Bacillus anthracis: Infectious substances pathogen safety data sheet

Section I – Infectious agent


Bacillus anthracis

Agent type









Synonym or cross-reference

Anthrax, Yamal disease and woolsorters' diseaseFootnote 1Footnote 2Footnote 3.


Brief description

B. anthracis is a member of the Bacillus cereus groupFootnote 1. B. anthracis are Gram-positive, rod-shaped (1.0 to 1.5 μm by 3 to 8 μm) cells that occur singly or in chainsFootnote 1. B. anthracis is a non-motile, facultative anaerobe that forms metabolically dormant, environmentally resistant endospores under adverse conditionsFootnote 1. Genomic DNA is circular (5.3 to 5.5 Mbp)Footnote 4. Clades A – E have been definedFootnote 5. Some strains produce a capsule and/or exotoxinsFootnote 1.


Upon entering a host, B. anthracis spores are internalized by phagocytes, triggering germinationFootnote 6. The germination process occurs rapidly (2 to 10 min)Footnote 7. Vegetative cells divide by binary fission and produce virulence factors that aid in the establishment of infection and local tissue damageFootnote 6. Virulence factors are encoded on two plasmids, pXO1 and pXO2; loss of either plasmid results in a considerable reduction in virulenceFootnote 7. Components of anthrax toxins (lethal toxin and edema toxin) are encoded by pXO1 and are largely responsible for the morbidity and mortality associated with anthrax diseaseFootnote 6Footnote 7, while pXO2 encodes capsule components.

Section II – Hazard identification

Pathogenicity and toxicity

Bacillus anthracis is the causative agent of anthrax, a toxin-mediated disease affecting humans, livestock, and wildlife. Disease in humans is characterized based on the portal of entry. Cutaneous, ingestion related (oropharyngeal/gastrointestinal), inhalational, and injection-associated forms of anthrax have been describedFootnote 7Footnote 8.

Cutaneous form of Bacillus anthracis accounts for about 95% of human anthrax casesFootnote 7. Disease can be mild to severe. Symptoms include headache, malaise, fever, and bullous lesions on face, neck, hands, and/or arms with localized erythema. Lesions develop into black eschars after 5 to 7 daysFootnote 8. If not treated, dissemination via the bloodstream occurs in 5-20% of casesFootnote 9. Complications include extensive edema and toxaemic shockFootnote 10. With treatment, mortality is less than 1%Footnote 8.

There are two forms of ingestion-associated anthrax. Oropharyngeal anthrax is characterized by lesions affecting the tongue, buccal cavity, tonsils, or posterior pharyngeal wall. Symptoms include sore throat, fever, swelling of neck and anterior chest wallFootnote 7Footnote 8. Gastrointestinal anthrax is characterized by occurrence of ulcers between the jejunum and the cecum and can be mild to severeFootnote 8. Symptoms include abdominal pain, anorexia, nausea, vomiting, and feverFootnote 8. In severe cases hematemesis, fluid collection in the abdomen, bloody diarrhoea, and toxic shock may occurFootnote 7Footnote 9. Mortality is estimated to be 4-50%Footnote 9.

Inhalational anthrax has a biphasic clinical course. Prodromal symptoms include headache, fever, malaise, myalgia, and cough for approximately 4 daysFootnote 8. This is followed by a fulminant phase characterized by high fever, shortness of breath, fluid collection in the chest, and septic shockFootnote 9. Development of meningoencephalitis occurs in approximately 38% of casesFootnote 11. Mortality is approximately 45% when treated early, and 97% if patients progress to fulminant phaseFootnote 11.

Intravenous anthrax is associated with tissue edema at the injection site and soft tissue infectionFootnote 8. Papules, vesicles, and eschars are usually absentFootnote 8. Severe cases develop hemorrhagic meningitis and have a high incidence of septic shockFootnote 8. Mortality is approximately 34%Footnote 8Footnote 12.

