and the most widespread species of the genus '' Bos''. Mature female cattle are calle ...
and
zebu The zebu (; ''Bos indicus''), also known as indicine cattle and humped cattle, is a species or subspecies of Bos taurus, domestic cattle originating in South Asia. Zebu, like many Sanga cattle breeds, differs from taurine cattle by a fatty hump ...
s, but has also been seen in
giraffe The giraffe is a large Fauna of Africa, African even-toed ungulate, hoofed mammal belonging to the genus ''Giraffa.'' It is the Largest mammals#Even-toed Ungulates (Artiodactyla), tallest living terrestrial animal and the largest ruminant on ...
s,
water buffalo The water buffalo (''Bubalus bubalis''), also called domestic water buffalo, Asian water buffalo and Asiatic water buffalo, is a large bovid originating in the Indian subcontinent and Southeast Asia. Today, it is also kept in Italy, the Balkans ...
, and
impala The impala or rooibok (''Aepyceros melampus'', lit. 'black-footed high-horn' in Ancient Greek) is a medium-sized antelope found in eastern and southern Africa. The only extant member of the genus '' Aepyceros'', and tribe Aepycerotini, it ...
s. Fine-skinned ''
Bos taurus Cattle (''Bos taurus'') are large, domesticated, bovid ungulates widely kept as livestock. They are prominent modern members of the subfamily Bovinae and the most widespread species of the genus '' Bos''. Mature female cattle are called ...
'' cattle breeds such as Holstein-Friesian and
Jersey Jersey ( ; ), officially the Bailiwick of Jersey, is an autonomous and self-governing island territory of the British Islands. Although as a British Crown Dependency it is not a sovereign state, it has its own distinguishing civil and gov ...
are the most susceptible to the disease. Thick-skinned '' Bos indicus'' breeds including the
Afrikaner Afrikaners () are a Southern African ethnic group descended from predominantly Dutch settlers who first arrived at the Cape of Good Hope in 1652.Entry: Cape Colony. ''Encyclopædia Britannica Volume 4 Part 2: Brain to Casting''. Encyclopæd ...
and Afrikaner cross-breeds show less severe signs of the disease. This is probably due to the decreased susceptibility to
ectoparasites Parasitism is a close relationship between species, where one organism, the parasite, lives (at least some of the time) on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The en ...
that ''Bos indicus'' breeds exhibit relative to ''Bos taurus'' breeds. Young calves and cows at peak
lactation Lactation describes the secretion of milk from the mammary glands and the period of time that a mother lactates to feed her young. The process naturally occurs with all sexually mature female mammals, although it may predate mammals. The process ...
show more severe clinical symptoms, but all age-groups are susceptible to the disease.


Transmission

Outbreaks of LSDV are associated with high temperature and high humidity. It is usually more prevalent during wet summer and autumn months, especially in low-lying areas or near bodies of water; however, outbreaks can also occur during the dry season. Blood-feeding insects such as mosquitos and flies act as mechanical vectors to spread the disease. A single species vector has not been identified. Instead, the virus has been isolated from '' Stomoxys'', ''Biomyia fasciata'',
Tabanidae Horse flies and deer flies are true flies in the family Tabanidae in the insect order Diptera. The adults are often large and agile in flight. Only females bite land vertebrates, including humans, to obtain blood. They prefer to fly in sunli ...
, '' Glossina'', and ''
Culicoides ''Culicoides'' is a genus of biting midges in the family Ceratopogonidae. There are over 1000 species in the genus,Connelly, C. RBiting midges: ''Culicoides'' spp.Featured Creatures, Entomology and Nematology Department, University of Florida IF ...
'' species. The particular role of each of these insects in the transmission of LSDV continues to be evaluated. Outbreaks of lumpy skin disease tend to be sporadic since they are dependent upon animal movements, immune status and wind and rainfall patterns, which affect the vector populations. The virus can be transmitted through blood, nasal discharge, lacrimal (tear duct) secretions, semen, and
saliva Saliva (commonly referred as spit or drool) is an extracellular fluid produced and secreted by salivary glands in the mouth. In humans, saliva is around 99% water, plus electrolytes, mucus, white blood cells, epithelial cells (from which ...
. The disease can also be transmitted through infected milk to suckling calves. In experimentally infected cattle, LSDV was found in saliva 11 days after the development of fever, in semen after 22 days, and in skin nodules after 33 days. The virus is not found in urine or stool. Like other pox viruses, which are known to be highly resistant, LSDV can remain viable in infected tissue for more than 120 days.


