For centuries, flies have been dismissed as nothing more than pests, buzzing around garbage or landing on food. Yet, these small insects hold a dark secret: they are among the most efficient carriers of disease in the natural world.
Recent research led by Dr. John G. Stoffolano Jr. reveals that synanthropic flies—species like houseflies (Musca domestica), blowflies (Calliphoridae), and flesh flies (Sarcophagidae) that thrive near humans—play a critical role in spreading pathogens between animals, humans, and the environment.
The Ancient Partnership Between Flies and Pathogens
Flies have existed for over 400 million years, long before humans walked the Earth. Their evolutionary journey has equipped them with unique tools to survive in diverse environments, including human settlements. One such tool is the diverticulated crop, a specialized organ in their digestive system.
The crop acts as a storage pouch, separate from the acidic midgut where digestion occurs. This separation allows pathogens ingested from feces, rotting meat, or open wounds to survive—and even multiply—for days.
For example, studies show that Bacillus anthracis, the bacterium causing anthrax (a deadly disease affecting both animals and humans), remains viable in the crop for up to four days. Similarly, the virus responsible for COVID-19, SARS-CoV-2, can cling to a housefly’s body long enough to infect new hosts.
Why the crop matters: Unlike the midgut, which breaks down food with enzymes, the crop’s neutral pH creates a safe haven for microbes. This organ essentially acts as a “biological backpack,” allowing flies to transport pathogens over distances.
For instance, when a blowfly feeds on an infected carcass, it stores harmful bacteria or spores in its crop and later regurgitates them onto plants, water, or food. This process, known as mechanical transmission, enables diseases to jump between species and environments.
This relationship between flies and pathogens is not new. Medieval records often linked fly swarms to plague outbreaks, though the science behind this connection was poorly understood. Today, advanced molecular tools confirm what ancient observers suspected: flies are far more than passive carriers. They actively shape disease dynamics by moving pathogens across ecosystems.
How Flies Spread Disease: A Closer Look
To understand why flies are such effective disease vectors, we must examine their feeding habits and biology. Flies lack teeth, so they liquefy food by regurgitating digestive enzymes onto it. This process, called “bubbling”, allows them to ingest nutrients but also leaves behind contaminated droplets. When a fly feeds on infected material—like animal feces or a decaying carcass—pathogens enter its crop. Later, these microbes are deposited onto human food, water, or wounds through regurgitation or defecation.
- Regurgitation: Flies vomit small droplets of crop fluid to lighten their bodies before flight. These droplets often contain live pathogens.
- Defecation: Undigested material from the midgut is expelled as feces, which can contaminate surfaces.
- Surface contact: Pathogens stick to a fly’s legs, wings, or mouthparts and are transferred during feeding or grooming.

A striking example comes from Kruger National Park in South Africa. Blowflies feeding on anthrax-infected animal carcasses were found to regurgitate spores onto nearby plants. Grazing animals like kudus then consumed these plants, sparking outbreaks.
In Thailand, houseflies were linked to 33% of diarrheal cases in a rural village, spreading bacteria like Shigella (which causes dysentery) and Salmonella (a common foodborne pathogen) through contaminated food. Even more alarming, lab experiments confirm that houseflies can mechanically transmit SARS-CoV-2, raising concerns about their role in COVID-19 spread in crowded, unsanitary environments.
The problem is amplified by flies’ ability to travel long distances. While a single fly might only move a few meters after feeding, swarms can disperse pathogens across entire regions. For instance, during the 2011 anthrax outbreak in Zambia, flies feeding on a dead hippopotamus likely transferred spores to nearby vegetation, contributing to 511 human cases and five deaths.
The Crop: A Hidden Hub for Pathogen Evolution
The diverticulated crop is not just a storage unit—it’s a hotspot for microbial activity. Here, bacteria and viruses evade the fly’s immune defenses and interact with other microbes. This environment fosters
horizontal gene transfer, a process where bacteria exchange genetic material, including antibiotic-resistant genes.For example, studies show that houseflies can spread drug-resistant E. coli between livestock and humans, with resistance genes transferring at a rate of 10⁻² (transfers per day) in their crops.
