''Orthohantavirus'' is a genus of viruses that includes all hantaviruses (family ''

Hantaviridae ''Hantaviridae'' is a family of viruses in the order ''Bunyavirales''. It is named for the Hantan River area in South Korea South Korea, officially the Republic of Korea (ROK), is a country in East Asia. It constitutes the southern half o ...

'') that cause disease in humans. Orthohantaviruses, hereafter referred to as hantaviruses, are naturally found primarily in

rodent Rodents (from Latin , 'to gnaw') are mammals of the Order (biology), order Rodentia ( ), which are characterized by a single pair of continuously growing incisors in each of the upper and Mandible, lower jaws. About 40% of all mammal specie ...

s. In general, each hantavirus is carried by one rodent species and each rodent that carries a hantavirus carries one hantavirus species. Hantaviruses in their

natural reservoir In Infection, infectious disease ecology and epidemiology, a natural reservoir, also known as a disease reservoir or a reservoir of infection, is the population of organisms or the specific environment in which an infectious pathogen naturally li ...

s usually cause an asymptomatic, persistent infection. In humans, however, hantaviruses cause two diseases:

hemorrhagic fever with renal syndrome Hantavirus hemorrhagic fever with renal syndrome (HFRS) is a hemorrhagic fever caused by hantaviruses. Symptoms usually occur 12–16 days after exposure to the virus and come in five distinct phases: febrile, hypotensive, low urine production (ol ...

(HFRS) and

hantavirus pulmonary syndrome Hantavirus pulmonary syndrome (HPS), also called hantavirus cardiopulmonary syndrome (HCPS), is a severe respiratory disease caused by hantaviruses. The main features of illness are microvascular leakage and acute respiratory distress syndrome. S ...

(HPS). HFRS is mainly caused by hantaviruses in Africa, Asia, and Europe, called Old World hantaviruses, and HPS is usually caused by hantaviruses in the Americas, called New World hantaviruses. Hantaviruses are transmitted mainly through

aerosol An aerosol is a suspension (chemistry), suspension of fine solid particles or liquid Drop (liquid), droplets in air or another gas. Aerosols can be generated from natural or Human impact on the environment, human causes. The term ''aerosol'' co ...

s and droplets that contain rodent excretions, as well as through contaminated food, bites, and scratches. Environmental factors such as rainfall, temperature, and humidity influence transmission. Human-to-human transmission does not occur. HFRS is marked by kidney disease with kidney swelling, excess protein in urine, and blood in urine. The case fatality rate of HFRS varies from less than 1% to 15% depending on the virus. A mild form of HFRS often called nephropathia epidemica is often caused by

Puumala virus Puumala virus (PUUV) is the main cause of hemorrhagic fever with renal syndrome (HFRS) in Europe and Russia. Puumala virus is transmitted by the bank vole (''Clethrionomys glareolus''). In its natural reservoir, PUUV causes a persistent infectio ...

and

Dobrava-Belgrade virus Dobrava-Belgrade virus (DOBV) is the main cause of hemorrhagic fever with renal syndrome (HFRS) in southern Europe. In its natural reservoirs, DOBV causes a persistent, asymptomatic infection and is spread through excretions, fighting, and groom ...

. For HPS, initial symptoms are flu-like, with fever, headache, and muscle pain, followed by sudden respiratory failure. HPS has a higher case fatality rate than HFRS, at 30–60%. For both HFRS and HPS, illness is the result of increased vascular permeability, decreased platelet count, and overreaction of the immune system. The hantavirus genome consists of three single-stranded negative-sense RNA segments that encode one protein each: an

RNA-dependent RNA polymerase RNA-dependent RNA polymerase (RdRp) or RNA replicase is an enzyme that catalyzes the self-replication, replication of RNA from an RNA template. Specifically, it catalyzes synthesis of the RNA strand Complementarity (molecular biology), compleme ...

(RdRp), a spike glycoprotein precursor, and the N protein. Segments are encased in N proteins to form ribonucleoprotein (RNP) complexes that each have a copy of RdRp attached. RNP complexes are surrounded by a lipid envelope that has

spike protein In virology, a spike protein or peplomer protein is a protein that forms a large structure known as a spike or peplomer projecting from the surface of an viral envelope, enveloped virus. as cited in The proteins are usually glycoproteins that ...

s emanating from its surface. Replication begins when spikes attach to the surface of cells. After entering the cell, the envelope fuses with

endosome Endosomes are a collection of intracellular sorting organelles in eukaryotic cells. They are parts of the endocytic membrane transport pathway originating from the trans Golgi network. Molecules or ligands internalized from the plasma membra ...

s and

lysosome A lysosome () is a membrane-bound organelle that is found in all mammalian cells, with the exception of red blood cells (erythrocytes). There are normally hundreds of lysosomes in the cytosol, where they function as the cell’s degradation cent ...

s, which empties RNPs into the cytoplasm. RdRp then transcribes the genome to produce

messenger RNA 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 synthesizing a protein. mRNA is created during the ...

(mRNA) for

translation Translation is the communication of the semantics, meaning of a #Source and target languages, source-language text by means of an Dynamic and formal equivalence, equivalent #Source and target languages, target-language text. The English la ...

by host ribosomes to produce viral proteins and replicates the genome for progeny viruses. Old World hantaviruses assemble in the

Golgi apparatus The Golgi apparatus (), also known as the Golgi complex, Golgi body, or simply the Golgi, is an organelle found in most eukaryotic Cell (biology), cells. Part of the endomembrane system in the cytoplasm, it protein targeting, packages proteins ...

and obtain their envelope from it, before being transported to the cell membrane to leave the cell via

exocytosis Exocytosis is a term for the active transport process that transports large molecules from cell to the extracellular area. Hormones, proteins and neurotransmitters are examples of large molecules that can be transported out of the cell. Exocytosis ...

. New World hantaviruses assemble near the cell membrane and obtain their envelope from it as they leave the cell by budding from its surface. Hantaviruses were first discovered following the

Korean War The Korean War (25 June 1950 – 27 July 1953) was an armed conflict on the Korean Peninsula fought between North Korea (Democratic People's Republic of Korea; DPRK) and South Korea (Republic of Korea; ROK) and their allies. North Korea was s ...