In herbivores, sudden death is often the first sign of anthraxFootnote 13. Death may be preceded by fever, dyspnea, edematous swelling in neck, or abdomenFootnote 13. Mortality rates in herbivores are highly variable, can be up to 90% for some unvaccinated species under certain environmental conditionsFootnote 7Footnote 14Footnote 15. Pigs, dogs, and carnivorous species are more resistant but can still die from infectionFootnote 7Footnote 13. Anthrax in birds is relatively rare; disease course is similar to that in herbivorous mammalsFootnote 13.


B. anthracis inhabits every continent except AntarcticaFootnote 7Footnote 16. Some regions have been characterized as hyperendemic, while other endemic regions can go years or decades without an anthrax outbreakFootnote 16Footnote 17Footnote 18Footnote 19. At least 2,000 cases of human anthrax occur annually worldwideFootnote 20. Approximately 95% of human anthrax cases are the cutaneous form acquired during the handling or processing of infected meat or animal products (e.g., hides, wool, bones)Footnote 7Footnote 9. Sporadic anthrax outbreaks associated with consumption of insufficiently cooked infected meat account for less than 5% of human casesFootnote 7Footnote 8. Several intravenous drug use-associated anthrax cases have been reported in which drugs were contaminated with B. anthracis sporesFootnote 12. A recent incident involving the malicious use of B. anthracis spores in 2001 in the United States where spores were mailed to several individuals resulted in 22 anthrax cases and 5 fatalitiesFootnote 7.

Introduction of livestock vaccines in the 1930s and other control measures drastically reduced anthrax incidenceFootnote 13. Anthrax outbreaks affecting livestock, particularly cattle, and herbivorous wildlife populations (e.g., bison, reindeer, hippopotami) are still reported in endemic and hyperendemic regionsFootnote 13Footnote 15Footnote 16Footnote 21. Seasonality of outbreaks varies according to affected species and ecosystemFootnote 16Footnote 17Footnote 22. Some sporadic outbreaks have been attributed to soil disturbance such as digging, dredging, or erosion-related exposure of sites previously used for carcass disposal or processing of animal products (e.g., tanneries)Footnote 7Footnote 13.

Unknown for humans. Seroprevalence studies indicate that herbivores are more severely affected than carnivorous wildlife, with the exceptions of mink and cheetahsFootnote 13Footnote 17. Some herbivore species (e.g., white-tailed deer, bison, and reindeer) are particularly prone to B. anthracis infectionFootnote 3Footnote 17. Animal stress has been suggested to play an important role in disease susceptibilityFootnote 3Footnote 22.

Host range

Natural host(s)

Humans, cattle, sheep, goats, swine, poultryFootnote 13. Various herbivorous mammals including bovines (e.g., buffaloFootnote 23, bisonFootnote 24), equines (e.g., horses, zebraFootnote 13), hippopotamiFootnote 15Footnote 23, elephantsFootnote 25, rhinosFootnote 26, antelopesFootnote 15Footnote 22Footnote 26, elks, and deersFootnote 27Footnote 28 are susceptible hosts. Other hosts include felinesFootnote 13Footnote 29, caninesFootnote 13Footnote 17, ostrichesFootnote 13, and snakesFootnote 30.

Other host(s)

Non-human primates, rabbits, guinea pigs, and rodents have been experimentally infectedFootnote 17Footnote 31.

Infectious dose

The infectious dose for anthrax varies from species to species and is dependent on route of exposure and the strainFootnote 31. The infectious dose for humans is unknown but in the absence of skin lesions (which lowers the infectious dose for cutaneous anthrax), the 50% infective doses (ID50) are in the thousands or tens of thousands of sporesFootnote 9. The infectious doses for cutaneous and injection-associated anthrax are considerably lowerFootnote 31.