Immunity


Artificial immunity

There have been two different approaches to
immunization Immunization, or immunisation, is the process by which an individual's immune system becomes fortified against an infectious agent (known as the antigen, immunogen). When this system is exposed to molecules that are foreign to the body, called ' ...
against LSDV. In
South Africa South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Africa. Its Provinces of South Africa, nine provinces are bounded to the south by of coastline that stretches along the Atlantic O ...
, the Neethling strain of the virus was first attenuated by 20 passages on the chorio-allantoic membranes of hens' eggs. Now the vaccine virus is propagated in
cell culture Cell culture or tissue culture is the process by which cell (biology), cells are grown under controlled conditions, generally outside of their natural environment. After cells of interest have been Cell isolation, isolated from living tissue, ...
. In
Kenya Kenya, officially the Republic of Kenya, is a country located in East Africa. With an estimated population of more than 52.4 million as of mid-2024, Kenya is the 27th-most-populous country in the world and the 7th most populous in Africa. ...
, the vaccine produced from sheep or goatpox viruses has been shown to provide immunity in cattle. However, the level of attenuation required for safe use in sheep and goats is not sufficient for cattle. For this reason the sheeppox and goatpox vaccines are restricted to countries where sheeppox or goatpox is already
endemic Endemism is the state of a species being found only in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also foun ...
since the live vaccines could provide a source of infection for the susceptible sheep and goat populations. In order to ensure adequate protection against LSDV, susceptible adult cattle should be vaccinated annually. Approximately 50% of cattle develop swelling ( in diameter) at the site of inoculation. This swelling disappears within a few weeks. Upon inoculation,
dairy cows Dairy cattle (also called dairy cows) are cattle bred with the ability to produce large quantities of milk, from which dairy products are made. Dairy cattle generally are of the species '' Bos taurus''. Historically, little distinction was ...
may also exhibit a temporary decrease in milk production.


Natural immunity

Most cattle develop lifelong immunity after recovery from a natural infection. Additionally, calves of immune cows acquire maternal antibodies and are resistant to clinical disease until about 6 months of age. To avoid interference with maternal antibodies, calves under 6 months of age whose dams were either naturally infected or were vaccinated should not be vaccinated. On the other hand, calves born from susceptible cows are also susceptible and should be vaccinated.