. During the war, HFRS was a common ailment in soldiers stationed near the Hantan river. In 1978 in South Korea, the first hantavirus was isolated,

Hantaan virus Hantaan virus (HTNV) is the main cause of hemorrhagic fever with renal syndrome (HFRS) in East Asia. Hantaan virus is transmitted by the striped field mouse (''Apodemus agrarius'') In its natural reservoir, HTNV causes a persistent, asymptomatic ...

, and was shown to be responsible for the outbreak during the war. Within a few years, other hantaviruses that cause HFRS were discovered throughout Eurasia. In 1982, the

World Health Organization The World Health Organization (WHO) is a list of specialized agencies of the United Nations, specialized agency of the United Nations which coordinates responses to international public health issues and emergencies. It is headquartered in Gen ...

gave HFRS its name, and in 1987, hantaviruses were classified for the first time. They collectively bear the name of Hantaan virus and the Hantan river. In 1993, an outbreak of HPS occurred in the Four Corners region in the United States, which led to the discovery of pathogenic New World hantaviruses and the second disease caused by hantaviruses. Since then, hantaviruses have been found not just in rodents but also moles,

shrew Shrews ( family Soricidae) are small mole-like mammals classified in the order Eulipotyphla. True shrews are not to be confused with treeshrews, otter shrews, elephant shrews, West Indies shrews, or marsupial shrews, which belong to dif ...

s, and

bat Bats are flying mammals of the order Chiroptera (). With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most birds, flying with their very long spread-out ...

Disease

Hantaviruses are sorted into Old World hantaviruses (OWHVs), which typically cause

(HFRS) in Africa, Asia, and Europe, and New World hantaviruses (NWHVs) which are associated with

(HPS) in the Americas. The case fatality rate of HFRS ranges from less than 1% to 15%, while for HPS it is 30–60%. The severity of symptoms of HFRS varies depending on the virus:

causes severe HFRS,

Seoul virus Seoul virus (SEOV) is one of the main causes of hemorrhagic fever with renal syndrome (HFRS). Seoul virus is transmitted by the brown rat (''Rattus norvegicus'') and the black rat (''Rattus rattus''). In its natural reservoirs, SEOV causes an asy ...

moderate HFRS,

mild HFRS, and

infection varies from mild to severe depending on genotype. The mild form of HFRS caused by Puumala virus and Dobrava-Belgrade virus is often called nephropathia epidemica (NE). Repeated infections of hantaviruses have not been observed, so recovering from infection likely grants life-long immunity. HFRS is characterized by five phases: febrile, hypotensive, low urine production (

oliguria Oliguria or hypouresis is the low output of urine specifically more than 80 ml/day but less than 400ml/day. The decreased output of urine may be a sign of dehydration, kidney failure, hypovolemic shock, hyperosmolar Hyperglycemic Nonketotic Syndro ...

), high urine production (

polyuria Polyuria () is excessive or an abnormally large production or Frequent urination, passage of urine (greater than 2.5 L or 3 L over 24 hours in adults). Increased production and passage of urine may also be termed as diuresis. Polyuria often appe ...

), and recovery. Symptoms usually occur 12–16 days after exposure to the virus. Acute kidney disease occurs with kidney swelling, excess protein in urine (

proteinuria Proteinuria is the presence of excess proteins in the urine. In healthy persons, urine contains very little protein, less than 150 mg/day; an excess is suggestive of illness. Excess protein in the urine often causes the urine to become fo ...

), and blood in urine (

hematuria Hematuria or haematuria is defined as the presence of blood or red blood cells in the urine. "Gross hematuria" occurs when urine appears red, brown, or tea-colored due to the presence of blood. Hematuria may also be subtle and only detectable with ...

). Other symptoms include headache, lower back pain, nausea, vomiting, diarrhea, bloody stool, the appearance of spots on the skin (

petechiae A petechia (; : petechiae) is a small red or purple spot ( 1 cm in diameter) and purpura (3 to 10 mm in diameter). The term is typically used in the plural (petechiae), since a single petechia is seldom noticed or significant. Causes Physical ...

), and hemorrhaging in the respiratory tract. Renal failure leads oliguria, and restoration of kidney health comes with polyuria. Recovery typically takes a few months. In more mild cases, the different phases of HFRS may be hard to distinguish, or some phases may be absent, while in more severe cases, the phases may overlap. HPS is mainly caused by two viruses: Andes virus and

Sin Nombre virus Sin Nombre virus (SNV) is the most common cause of hantavirus pulmonary syndrome (HPS) in North America. Sin Nombre virus is transmitted mainly by the western deer mouse (''Peromyscus sonoriensis''). In its natural reservoir, SNV causes an asymp ...

. The disease has three phases: prodromal (early), cardiopulmonary, and recovery. Symptoms occur about 1–8 weeks after exposure to the virus. Early symptoms include fever, headache, muscle pain, shortness of breath (

dyspnea Shortness of breath (SOB), known as dyspnea (in AmE) or dyspnoea (in BrE), is an uncomfortable feeling of not being able to breathe well enough. The American Thoracic Society defines it as "a subjective experience of breathing discomfort that ...

), and low platelet count (

thrombocytopenia In hematology, thrombocytopenia is a condition characterized by abnormally low levels of platelets (also known as thrombocytes) in the blood. Low levels of platelets in turn may lead to prolonged or excessive bleeding. It is the most common coag ...

). During the cardiopulmonary phase, there is elevated heart rate (

tachycardia Tachycardia, also called tachyarrhythmia, is a heart rate that exceeds the normal resting rate. In general, a resting heart rate over 100 beats per minute is accepted as tachycardia in adults. Heart rates above the resting rate may be normal ...

), irregular heartbeats (

arrhythmia Arrhythmias, also known as cardiac arrhythmias, are irregularities in the cardiac cycle, heartbeat, including when it is too fast or too slow. Essentially, this is anything but normal sinus rhythm. A resting heart rate that is too fast – ab ...

s), and

cardiogenic shock Cardiogenic shock is a medical emergency resulting from inadequate blood flow to the body's organs due to the dysfunction of the heart. Signs of inadequate blood flow include low urine production (<30 mL/hour), cool arms and legs, and decreased ...

. Pulmonary capillary leakage can lead to

acute respiratory distress syndrome Acute respiratory distress syndrome (ARDS) is a type of respiratory failure characterized by rapid onset of widespread inflammation in the lungs. Symptoms include shortness of breath (dyspnea), rapid breathing (tachypnea), and bluish skin co ...