Incubation period

In humans, the incubation period for anthrax ranges from 12 hours to 7 days for cutaneous anthrax; 2 to 5 days for ingestion-associated anthrax; 1 to 6 days for inhalational anthrax; and 1 to 10 days for intravenous anthraxFootnote 9. For trade purposes, the World organisation for Animal Health (WOAH- founded as OIE) incubation period for animals is up to 20 daysFootnote 7.


Anthrax occurs from human exposure to B. anthracis spores via contact with mucous membranes or broken skin, ingestion, inhalation, or injectionFootnote 7Footnote 8. Human exposure commonly occurs due to handling or consumption of animal meat or handling of animal products (e.g., hides, hair, bones, wool) containing B. anthracis sporesFootnote 7. Human-to-human transmission is rare and has not been reported for gastrointestinal or inhalation anthrax. Maternal-fetal transmission is possibleFootnote 32Footnote 33.

The primary route of infection with B. anthracis in herbivores is via inhalation or ingestion of spores while grazing or via ingestion of contaminated feedFootnote 13. In some areas, transmission may be aided by fliesFootnote 13Footnote 17. Carnivorous and omnivorous animals can become infected via consumption of B. anthracis-infected animalsFootnote 13.

Section III – Dissemination


B. anthracis spores can remain viable in animal carcasses, animal products, and soil for many yearsFootnote 7.


Anthrax is primarily a disease of herbivorous mammals. It can be transmitted to humans via contact with infected animal meat and animal productsFootnote 7.


Insects (e.g., tabanids) may play a role as a mechanical vector capable of transmitting B. anthracis in some ecosystemsFootnote 17Footnote 22Footnote 31Footnote 34Footnote 35Footnote 36.

Section IV – Stability and viability

Drug susceptibility/resistance

B. anthracis is susceptible to penicillinFootnote 7; ampicillinFootnote 7; amoxicillinFootnote 7; fluoroquinolones (e.g., ciprofloxacin, levofloxacin, gatifloxacin, ofloxacin)Footnote 7Footnote 37; macrolides (e.g., clindamycin, clarithromycin, erythromycin)Footnote 7; aminoglycosides (e.g., streptomycin, gentamicin)Footnote 1Footnote 7; tetracyclines (e.g., doxycycline, omadacycline)Footnote 7Footnote 38Footnote 39; vancomycinFootnote 7Footnote 40; and rifampinFootnote 7Footnote 41Footnote 42. Other promising antimicrobial agents effective against B. anthracis include daptomycin and dalbavancinFootnote 37.

Naturally occurring penicillin-resistant B. anthracis strains have been isolated occasionallyFootnote 43Footnote 44Footnote 45. Many B. anthracis strains are resistant to some cephalosporins (e.g., cefuroxime, ceftriaxone, ceftazidime)Footnote 45Footnote 46Footnote 47.

Susceptibility to disinfectants

Vegetative cells are highly susceptible to a range of disinfectants, including ethanol and chlorine, whereas B. anthracis spores require high doses and/or extensive contact time to achieve adequate reduction of viable sporesFootnote 48Footnote 49. Ozone gas (9,800-12,000 ppm for 6 to 12 hours, 85% relative humidity)Footnote 49; hydrogen peroxide vapour treatment for 90 minutesFootnote 49Footnote 50; iodine vapour (2.1 mg/L for 24 hours, 90% relative humidity)Footnote 51; chlorine dioxide gas (100-300 ppm for 3 to 12 hours, 75% relative humidity)Footnote 52; formaldehyde gas (10.5 g solid paraformaldehyde per m3)Footnote 49; methyl bromide gas (300 mg/L for 18 hours, 75% relative humidity)Footnote 49; methyl iodide gas (100-400 mg/L, 12 to 48 hours)Footnote 53; and metam sodiumFootnote 54 completely inactivated or achieved ≥6-log reduction of B. anthracis (or surrogate) spores. Sodium hypochlorite (2%)Footnote 49, formaldehyde (5-38%)Footnote 49Footnote 55, and peracetic acid (2%)/0.2% surfactantFootnote 49Footnote 56 also effectively inactivated B. anthracis spores.