History

Lumpy skin disease was first seen as an
epidemic An epidemic (from Greek ἐπί ''epi'' "upon or above" and δῆμος ''demos'' "people") is the rapid spread of disease to a large number of hosts in a given population within a short period of time. For example, in meningococcal infection ...
in
Zambia Zambia, officially the Republic of Zambia, is a landlocked country at the crossroads of Central Africa, Central, Southern Africa, Southern and East Africa. It is typically referred to being in South-Central Africa or Southern Africa. It is bor ...
in 1929. Initially, it was thought to be the result of either poisoning or a hypersensitivity to insect bites. Additional cases occurred between 1943 and 1945 in
Botswana Botswana, officially the Republic of Botswana, is a landlocked country in Southern Africa. Botswana is topographically flat, with approximately 70 percent of its territory part of the Kalahari Desert. It is bordered by South Africa to the sou ...
,
Zimbabwe file:Zimbabwe, relief map.jpg, upright=1.22, Zimbabwe, relief map Zimbabwe, officially the Republic of Zimbabwe, is a landlocked country in Southeast Africa, between the Zambezi and Limpopo Rivers, bordered by South Africa to the south, Bots ...
, and South Africa. Approximately, 8 million cattle were affected in a
panzootic A panzootic (from Greek παν all + ζόιον animal) is an epizootic (an outbreak of an infectious disease of non-human animals) that spreads across a large region (for example a continent), or even worldwide. The equivalent in human populati ...
infection in South Africa in 1949, causing enormous economic losses. LSD spread throughout Africa between the 1950s and 1980s, affecting cattle in Kenya,
Sudan Sudan, officially the Republic of the Sudan, is a country in Northeast Africa. It borders the Central African Republic to the southwest, Chad to the west, Libya to the northwest, Egypt to the north, the Red Sea to the east, Eritrea and Ethiopi ...
,
Tanzania Tanzania, officially the United Republic of Tanzania, is a country in East Africa within the African Great Lakes region. It is bordered by Uganda to the northwest; Kenya to the northeast; the Indian Ocean to the east; Mozambique and Malawi to t ...
,
Somalia Somalia, officially the Federal Republic of Somalia, is the easternmost country in continental Africa. The country is located in the Horn of Africa and is bordered by Ethiopia to the west, Djibouti to the northwest, Kenya to the southwest, th ...
, and
Cameroon Cameroon, officially the Republic of Cameroon, is a country in Central Africa. It shares boundaries with Nigeria to the west and north, Chad to the northeast, the Central African Republic to the east, and Equatorial Guinea, Gabon, and the R ...
. In 1989 there was an LSD outbreak in
Israel Israel, officially the State of Israel, is a country in West Asia. It Borders of Israel, shares borders with Lebanon to the north, Syria to the north-east, Jordan to the east, Egypt to the south-west, and the Mediterranean Sea to the west. Isr ...
. This outbreak was the first instance of LSD north of the Sahara desert and outside of the African continent. This particular outbreak was thought to be the result of infected '' Stomoxys calcitrans'' being carried on wind from Ismailiya in Egypt. During a period of 37 days between August and September 1989, fourteen of the seventeen dairy herds in Peduyim became infected with LSD. All of the cattle as well as small flocks of sheep and goats in the village were slaughtered. Throughout the past decade, LSD occurrences have been reported in Middle Eastern, European, and west Asian regions. LSD was first reported to the Bangladesh livestock department in July 2019. Eventually 500,000 head were estimated to have been infected in this outbreak. The
Food and Agriculture Organization The Food and Agriculture Organization of the United Nations; . (FAO) is a specialized agency of the United Nations that leads international efforts to defeat hunger and improve nutrition and food security. Its Latin motto, , translates ...
(FAO) recommended mass vaccination. As a result of the introduction of
fall armyworm The fall armyworm (''Spodoptera frugiperda'') is a species in the order Lepidoptera and one of the species of the fall armyworm moths distinguished by their larval life stage. The term "armyworm" can refer to several species, often describing the ...
and this cattle plague within a few months of each other, the FAO, the
World Food Programme The World Food Programme (WFP) is an international organization within the United Nations that provides food assistance worldwide. It is the world's largest humanitarian organization and the leading provider of school meals. Founded in 1961 ...
,
Bangladesh Government The government of the People's Republic of Bangladesh () is the central government of Bangladesh. The government was constituted by the Constitution of Bangladesh comprising the executive (the president, prime minister and cabinet), the le ...
officials, and others agreed to begin improving Bangladesh's livestock disease surveillance and emergency response capabilities. Method of entry of the virus into Bangladesh remains unknown. In 2022 a lumpy skin disease outbreak in Pakistan killed over 7000 cattle. In India, between July and September 2022 the lumpy skin disease outbreak in India resulted in the death of over 80,000 cattle. The state of
Rajasthan Rajasthan (; Literal translation, lit. 'Land of Kings') is a States and union territories of India, state in northwestern India. It covers or 10.4 per cent of India's total geographical area. It is the List of states and union territories of ...
has seen a majority of the deaths. Inter-
state State most commonly refers to: * State (polity), a centralized political organization that regulates law and society within a territory **Sovereign state, a sovereign polity in international law, commonly referred to as a country **Nation state, a ...
and inter-
district A district is a type of administrative division that in some countries is managed by the local government. Across the world, areas known as "districts" vary greatly in size, spanning regions or county, counties, several municipality, municip ...
movement of cattle in a number of states has been restricted.
Indian Council of Agricultural Research The Indian Council of Agricultural Research (ICAR) is an autonomous body responsible for co-ordinating agricultural education and research in India. It reports to the Department of Agricultural Research and Education, Ministry of Agriculture. Th ...
labs have undertaken creation of an indigenous vaccine. A goat pox vaccine is being used; 15 million doses had been administered by September 2022. There are at least three centres manufacturing the goat pox vaccine in India. Institutions with authority to test have been expanded. In 2023, there was an outbreak of LSD in Nepal, during which more than 50,000 cattle and buffalo died from the disease.


References


External links


Current status of Lumpy skin disease worldwide
at OIE. WAHID Interface - OIE World Animal Health Information Database
Disease card

Lumpy Skin Disease
Food and Agriculture Organization The Food and Agriculture Organization of the United Nations; . (FAO) is a specialized agency of the United Nations that leads international efforts to defeat hunger and improve nutrition and food security. Its Latin motto, , translates ...
of the
United Nations The United Nations (UN) is the Earth, global intergovernmental organization established by the signing of the Charter of the United Nations, UN Charter on 26 June 1945 with the stated purpose of maintaining international peace and internationa ...
{{DEFAULTSORT:Lumpy skin Disease Bovine diseases Chordopoxvirinae Animal viral diseases Virus-related cutaneous conditions Lumpy skin disease