, buildup of fluids in the lungs (

pulmonary edema Pulmonary edema (British English: oedema), also known as pulmonary congestion, is excessive fluid accumulation in the tissue or air spaces (usually alveoli) of the lungs. This leads to impaired gas exchange, most often leading to shortness ...

), hypotension, and buildup of fluid in the chest cavity (

pleural effusion A pleural effusion is accumulation of excessive fluid in the pleural space, the potential space that surrounds each lung. Under normal conditions, pleural fluid is secreted by the parietal pleural capillaries at a rate of 0.6 millilitre per kilog ...

). These symptoms can cause sudden death. After the cardiopulmonary phase is resolved, polyuria occurs while recovery takes months. While HFRS is associated with renal disease and HPS with cardiopulmonary disease, HFRS may sometimes include cardiopulmonary symptoms associated with HPS and HPS may sometimes include renal symptoms associated with HFRS.

Transmission

Hantaviruses that cause illness in humans are mainly transmitted by rodents. In rodents, hantaviruses usually cause an asymptomatic, persistent infection. Infected animals can spread the virus to uninfected animals through aerosols or droplets from their feces, urine, saliva, and blood, through consumption of contaminated food, from virus particles shed from skin or fur, via grooming, or through biting and scratching. Hantaviruses can also spread through the fecal-oral route and across the placenta during pregnancy from mother to child. They can survive for 10 days at room temperature, 15 days in a temperate environment, and more than 18 days at 4 degrees Celsius (39.2 degrees Fahrenheit), which aids in the transmission of the virus. Environmental conditions favorable to the reproduction and spread of rodents are known to increase disease transmission. Living in a rural environment, in unhygienic settings, and interacting with environments shared with hosts are the biggest risk factors for infection, especially among people who are hikers, farmers, and forestry workers, as well as those in mining, the military, and

zoology Zoology ( , ) is the scientific study of animals. Its studies include the anatomy, structure, embryology, Biological classification, classification, Ethology, habits, and distribution of all animals, both living and extinction, extinct, and ...

. Rodents can transmit hantaviruses to humans through

s or droplets from the excretions and through consumption of contaminated food. Rodent bites and scratches are also an important means of transmission to humans. The prevalence of hantavirus among rodent breeders and rodent pet owners is up to 80%. In one outbreak in North America in 2017,

infected 31 people through contact with pet rats. Andes virus has often been claimed by researchers to be the only hantavirus known to be spread from person to person, usually after coming into close contact with an infected person. It can also reportedly spread through human saliva, airborne droplets from coughing and sneezing, and to newborns through breast milk and the placenta. A 2021 systematic review, however, found human-to-human transmission of the Andes virus to not be strongly supported by evidence but nonetheless possible in limited circumstances, especially between close household contacts such as sexual partners. There is also suspicion that Puumala virus can spread from person to person through blood and platelet transfusions. Hantaviruses that cause HFRS can be transmitted through the bites of

mite Mites are small arachnids (eight-legged arthropods) of two large orders, the Acariformes and the Parasitiformes, which were historically grouped together in the subclass Acari. However, most recent genetic analyses do not recover the two as eac ...

s and

tick Ticks are parasitic arachnids of the order Ixodida. They are part of the mite superorder Parasitiformes. Adult ticks are approximately 3 to 5 mm in length depending on age, sex, and species, but can become larger when engorged. Ticks a ...

s. Research has also shown that pigs can be infected with Hantaan virus without severe symptoms and sows can transmit the virus to offspring through the placenta. Pig-to-human transmission may also be possible, as one swine breeder was infected with hantavirus with no contact with rodents or mites. Hantaan virus and Puumala virus have been detected in cattle, deer, and rabbits, and antibodies to Seoul virus have been detected in cats and dogs, but the role of these hosts for hantaviruses is unknown. Infection in these other animals can potentially facilitate the evolution of hantaviruses by genome reassortment. In addition to rodents, some hantaviruses are found in small insectivorous mammals, such as moles, shrews, and bats. Hantavirus antigen has also been detected in a variety of bird species, indicative of infection. Human built environments are important in hantavirus transmission. Deforestation and excess agriculture may destroy rodents' natural habitat. The expansion of agricultural land is associated with a decline in predator populations, which enables hantavirus host species to use farm monocultures as nesting and foraging sites. Agricultural sites built in close proximity to rodents' natural habitats can facilitate the proliferation of rodents as they may be attracted to animal feed. Sewers and stormwater drainage systems may be inhabited by rodents, especially in areas with poor solid waste management. Maritime trade and travel have also been implicated in the spread of hantaviruses. Research results are inconsistent on whether urban living increases or decreases hantavirus incidence.

Seroprevalence Seroprevalence is the number of persons in a population who test positive for a specific disease based on serology (blood serum) specimens, often presented as a percent of the total specimens tested or as a proportion per 100,000 persons tested. As ...

, which shows past infection to hantavirus, is consistently higher in occupations and areas that have greater exposure to rodents. Poor living conditions on battlefields, in military camps, and in refugee camps make soldiers and refugees at great risk of exposure as well.

Environment

Rodent species that carry hantaviruses inhabit a diverse range of habitats, including desert-like biomes, equatorial and tropical forests, swamps, savannas, fields, and salt marshes. The

seroprevalence Seroprevalence is the number of persons in a population who test positive for a specific disease based on serology (blood serum) specimens, often presented as a percent of the total specimens tested or as a proportion per 100,000 persons tested. As ...

of hantaviruses in their host species has been observed to range from 5.9% to 38% in the Americas, and 3% to about 19% worldwide, depending on testing method and location. In some places, such as South Korea, routine trapping of wild rodents is performed to surveil hantavirus circulation. High humidity can benefit rodent populations in warm climates, where it may positively impact plant growth and thus food availability. Increased forest coverage is associated with increased hantavirus incidence, particularly in Europe.