Physical inactivation

Heat treatment (75-80 °C, 70-90% relative humidity) for 7 days resulted in a 7-log reduction of surrogate spores on surfacesFootnote 57. Gamma irradiation (>30 kGy) has been used to achieve 6-log reduction in B. anthracis sporesFootnote 58.

Survival outside host

B. anthracis spores can persist for years on various materials, particularly under dry conditionsFootnote 7Footnote 59. B. anthracis spores can remain viable in dry soil for 60 yearsFootnote 60. Viable B. anthracis spores were recovered from bones that were over 200 years oldFootnote 7.

Section V – First aid/medical


B. anthracis can be isolated from clinical samples using culture-based methodsFootnote 7Footnote 8. Blood smears can be stained and examined microscopically for presence of capsulated, Gram-positive bacilliFootnote 7. Gamma phage susceptibility assay, PCR, and immunoassays can also be used to confirm presence of B. anthracisFootnote 8Footnote 61Footnote 62. Xenodiagnosis has also been usedFootnote 22.

Note: The specific recommendations for surveillance in the laboratory should come from the medical surveillance program, which is based on a local risk assessment of the pathogens and activities being undertaken, as well as an overarching risk assessment of the biosafety program as a whole. More information on medical surveillance is available in the Canadian Biosafety Handbook (CBH).

First aid/treatment

Human anthrax can be treated successfully with appropriate antibiotics and supportive care to manage symptomsFootnote 7. Duration of treatment can range from 3 to 60 daysFootnote 7Footnote 9. Some licensed immunoglobulin products have been approved in Canada for use in combination with antibiotics for the treatment of toxaemia associated with inhalational anthrax, including Anthrasil® (AIGIV, Emergent BioSolutions, Winnipeg, MB)Footnote 9Footnote 63. Surgical treatment to remove damaged/necrotic tissue and reconstructive surgery may be necessary in some casesFootnote 12Footnote 64.

Similarly, animals can be treated successfully with appropriate antibiotics, particularly when administered early in the course of infectionFootnote 7.

Note: The specific recommendations for first aid/treatment in the laboratory should come from the post-exposure response plan, which is developed as part of the medical surveillance program. More information on the post-exposure response plan can be found in the CBH.


Human vaccines include BioThrax® (Anthrax Vaccine Adsorbed, Emergent Biosolutions, Winnipeg, MB) approved in the United States and Canada, Anthrax Vaccine Precipitated in the United KingdomFootnote 7, Russian live attenuated anthrax vaccine, and live attenuated strain A16R in ChinaFootnote 7Footnote 65Footnote 66.

Most veterinary vaccines use a live attenuated, toxigenic, non-capsulating B. anthracis Sterne strain 34F2 or an analogous strainFootnote 7. About 200 million doses of anthrax vaccine are used for livestock globally every yearFootnote 16.

Note: More information on the medical surveillance program can be found in the CBH, and by consulting the Canadian Immunization Guide.


In the United States and Canada, BioThrax® is licensed for post-exposure prophylaxis of anthrax in combination with antimicrobial drugs (e.g., ciprofloxacin, doxycycline) for up to 60 daysFootnote 7Footnote 8Footnote 67.

Note: More information on prophylaxis as part of the medical surveillance program can be found in the CBH.

Section VI – Laboratory hazard

Laboratory-acquired infections

Prior to 1975, forty-five cases of laboratory-acquired anthrax were reportedFootnote 68. Five of the infections were fatalFootnote 68. In 2002, a laboratory worker in the United States developed cutaneous anthrax after handling a contaminated vial with bare handsFootnote 69.

Note: Please consult the Canadian Biosafety Standard (CBS) and CBH for additional details on requirements for reporting exposure incidents. A Canadian biosafety guideline describing notification and reporting procedures is also available.


Blood, skin lesion exudates, cerebrospinal fluid, pleural fluid, sputum, fecesFootnote 8.

Primary hazards

Primary hazards in a laboratory setting are exposure of broken skin or mucous membranes to infectious material (e.g., cultures, contaminated laboratory surfaces), autoinoculation, and exposure to infectious aerosolsFootnote 70.

Special hazards


Section VII – Exposure controls/personal protection

Risk group classification

B. anthracis is a Risk Group 3 (RG3) Human Pathogen, a RG3 Animal Pathogen, and a Security Sensitive Biological Agent (SSBA)Footnote 71Footnote 72. B. anthracis Sterne strain, Weybridge strain, and Ames35 strain are RG2 human pathogens and RG2 animal pathogensFootnote 72.

Containment requirements

Containment Level 3 facilities, equipment, and operational practices outlined in the CBS are required for work involving infectious or potentially infectious materials, animals, or cultures.

Containment Level 2 facilities, equipment, and operational practices outlined in the CBS are required for work involving B. anthracis Sterne strain, Weybridge strain, and Ames35 strain.

Note: There are additional security requirements, such as obtaining a Human Pathogens and Toxins Act Security Clearance, for work involving SSBAs.

Protective clothing

The applicable Containment Level 2 or Containment Level 3 requirements for personal protective equipment (PPE) and clothing outlined in the CBS to be followed for work involving RG2 and RG3 B. anthracis strains, respectively. At minimum, use of full body coverage dedicated protective clothing, dedicated protective footwear and/or additional protective footwear, gloves when handling infectious materials or animals, face protection when there is a known or potential risk of exposure to splashes or flying objects, respirators when there is a risk of exposure to infectious aerosols, and an additional layer of protective clothing prior to work with infectious materials or animals.

A local risk assessment will identify the appropriate hand, foot, head, body, eye/face, and respiratory protection, and the personal protective equipment requirements for the containment zone and work activities must be documented.

Other precautions

All activities involving open vessels of pathogens are to be performed in a certified biological safety cabinet (BSC) or other appropriate primary containment device. The use of needles, syringes, and other sharp objects are to be strictly limited. Additional precautions must be considered with work involving animals or large-scale activities.

Additional information

For diagnostic laboratories handling primary specimens that may contain B. anthracis, the following resources may be consulted:

Section VIII – Handling and storage


Allow aerosols to settle. Wearing personal protective equipment, gently cover the spill with absorbent paper towel and apply suitable disinfectant, starting at the perimeter and working towards the centre. Allow sufficient contact time before clean up (CBH).


All materials/substances that have come in contact with the regulated materials should be completely decontaminated before they are removed from the containment zone or standard operating procedures (SOPs) to be in place to safely and securely move or transport waste out of the containment zone to a designated decontamination area / third party. This can be achieved by using decontamination technologies and processes that have been demonstrated to be effective against the regulated material, such as chemical disinfectants, autoclaving, irradiation, incineration, an effluent treatment system, or gaseous decontamination (CBH).


The applicable Containment Level 2 or Containment Level 3 requirements for storage outlined in the CBS are to be followed for work involving RG2 and RG3 B. anthracis strains, respectively. Primary containers of regulated materials removed from the containment zone to be stored in a labelled, leak-proof, impact-resistant secondary container, and kept either in locked storage equipment or within an area with limited access.

Containers of SSBAs stored outside the containment zone must be labelled, leakproof, impact resistant, and kept in locked storage equipment that is fixed in place (i.e., non-movable) and within an area with limited access.

An inventory of RG3 and SSBA in long-term storage, to be maintained and to include:

  • specific identification of the pathogens, toxins, and other regulated materials
  • a means to allow for the detection of a missing or stolen sample in a timely manner

Section IX – Regulatory and other information

Canadian regulatory information

Controlled activities with B. anthracis require a Human Pathogens and Toxins licence, issued by the Public Health Agency of CanadaFootnote 71. Anthrax is a World organisation for Animal health (WOAH) listed disease (founded as OIE), and a nationally notifiable disease and a reportable disease in CanadaFootnote 71Footnote 73.