Climate change Present-day climate change includes both global warming—the ongoing increase in Global surface temperature, global average temperature—and its wider effects on Earth's climate system. Climate variability and change, Climate change in ...

and environmental degradation increase contact areas between rodent hosts and humans, which increases potential exposure to hantaviruses. An example of this was the 1993 Four Corners outbreak in the United States, which was immediately preceded by elevated rainfall from the 1992-1993

El Niño EL, El or el may refer to: Arts and entertainment Fictional entities * El, a character from the manga series ''Shugo Chara!'' by Peach-Pit * Eleven (''Stranger Things'') (El), a fictional character in the TV series ''Stranger Things'' * El, fami ...

warming period. This caused a substantial growth in the food supply for rodents, which led to rapid growth in their population and facilitated greater spread of the hantavirus that caused that outbreak. Rainfall is consistently associated with hantavirus incidence in various patterns. Heavy rainfall is a risk factor for outbreaks in the following months, but may negatively affect incidence by flooding rodent burrows and nests. In places that have wet and dry seasons, infections are more common in the wet season than in the dry season. Low rainfall and drought are associated with decreased incidence since such conditions result in a smaller rodent population, but displacement of rodent populations via drought or flood can lead to an increase in rodent-human interactions and infections. In Europe, however, no association between rainfall and disease incidence has been found. Temperature has varying effects on hantavirus transmission. Higher temperatures create unfavorable environments for virus survival and decreases activity levels of

Neotropic The Neotropical realm is one of the eight biogeographic realms constituting Earth's land surface. Physically, it includes the tropical terrestrial ecoregions of the Americas and the entire South American temperate zone. Definition In biogeogra ...

rodents, but it can cause rodents to seek shelter from heat in human settings and is beneficial for aerosol production. Lower temperature can prolong virus survival outside a host. Higher average winter temperature is associated with reduced survival of bank voles, the natural reservoir of Puumala virus, but increased survival of striped field mice in China, the natural reservoirs of Hantaan virus. Extreme temperatures, whether hot or cold, are associated with lower disease incidence.

Genome and structure

The genome of hantaviruses is segmented into three parts: the large (L), medium (M), and small (S) segments. Each part is a single-stranded negative-sense RNA strand and consists of 10,000–15,000 nucleotides in total. The segments form into circles via non-covalent bonding of the ends of the genome. The L segment is about 6.6 kilobases (kb) in length and encodes a viral

(RdRp), which mediates

transcription Transcription refers to the process of converting sounds (voice, music etc.) into letters or musical notes, or producing a copy of something in another medium, including: Genetics * Transcription (biology), the copying of DNA into RNA, often th ...

and replication of viral RNA. The M segment, about 3.7 kb in length, encodes a glycoprotein precursor that is co-translated and cleaved into Gn and Gc. Gn and Gc bind to cell receptors, regulate immune responses, and induce protective antibodies. The S segment is around 2.1 kb in length and encodes the

nucleocapsid A capsid is the protein shell of a virus, enclosing its genetic material. It consists of several oligomeric (repeating) structural subunits made of protein called protomers. The observable 3-dimensional morphological subunits, which may or ma ...

protein N, which binds to and protects viral RNA. An

open reading frame In molecular biology, reading frames are defined as spans of DNA sequence between the start and stop codons. Usually, this is considered within a studied region of a prokaryotic DNA sequence, where only one of the six possible reading frames ...

in the N gene on the S segment of some orthohantaviruses also encodes the non-structural protein NS that inhibits

interferon Interferons (IFNs, ) are a group of signaling proteins made and released by host cells in response to the presence of several viruses. In a typical scenario, a virus-infected cell will release interferons causing nearby cells to heighten ...

production in host cells. The untranslated regions at the ends of the genome are highly conserved and participate in the replication and transcription of the genome. Individual hantavirus particles (virions) are usually spherical, but may be oval, pleomorphic, or tubular. The diameter of the virion is 70–350 nanometers (nm). The outer part of the virion is a lipid envelope that is about 5 nm thick. Embedded in the envelope are the surface spike glycoproteins Gn and Gc, which are arranged in a lattice pattern. Each surface spike is composed of a tetramer of Gn and Gc (four units each) that has four-fold rotational symmetry, and extends about 10 nm out from the envelope. Gn forms the stalk of the spike and Gc the head. Inside the envelope are helical nucleocapsids made of many copies of the nucleocapsid protein N, which are attached to the virus's genome to form ribonucleoprotein (RNP) complexes. Each RNP complex has a copy of RdRp attached to it. Hantaviruses do not encode matrix proteins to assist with structuring the virion, so how surface proteins organize into a sphere with a symmetrical lattice is not yet known.

Life cycle

Sin Nombre hanta virus TEM PHIL 1136 lores

Vascular

endothelial cell The endothelium (: endothelia) is a single layer of squamous endothelial cells that line the interior surface of blood vessels and lymphatic vessels. The endothelium forms an interface between circulating blood or lymph in the lumen and th ...

s and

macrophage Macrophages (; abbreviated MPhi, φ, MΦ or MP) are a type of white blood cell of the innate immune system that engulf and digest pathogens, such as cancer cells, microbes, cellular debris and foreign substances, which do not have proteins that ...

s are the primary cells infected by hantaviruses.

Podocyte Podocytes are cells in Bowman's capsule in the kidneys that wrap around capillaries of the glomerulus. Podocytes make up the epithelial lining of Bowman's capsule, the third layer through which filtration of blood takes place. Bowman's capsule ...

s, tubular cells,

dendritic cell A dendritic cell (DC) is an antigen-presenting cell (also known as an ''accessory cell'') of the mammalian immune system. A DC's main function is to process antigen material and present it on the cell surface to the T cells of the immune system ...

s, and

lymphocyte A lymphocyte is a type of white blood cell (leukocyte) in the immune system of most vertebrates. Lymphocytes include T cells (for cell-mediated and cytotoxic adaptive immunity), B cells (for humoral, antibody-driven adaptive immunity), an ...

s can also be infected. Attachment and entry into the host cell is mediated by the binding of the viral glycoprotein spikes to host cell receptors, particularly β3 integrins. Decay acceleration factors, complement receptors, and, for New World hantaviruses,

protocadherin Protocadherins (Pcdhs) are the largest mammalian subgroup of the cadherin superfamily of homophilic cell-adhesion proteins. They were discovered by Shintaro Suzuki's group, when they used PCR to find new members of the cadherin family. The PCR fra ...

-1 have also been proposed to be involved in attachment. After attachment, hantaviruses rely on several ways to enter a cell, including micropinocytosis,

clathrin Clathrin is a protein that plays a role in the formation of coated vesicles. Clathrin was first isolated by Barbara Pearse in 1976. It forms a triskelion shape composed of three clathrin heavy chains and three light chains. When the triskel ...