The following is a non-exhaustive list of applicable designations, regulations, or legislations:

Last file update


Prepared by

Centre for Biosecurity, Public Health Agency of Canada.


The scientific information, opinions, and recommendations contained in this Pathogen Safety Data Sheet have been developed based on or compiled from trusted sources available at the time of publication. Newly discovered hazards are frequent and this information may not be completely up to date. The Government of Canada accepts no responsibility for the accuracy, sufficiency, or reliability or for any loss or injury resulting from the use of the information.

Persons in Canada are responsible for complying with the relevant laws, including regulations, guidelines and standards applicable to the import, transport, and use of pathogens in Canada set by relevant regulatory authorities, including the Public Health Agency of Canada, Health Canada, Canadian Food Inspection Agency, Environment and Climate Change Canada, and Transport Canada. The risk classification and related regulatory requirements referenced in this Pathogen Safety Data Sheet, such as those found in the Canadian Biosafety Standard, may be incomplete and are specific to the Canadian context. Other jurisdictions will have their own requirements.

Copyright © Public Health Agency of Canada, 2023, Canada


Footnote 1

Logan, N. A., and P. de Vos. 2009. Genus I. Bacillus, p. 21. P. de Vos, G. M. Garrity, D. Jones, N. R. Krieg, W. Ludwig, F. A. Rainey, K. H. Schleifer, and W. B. Whitman (eds.), Bergey's Manual of Systemic Bacteriology, Second Edition, Volume Three: The Firmicutes. Springer.

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Footnote 2

Laforce, F. M. 1978. Woolsorters' disease in England. Bull. N. Y. Acad. Med. 54:956-963.

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Footnote 3

Gainer, R. 2016. Yamal and anthrax. Can. Vet. J. 57:985-987.

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Footnote 4

Sommer, D. D., S. Ratnayake, D. Radune, K. Parker, S. Enke, T. M. Ferguson, M. Lovett, A. Mallonee, Z. Rae, M. J. Rosovitz, L. F. Diviak, M. B. Friss, J. P. Klubnik, K. H. Fronda, G. P. Horn, T. E. Blank, R. K. Pope, P. C. Hanna, N. H. Bergman, and A. L. Bazinet. 2020. Draft Genome Sequences of Five Historical Bacillus anthracis Strains. Microbiol. Resour. Announc. 9:e01130-19.

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Footnote 5

Yang, A., J. C. Mullins, M. Van Ert, R. A. Bowen, T. L. Hadfield, and J. K. Blackburn. 2020. Predicting the Geographic Distribution of the Bacillus anthracis A1.a/Western North American Sub-Lineage for the Continental United States: New Outbreaks, New Genotypes, and New Climate Data. Am. J. Trop. Med. Hyg. 102:392-402.

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Footnote 6

Moayeri, M., and S. H. Leppla. 2009. Cellular and systemic effects of anthrax lethal toxin and edema toxin. Mol. Aspects Med. 30:439-455.

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Footnote 7

World Health Organization, and International Office of Epizootics. 2008. Anthrax in humans and animals. World Health Organization.

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Footnote 8

Sweeney, D. A., C. W. Hicks, X. Cui, Y. Li, and P. Q. Eichacker. 2011. Anthrax infection. American Journal of Respiratory and Critical Care Medicine. 184:1333-1341.

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Footnote 9

Savransky, V., B. Ionin, and J. Reece. 2020. Current Status and Trends in Prophylaxis and Management of Anthrax Disease. Pathogens. 9:370.

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Footnote 10

Doganay, M., and H. Demiraslan. 2015. Human anthrax as a re-emerging disease. Recent. Pat. Antiinfect Drug Discov. 10:10-29.

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Footnote 11

Holty, J. E., D. M. Bravata, H. Liu, R. A. Olshen, K. M. McDonald, and D. K. Owens. 2006. Systematic review: a century of inhalational anthrax cases from 1900 to 2005. Ann. Intern. Med. 144:270-280.

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Footnote 12

Zasada, A. A. 2018. Injectional anthrax in human: A new face of the old disease. Adv. Clin. Exp. Med. 27:553-558.

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Footnote 13

Beyer, W., and P. Turnbull. 2009. Anthrax in animals. Mol. Aspects Med. 30:481-489.

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Footnote 14

Prins, H. H. T., and F. J. Weyerhaeuser. 1987. Epidemics in Populations of Wild Ruminants: Anthrax and Impala, Rinderpest and Buffalo in Lake Manyara National Park, Tanzania. Oikos. 49:28-38.

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Footnote 15

Driciru, M., I. B. Rwego, B. Asiimwe, D. A. Travis, J. Alvarez, K. VanderWaal, and K. Pelican. 2018. Spatio-temporal epidemiology of anthrax in Hippopotamus amphibious in Queen Elizabeth Protected Area, Uganda. PLoS One. 13:e0206922.

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Footnote 16

Carlson, C. J., I. T. Kracalik, N. Ross, K. A. Alexander, M. E. Hugh-Jones, M. Fegan, B. T. Elkin, T. Epp, T. K. Shury, W. Zhang, M. Bagirova, W. M. Getz, and J. K. Blackburn. 2019. The global distribution of Bacillus anthracis and associated anthrax risk to humans, livestock and wildlife. Nat. Microbiol. 4:1337-1343.

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Footnote 17

Hugh-Jones, M., and J. Blackburn. 2009. The ecology of Bacillus anthracis. Mol. Aspects Med. 30:356-367.

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Footnote 18

Turner, W. C., P. Imologhome, Z. Havarua, G. P. Kaaya, J. K. E. Mfune, I. D. T. Mpofu, and W. M. Getz. 2013. Soil ingestion, nutrition and the seasonality of anthrax in herbivores of Etosha National Park. Ecosphere. 4:art13.

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Footnote 19

Lewerin, S. S., M. Elvander, T. Westermark, L. N. Hartzell, A. K. Norström, S. Ehrs, R. Knutsson, S. Englund, A. C. Andersson, M. Granberg, S. Bäckman, P. Wikström, and K. Sandstedt. 2010. Anthrax outbreak in a Swedish beef cattle herd--1st case in 27 years: Case report. Acta Vet. Scand. 52:7-0147-52-7.

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Footnote 20

Berger, S. 2019. Anthrax: Global Status. Gideon Informatics, Inc.

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Footnote 21

Hueffer, K., D. Drown, V. Romanovsky, and T. Hennessy. 2020. Factors Contributing to Anthrax Outbreaks in the Circumpolar North. Ecohealth. 17:174-180.

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Footnote 22

Gainer, R. S. 2018. Spore concentration and modified host resistance as cause of anthrax outbreaks: A practitioner's perspective. Can. Vet. J. 59:185-187.

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Footnote 23

Cossaboom, C. M., S. Khaiseb, B. Haufiku, P. Katjiuanjo, A. Kannyinga, K. Mbai, T. Shuro, J. Hausiku, A. Likando, R. Shikesho, K. Nyarko, L. A. Miller, S. Agolory, A. R. Vieira, J. S. Salzer, W. A. Bower, L. Campbell, C. B. Kolton, C. Marston, J. Gary, B. C. Bollweg, S. R. Zaki, A. Hoffmaster, and H. Walke. 2019. Anthrax Epizootic in Wildlife, Bwabwata National Park, Namibia, 2017. Emerg. Infect. Dis. 25:947-950.

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Footnote 24

Nekorchuk, D. M., L. R. Morris, V. Asher, D. L. Hunter, S. J. Ryan, and J. K. Blackburn. 2019. Potential Bacillus anthracis Risk Zones for Male Plains Bison in Southwestern Montana, USA. J. Wildl. Dis. 55:136-141.