-independent receptor-mediated

endocytosis Endocytosis is a cellular process in which Chemical substance, substances are brought into the cell. The material to be internalized is surrounded by an area of cell membrane, which then buds off inside the cell to form a Vesicle (biology and chem ...

and

cholesterol Cholesterol is the principal sterol of all higher animals, distributed in body Tissue (biology), tissues, especially the brain and spinal cord, and in Animal fat, animal fats and oils. Cholesterol is biosynthesis, biosynthesized by all anima ...

- or

caveolae In biology, caveolae (Latin for "little caves"; singular, caveola), which are a special type of lipid raft, are small (50–100 nanometer) invaginations of the plasma membrane in the cells of many vertebrates. They are the most abundant surface fe ...

-dependent endocytosis. Old World hantaviruses use clathrin-dependent endocytosis while New World hantaviruses use clathrin-independent endocytosis. After entering a cell, virions form vesicles that are transported to early

s, then late endosomes and lysosomal compartments. A decrease in pH then causes the viral envelope to fuse with the endosome or

. This fusion releases viral ribonucleoprotein complexes into the cell cytoplasm, which initiates transcription and replication by RdRp. RdRp transcribes viral -ssRNA into complementary positive-sense strands, then snatches 5′ ("five prime") ends of host

(mRNA) to prepare mRNA for translation by host ribosomes to produce viral proteins. Complementary RNA strands are also used to produce copies of the genome, which are encapsulated by N proteins to form RNPs. During virion assembly, the glycoprotein precursor is cleaved in the endoplasmic reticulum into the Gn and Gc glycoproteins by host cell

signal peptidase Signal peptidases are enzymes that convert secretory and some membrane proteins to their mature or pro forms by cleaving their signal peptides from their N-termini. Signal peptidases were initially observed in endoplasmic reticulum (ER)-deri ...

s. Gn and Gc are modified by N-glycan chains, which stabilize the spike structure and assist in assembly in the Golgi apparatus for Old World hantaviruses or at the cell membrane for New World hantaviruses. Old World hantaviruses obtain their viral envelope from the

and are then transported to the cell membrane in vesicles to leave the cell via

. On the other hand, New World hantavirus RNPs are transported to the cell membrane, where they bud from the surface of the cell to obtain their envelope and leave the cell.

Evolution

The most common form of evolution for hantaviruses is

mutation In biology, a mutation is an alteration in the nucleic acid sequence of the genome of an organism, virus, or extrachromosomal DNA. Viral genomes contain either DNA or RNA. Mutations result from errors during DNA or viral replication, ...

s through single nucleotide substitutions, insertions, and deletions. Hantaviruses are usually restricted to individual

species and evolve alongside their hosts, but this one-species-one-hantavirus relationship is not true for all hantaviruses. The exact evolutionary history of hantaviruses is likely obscured by many instances of genome reassortment, host spillover, and host-switching. Within species, geography has affected the evolution of hantaviruses. For example, Hantaan virus and Seoul virus have both formed multiple lineages corresponding to their geographic distribution. Because hantaviruses have segmented genomes, they are capable of

genetic recombination Genetic recombination (also known as genetic reshuffling) is the exchange of genetic material between different organisms which leads to production of offspring with combinations of traits that differ from those found in either parent. In eukaryot ...

and

reassortment Reassortment is the mixing of the genetic material of a species into new combinations in different individuals. The product of reassortment is called a reassortant. It is particularly used when two similar viruses that are infecting the same cell ...

in which segments from different viruses can combine to form new viruses. This occurs often in nature and facilitates the adaptation of hantaviruses to multiple hosts and ecosystems. Recombination in OWHVs of the S and M segments is usually observed amongst viruses within species, but can occur between species. Reassortment in NWHVs of the S and M segments has been observed in rodents. Among Puumala viruses isolated from rodents in 2005-2009, 19.1% of them were identified as reassortments. Diploid progeny are also possible, in which virions may possess two of the same segment from two parent viruses.

Classification

''Orthohantavirus'' belongs to the family ''

'', which contains all hantaviruses. The genus has 37 species, listed hereafter with the exemplar virus of the species. In general, species bear the name of the exemplar virus with the suffix -''ense''. *''Orthohantavirus andesense'', Andes virus *''Orthohantavirus artybashense'',

Artybash virus Artybash (; , ''Artıbaş'') is a rural locality (a settlement) in Artybashskoye Rural Settlement of Turochaksky District, the Altai Republic, Russia. The population was 602 as of 2016. There are 46 streets. Geography Artybash is located at the ...

*''Orthohantavirus asamaense'',

Asama virus Asama virus (ASAV) is a single-stranded, enveloped, segmented negative-sense RNA hantavirus.Arai S, Ohdachi SD, Asakawa M, Kang HJ, Mocz G, Arikawa J, Okabe N, Yanagihara R.Molecular phylogeny of a newfound hantavirus in the Japanese shrew mole ( ...

*''Orthohantavirus asikkalaense'',

Asikkala virus Asikkala virus (ASIV) is an Old World hantavirus isolated from ''Sorex araneus'' (pygmy shrew) in Germany and the Czech Republic The Czech Republic, also known as Czechia, and historically known as Bohemia, is a landlocked country in Cen ...

*''Orthohantavirus bayoui'', Bayou virus *''Orthohantavirus boweense'', Bowé virus *''Orthohantavirus brugesense'',

Bruges virus Bruges ( , ; ; ) is the capital and largest city of the province of West Flanders, in the Flemish Region of Belgium. It is in the northwest of the country, and is the sixth most populous city in the country. The area of the whole city amoun ...

*''Orthohantavirus caobangense'', Cao Bằng virus *''Orthohantavirus carrizalense'',

Carrizal virus __NOTOC__ Carrizal may refer to: Places Europe * Cerecinos del Carrizal, municipality of Zamora, Spain North America * Carrizal, Chihuahua, Mexico, a small town between Ciudad Chihuahua and Juarez ** Battle of Carrizal The Battle of Carrizal ...

*''Orthohantavirus chocloense'',

Choclo virus Choclo virus (CHOV) is a single-stranded, negative-sense RNA zoonotic New World hantavirus. It was first isolated in 1999 in western Panama. The finding marked the first time Hantavirus pulmonary syndrome (HPS) was found in Central America. Dur ...

*''Orthohantavirus dabieshanense'', Dàbiéshān virus *''Orthohantavirus delgaditoense'', Caño Delgadito virus *''Orthohantavirus dobravaense'',

The exemplar virus of ''Orthohantavirus dobravaense'' is Dobrava virus, a genotype of Dobrava-Belgrade virus. In scientific papers, "Dobrava-Belgrade virus" is essentially used as a synonym for ''Orthohantavirus dobravaense''. *''Orthohantavirus fugongense'', Fúgòng virus *''Orthohantavirus hantanense'',

*''Orthohantavirus jejuense'', Jeju virus *''Orthohantavirus kenkemeense'',

Kenkeme virus The Kenkeme (; , ''Keŋkeme'') is a river in Yakutia (Sakha Republic), Russia. It is a tributary of the Lena (river), Lena with a length of — together with the Yagas-Yyaabyt at its head— and a drainage basin area of . The Mirny, Sakha Repub ...

*''Orthohantavirus khabarovskense'', Khabarovsk virus *''Orthohantavirus lankaense'',

Lanka virus Lanka (; ) is the name given in Hindu epics to the island fortress capital of the legendary Rakshasa king Ravana in the epics of the ''Ramayana'' and the ''Mahabharata''. The fortress was situated on a plateau between three mountain peaks kn ...

*''Orthohantavirus luxiense'', Lúxī virus *''Orthohantavirus mamorense'', Rio Mamoré virus *''Orthohantavirus maporalense'', Maporal virus *''Orthohantavirus montanoense'', Montaño virus *''Orthohantavirus nigrorivense'', Black Creek Canal virus *''Orthohantavirus ozarkense'',

Ozark virus The Ozarks, also known as the Ozark Mountains, Ozark Highlands or Ozark Plateau, is a physiographic region in the U.S. states of Missouri, Arkansas, and Oklahoma, as well as a small area in the southeastern corner of Kansas. The Ozarks cover ...

*''Orthohantavirus prospectense'', Prospect Hill virus *''Orthohantavirus puumalaense'',

*''Orthohantavirus rockportense'',

Rockport virus Rockport virus (RKPV) is a single-stranded, enveloped, negative-sense RNA orthohantavirus. Natural reservoir Rockport virus was first isolated in archival tissues of four Eastern moles found in and around Rockport, Texas. Virology Phylogene ...

*''Orthohantavirus sagercreekense''

Sager Creek virus Sager is a surname. Notable people with the surname include: * Bobby Sager, entrepreneur turned philanthropist, and inspiration for ''The Philanthropist'' television series * Carole Bayer Sager (born 1944), American lyricist, songwriter, and singer ...

*''Orthohantavirus sangassouense'', Sangassou virus *''Orthohantavirus seoulense'',

*''Orthohantavirus sinnombreense'',

*''Orthohantavirus tatenalense'', Tatenale virus *''Orthohantavirus thailandense'', which contains

Anjozorobe virus Anjozorobe is a large town in the Analamanga Region, Madagascar, approximately 90 kilometers north-east of the capital Antananarivo. It has a population of 24,117 inhabitants in 2018. Anjozorobe-Angavo Reserve is one of the last high plateau fo ...

and

Thailand virus Thailand virus (THAIV) is a single-stranded, enveloped, negative-sense RNA orthohantavirus. Natural reservoir THAIV was first isolated from rodents in two Thai provinces, Nakhon Pathom Province and Nakhon Ratchasima Province, in 1994. The gre ...

''Orthohantavirus thailandense'' bears the name of Thailand virus but its exemplar virus is Anjozorobe virus. *''Orthohantavirus tigrayense'',

Tigray virus The Tigray Region (or simply Tigray; officially the Tigray National Regional State) is the northernmost regional state in Ethiopia. The Tigray Region is the homeland of the Tigrayan, Irob and Kunama people. Its capital and largest city is Me ...

*''Orthohantavirus tulaense'', Tula virus *''Orthohantavirus wufangense'', Wùfeng Chodsigoa smithii orthohantavirus 1 Many other hantaviruses are unclassified, though some may be isolates of other viruses: * Academ virus *

Adler virus Adler may refer to: Places *Adler, Alabama, an unincorporated community in Perry County *Adler Planetarium, Chicago, Illinois, USA *Adler Township, Nelson County, North Dakota, USA *Adler University, formerly Adler School of Professional Psycholo ...

Alto Paraguay virus The musical term alto, meaning "high" in Italian (Latin: ''altus''), historically refers to the contrapuntal part higher than the tenor and its associated vocal range. In four-part voice leading alto is the second-highest part, sung in choruses ...

Amga virus Amga or AMGA may refer to: *Amga (rural locality), a rural locality (a ''selo'') in the Amginsky District of the Sakha Republic, Russia *Amga (river), a river in the Sakha Republic, Russia * American Medical Group Association (AMGA) *American Mounta ...

/ Seewis virus *

Anajatuba virus Anajatuba is a municipality in the state of Maranhão in the Northeast region of Brazil. The municipality contains a small part of the Baixada Maranhense Environmental Protection Area, a sustainable use conservation unit created in 1991 that has ...

Ash River virus Ash is the solid remnants of fires. Specifically, ''ash'' refers to all non-aqueous, non-gaseous residues that remain after something burns. In analytical chemistry, to analyse the mineral and metal content of chemical samples, ash is the n ...

Asturias virus Asturias (; ; ) officially the Principality of Asturias, is an autonomous communities of Spain, autonomous community in northwest Spain. It is coextensive with the provinces of Spain, province of Asturias and contains some of the territory t ...

Azagny virus Azagny virus (AZGV) is an ''Orthohantavirus ''Orthohantavirus'' is a genus of viruses that includes all hantaviruses (family ''Hantaviridae'') that cause disease in humans. Orthohantaviruses, hereafter referred to as hantaviruses, are naturall ...

Belgrade virus Belgrade is the Capital city, capital and List of cities in Serbia, largest city of Serbia. It is located at the confluence of the Sava and Danube rivers and at the crossroads of the Pannonian Basin, Pannonian Plain and the Balkan Peninsula. T ...

Biya river virus Biya may refer to: Places * Biya (river), a tributary of the Ob in Siberia, Russia; * Al Hoceima or Biya, a Moroccan port on the Mediterranean Sea; People * Paul Biya (born 1933), president of Cameroon 1982–present ** Jeanne-Irène Biya (1935 ...

* Bloodland Lake virus *

Blue River virus Blue is one of the three primary colours in the RYB colour model (traditional colour theory), as well as in the RGB (additive) colour model. It lies between violet and cyan on the spectrum of visible light. The term ''blue'' generally descr ...