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Footnote 25

Walsh, M. G., S. M. Mor, and S. Hossain. 2019. The elephant-livestock interface modulates anthrax suitability in India. Proc. Biol. Sci. 286:20190179.

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Footnote 26

Muturi, M., J. Gachohi, A. Mwatondo, I. Lekolool, F. Gakuya, A. Bett, E. Osoro, A. Bitek, S. M. Thumbi, P. Munyua, H. Oyas, O. N. Njagi, B. Bett, and M. K. Njenga. 2018. Recurrent Anthrax Outbreaks in Humans, Livestock, and Wildlife in the Same Locality, Kenya, 2014-2017. Am. J. Trop. Med. Hyg. 99:833-839.

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Footnote 27

Mullins, J. C., M. Van Ert, T. Hadfield, M. P. Nikolich, M. E. Hugh-Jones, and J. K. Blackburn. 2015. Spatio-temporal patterns of an anthrax outbreak in white-tailed deer, Odocoileus virginanus, and associated genetic diversity of Bacillus anthracis. BMC Ecol. 15:23-015-0054-8.

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Footnote 28

Morris, L. R., K. M. Proffitt, V. Asher, and J. K. Blackburn. 2016. Elk Resource Selection and Implications for Anthrax Management in Montana. J. Wildl. Manage. 80:235-244.

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Footnote 29

Ekebas, G., A. Atasever, D. Y. Gram, E. Karakaya, S. Abay, F. Aydin, K. S. Gumussoy, and M. Sahin. 2018. A case of Anthrax in two captive pumas (Puma concolor). J. Vet. Med. Sci. 80:1875-1880.

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Footnote 30

Padhi, L., S. K. Panda, P. P. Mohapatra, and G. Sahoo. 2020. Antibiotic susceptibility of cultivable aerobic microbiota from the oral cavity of Echis carinatus from Odisha (India). Microb. Pathog. 143:104121.

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Footnote 31

Watson, A., and D. Keir. 1994. Information on which to base assessments of risk from environments contaminated with anthrax spores. Epidemiol. Infect. 113:479-490.

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Footnote 32

Meaney-Delman, D., M. E. Zotti, S. A. Rasmussen, S. Strasser, S. Shadomy, R. M. Turcios-Ruiz, G. D. Wendel Jr, T. A. Treadwell, and D. J. Jamieson. 2012. Anthrax cases in pregnant and postpartum women: a systematic review. Obstet. Gynecol. 120:1439-1449.

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Footnote 33

Dettwiler, M., K. Mehinagic, S. Gobeli Brawand, A. Thomann, S. Feyer, L. Hüsser, G. Theubet, J. Gigandet, S. Rottenberg, and H. Posthaus. 2018. Bacillus anthracis as a cause of bovine abortion - a necropsy case requiring special biosafety measures. Schweiz. Arch. Tierheilkd. 160:547-552.

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Footnote 34

Gogarten, J. F., A. Düx, B. Mubemba, K. Pléh, C. Hoffmann, A. Mielke, J. Müller-Tiburtius, A. Sachse, R. M. Wittig, S. Calvignac-Spencer, and F. H. Leendertz. 2019. Tropical rainforest flies carrying pathogens form stable associations with social nonhuman primates. Mol. Ecol. 28:4242-4258.

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Footnote 35

Blackburn, J. K., M. Van Ert, J. C. Mullins, T. L. Hadfield, and M. E. Hugh-Jones. 2014. The necrophagous fly anthrax transmission pathway: empirical and genetic evidence from wildlife epizootics. Vector Borne Zoonotic Dis. 14:576-583.

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Footnote 36

Fasanella, A., G. Garofolo, M. Galella, P. Troiano, C. De Stefano, L. Pace, A. Aceti, L. Serrecchia, and R. Adone. 2013. Suspect vector transmission of human cutaneous anthrax during an animal outbreak in Southern Italy. Vector Borne Zoonotic Dis. 13:769-771.

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Footnote 37

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