Boginia virus Boginia is a village in the administrative district of Gmina Nowosolna, within Łódź East County, Łódź Voivodeship, in central Poland. It lies approximately north-east of the regional capital Łódź Łódź is a city in central Poland ...

Calabazo virus Calabazo virus is an enveloped, single-stranded, negative-sense RNA hantavirus species of the order ''Bunyavirales''. It is a novel New World microtine rodent-borne hantavirus discovered in Central America on the Azuero Peninsula of Panama in ea ...

* Camp Ripley virus *

Castelo dos Sonhos virus Castelo may refer to: Places Brazil * Castelo, Espírito Santo, a municipality in the State of Espírito Santo * Castelo (Rio de Janeiro), a neighbourhood in the city of Rio de Janeiro Portugal * Castelo (Lisbon), a civil parish in the municipalit ...

* CGRn9415 virus *

Dode virus Dode may refer to: Geography *Dode, Jalandhar, India *Dode, Kent, United Kingdom People * Dode, Abbess of Saint Pierre de Reims * Dode Criss * Dode Paskert *Guillaume Dode de la Brunerie Guillaume Dode de la Brunerie, Viscount of Martignac (Apr ...

* El Moro Canyon virus *

Fox Creek virus Foxes are small-to-medium-sized omnivorous mammals belonging to several genera of the family Canidae. They have a flattened skull; upright, triangular ears; a pointed, slightly upturned snout; and a long, bushy tail ("brush"). Twelve species ...

Fusong virus Fusong County () is a county in southern Jilin province, China. It is under the administration of Baishan City, with a population of 310,000 residing in an area of . The county contains the Changbaishan Airport, which opened on 3 August 2008, and ...

* Gōu virus * hantavirus sp. strain Tamarin/BRA/SM22/2014 *

HoJo virus Hojo or Hōjō may refer to: Hojo or HoJo: *Howard Johnson's, a U.S. chain of restaurants and hotels *A nickname for a number of people named Howard Johnson *A nickname for Howard Jones (British musician), an synthpop singer, musician and songwr ...

* Iamonia virus * Isla Vista virus *

Jemez Springs virus Jemez or Jémez may refer to *Jemez Pueblo, New Mexico, a census-designated place in the United States **Jemez Springs, New Mexico, a village **Jemez Mountains **Jemez Mountains salamander (''Plethodon neomexicanus'') **Jemez Mountains Electric C ...

Jerboa hantavirus Jerboas () are the members of the family Dipodidae. They are hopping desert rodents found throughout North Africa and Asia. They tend to live in hot deserts. When chased, jerboas can run at up to . Some species are preyed on by little owls (''A ...

Jurong virus Jurong () is a major Region, geographical region located at the south-westernmost point of the West Region, Singapore, West Region of Singapore. Although mostly vaguely defined, the region's extent roughly covers the Planning Areas of Singapore, ...

Kielder hantavirus Kielder is a small, remote village in western Northumberland, England. Located at the head of Kielder Water and in the north west of Kielder Forest, the village is within of the Scottish border. Etymology ''Kielder'' is thought to take its na ...

* Laguna Negra virus *

Landiras virus Landiras (; ) is a Communes of France, commune in the Gironde Departments of France, department in the Nouvelle-Aquitaine Regions of France, region in Southwestern France. Population Notable people Jules Koundé (born 1998), professional footb ...

Leakey virus Leakey may refer to: *Leakey, Texas, U.S., a city *Leakey (crater), a lunar impact crater *Leakey (surname) *7958 Leakey, an asteroid *Leakey Independent School District The Leakey Independent School District is a public school district based i ...

* Lechiguanas virus * Liánghé virus *

Lohja virus Lohja (; ) is a town in Finland, located in the southern interior of the country. Lohja is situated in the western part of the Uusimaa region. The population of Lohja is approximately . It is the most populous municipality in Finland. Lohja is ...

Malacky virus Malacky (German: ''Malatzka'', Hungarian: ''Malacka'') is a town and municipality in western Slovakia around north of Slovakia’s capital, Bratislava. From the second half of the 10th century until 1918, it was part of the Kingdom of Hungary. ...

* Muleshoe virus * Necocli virus * Orán virus *

Oxbow virus Oxbow virus (OXBV) is a single-stranded, enveloped, negative-sense RNA orthohantavirus.Kang HJ, Bennett SN, Dizney L, Sumibcay L, Arai S, Ruedas LA, Song JW, Yanagihara R.Host switch during evolution of a genetically distinct hantavirus in the A ...

* Playa de Oro virus *

Powell Butte virus Powell may refer to: People * Powell (surname) * Powell (given name) * Powell baronets, several baronetcies * Colonel Powell (disambiguation), several military officers * General Powell (disambiguation), several military leaders * Governor Powell ...

* Prairie vole virus * Qiān Hú Shān virus/ Qiāndǎo Lake virus *

Rio Mearim virus Rio or Río is the Portuguese and Spanish word for "river". The word also exists in Italian, but is largely obsolete and used in a poetical or literary context to mean "stream". Rio, RIO or Río may also refer to: Places United States * Rio, Flo ...

* Río Segundo virus *

Sapporo rat virus is a designated city in Hokkaido, Japan. Located in the southwest of Hokkaido, it lies within the alluvial fan of the Toyohira River, a tributary of the Ishikari River. Sapporo is the capital of Hokkaido Prefecture and Ishikari Subprefecture. ...

Sarufutsu virus file:Sarufutsu village office.JPG, 290px, Sarufutsu Town Hall file:Indigirka Issue Distress Memorial, Hokkaido, Japan.jpg, 290px, Indigirka Memorial is a List of villages in Japan, village located in Sōya Subprefecture, Hokkaido, Japan. , the ...

Serang virus Serang virus (SERV) is a single-stranded, negative-sense, enveloped, novel RNA orthohantavirus. Natural reservoir SERV was first isolated from the Asian house rat (''R.Tanezumi'') in Serang, Indonesia in 2000. Virology Phylogenetic analysis ...

* Shěnyáng virus * Taimyr virus * Tanganya virus *

Tualatin River virus Tualatin may refer to: Places Oregon, United States * Tualatin, Oregon, a city in the Tualatin Valley * Tualatin Academy, a building and former school in Forest Grove * Tuality Community Hospital, a hospital in Hillsboro * Tualatin Mountains, a se ...

Uurainen virus Uurainen (, also ) is a municipality of Finland. It is part of the Central Finland region. The municipality has a population of () and covers an area of of which , or roughly 6.5%, is water. The population density is . Neighbouring municipali ...

Vladivostok virus Vladivostok ( ; , ) is the largest city and the administrative center of Primorsky Krai and the capital of the Far Eastern Federal District of Russia. It is located around the Golden Horn Bay on the Sea of Japan, covering an area of , with a p ...

Yakeshi virus Yakeshi (; zh, c=牙克石) is a county-level city of Hulunbuir, Inner Mongolia, China. Yakeshi has a population of 391,627 and an area of . It is situated next to the Hailar River east of Hailar District, the seat of Hulunbuir, and on the nor ...

* Yuánjiāng virus

History

Hantavirus hemorrhagic disease was likely first described in the ''

Huangdi Neijing ' (), literally the ''Inner Canon of the Yellow Emperor'' or ''Esoteric Scripture of the Yellow Emperor'', is an ancient Chinese medical text or group of texts that has been treated as a fundamental doctrinal source for Chinese medicine for mo ...

'', an ancient Chinese medical text, in Imperial China during the

Warring States Period The Warring States period in history of China, Chinese history (221 BC) comprises the final two and a half centuries of the Zhou dynasty (256 BC), which were characterized by frequent warfare, bureaucratic and military reforms, and ...

of 475–221 BCE. Hantaviruses have been suggested as a cause of "trench nephritis" in soldiers during the

US Civil War The American Civil War (April 12, 1861May 26, 1865; also known by other names) was a civil war in the United States between the Union ("the North") and the Confederacy ("the South"), which was formed in 1861 by states that had seceded f ...

and in British soldiers in Flanders, Belgium during the

First World War World War I or the First World War (28 July 1914 – 11 November 1918), also known as the Great War, was a World war, global conflict between two coalitions: the Allies of World War I, Allies (or Entente) and the Central Powers. Fighting to ...

. The disease was also mentioned in East Asia, where it was probably endemic, and was first described scientifically in

Vladivostok Vladivostok ( ; , ) is the largest city and the administrative center of Primorsky Krai and the capital of the Far Eastern Federal District of Russia. It is located around the Zolotoy Rog, Golden Horn Bay on the Sea of Japan, covering an area o ...

in 1913–1914. During the

Second World War World War II or the Second World War (1 September 1939 – 2 September 1945) was a World war, global conflict between two coalitions: the Allies of World War II, Allies and the Axis powers. World War II by country, Nearly all of the wo ...

in 1942, an outbreak of disease with symptoms characteristic of hantavirus infection occurred in

Salla Salla, known as Kuolajärvi until 1936, is a municipality of Finland, located in Lapland. The municipality has a population of () and covers an area of of which is water. The population density is . The nearby settlement of Sallatunturi is ...

, Eastern Lapland, Finland among German and Finnish soldiers. This outbreak was later reported in 1980 to be caused by a virus transmitted by bank voles and was named Puumala virus. Also during the war, around 10,000 Japanese soldiers stationed in

Manchuria Manchuria is a historical region in northeast Asia encompassing the entirety of present-day northeast China and parts of the modern-day Russian Far East south of the Uda (Khabarovsk Krai), Uda River and the Tukuringra-Dzhagdy Ranges. The exact ...

developed HFRS. Around 3,200 cases of HFRS occurred among United Nations soldiers stationed near the Hantan river during the

, where it was first identified in 1951 and named "Korean hemorrhagic fever" and "epidemic hemorrhagic fever". After the war, in 1976 in South Korea,

Ho Wang Lee Ho Wang Lee (26 October 1928 – 5 July 2022) was a South Korean physician, epidemiologist, and virologist. He was the first person in the history of medicine to be the one chiefly responsible for all 3 of the following steps: (1) discovery of ...

(Korean: 이호황) tested striped field mice and showed that antigens from their lungs were reactive to antibodies in sera from war survivors. In 1978, the virus was isolated for the first time and named Hantaan virus after the river. Retrospective analysis showed that Hantaan virus was responsible for the war outbreak. Other hantaviruses that caused by HFRS were then discovered throughout Eurasia. The disease had a variety of names, so in 1982, the World Health Organization officially named it "hemorrhagic fever with renal syndrome". In 1985, this group of viruses were named "hantaviruses" after Hantaan virus, and in 1987, the genus ''Hantavirus'' was established to accommodate them in the then-family ''Bunyaviridae''. In 1993, an outbreak of highly lethal acute respiratory distress syndrome occurred in the

Four Corners Four Corners is a region of the Southwestern United States consisting of the southwestern corner of Colorado, southeastern corner of Utah, northeastern corner of Arizona, and northwestern corner of New Mexico. Most of the Four Corners regio ...

region of the United States. This outbreak was determined to be caused by a hantavirus, now named

, and represented the first confirmed instance of pathogenic hantaviruses in the Americas as well as the discovery of a new type of disease caused by hantaviruses. The new disease was named "hantavirus pulmonary syndrome". During subsequent years, numerous other hantaviruses were discovered in the Americas, including Andes virus, which has been claimed to be transmissible from person to person. HFRS, however, remains much more common than HPS—more than 100,000 cases of HFRS occur each year, compared to only a few hundred cases of HPS annually. Over time, hundreds of bunyaviruses were discovered but could not be accommodated within the genera of the ''Bunyaviridae'' family. To address this, in 2017 bunyaviruses were elevated to the rank of order, ''Bunyavirales'', and hantaviruses, along with the other bunyavirus genera, were elevated to the rank of family. Hantaviruses, also called hantavirids, now also refer to members of the family ''Hantaviridae''. The prior genus of ''Hantavirus'' was renamed ''Orthohantavirus'' to distinguish them from members of the family, and the genus's members are often called orthohantaviruses. In 2019, additional genera, subfamilies, and families were created to classify non-rodent hantaviruses, and in 2023

binomial nomenclature In taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin grammatical forms, altho ...

was adopted for hantaviruses.

Notes

References

External links

CDC's Hantavirus Fact Sheet (PDF)

Virus Pathogen Database and Analysis Resource (ViPR): Hantaviridae
{{Authority control Hantaviridae Hemorrhagic fevers Rodent-carried diseases Viral diseases Virus genera