Horizontal gene transfer explained: This process allows bacteria to share traits like antibiotic resistance, even between unrelated species. In the crop, resistant bacteria can pass genes to harmless microbes, creating “superbugs” that drugs cannot treat.
Moreover, the crop’s neutral pH allows pathogens to thrive. For example, Chlamydia trachomatis, which causes eye infections, survives in the crop for 24 hours. When flies feed on eye secretions, they deposit the bacterium into new hosts.
Similarly, the orf virus, which causes painful sores in livestock, remains infectious in fly regurgitant for six hours. These findings underscore the crop’s role as a “bioreactor” where pathogens adapt and evolve.
Flies as Bridges Between Species
Flies do not discriminate between hosts. They feed on bats, birds, livestock, and humans, creating pathways for zoonotic diseases—those that jump from animals to humans. Bats, which harbor over 320,000 undiscovered viruses (Anthony et al., 2013), are a prime example.
Flies feeding on bat guano (feces) or half-eaten fruits can transfer viruses like Nipah (a deadly virus causing brain inflammation) to humans. In Uganda, flies were implicated in spreading Bacillus cereus Biovar anthracis from mangabey monkeys to humans, causing sylvatic anthrax—a form of the disease that circulates in wildlife.

Non-human primates (NHPs) are equally vulnerable. Flies follow chimpanzee troops for up to 1.3 kilometers, feeding on their feces and wounds. This behavior facilitates the spread of diseases like yaws, a debilitating infection caused by Treponema pallidum pertenue. DNA analysis confirmed the pathogen in fly crops, linking their feeding habits to outbreaks in wildlife and humans.
Why Female Flies Pose a Greater Threat
Female flies are disproportionately responsible for disease transmission. Their need for protein to develop eggs drives them to seek out nutrient-rich—and often pathogen-laden—sources like carcasses and feces. Gravid females (those carrying eggs) produce 6.5 times more defecation spots than males, increasing contamination risks.
Larger species, such as blowflies, pose an even greater threat due to their bigger crops, which can store 18 microliters of fluid—enough to hold millions of bacterial spores. In agricultural settings, female flies exacerbate foodborne illnesses. A study in the U.S. found that houseflies contaminated 65% of spinach samples with E. coli, highlighting their role in farm-to-table disease spread.
Climate Change and Urbanization: Fueling the Fly-Disease Cycle
As global temperatures rise, flies are thriving. A 1°C increase in temperature boosts fly activity by 10% in tropical regions, accelerating their breeding cycles. Urbanization compounds the problem. Slums and wet markets (where live animals are sold) create ideal habitats for flies.
In Malaysia, researchers collected 1,037 flies from garbage sites, kitchens, and cafeterias, with Muscidae (houseflies) being the most common. These environments are ticking time bombs for disease emergence.
Antibiotic resistance adds another layer of complexity. Flies frequent hospitals, farms, and waste sites, picking up drug-resistant bacteria. In Bangladesh, 75% of houseflies carried multidrug-resistant E. coli (strains immune to multiple antibiotics), posing a dire threat to public health.
Combating Fly-Borne Diseases: Strategies for the Future
Addressing fly-borne diseases requires a multifaceted approach. Improved sanitation is the first line of defense. Reducing organic waste disrupts fly breeding sites, while sealing food and managing landfills limits their access to pathogens. In rural Zambia, where bushmeat consumption triggered an anthrax outbreak, education campaigns could reduce risks by promoting safer food practices.
Technology also plays a role. DNA metabarcoding, a technique that identifies pathogens in fly populations using genetic markers, could revolutionize disease surveillance. By analyzing fly samples from outbreak zones, scientists can detect emerging threats before they spread.
Finally, global collaboration is essential. Flies know no borders, and neither do pathogens. International efforts to monitor fly populations in high-risk areas—like tropical forests and urban slums—could prevent the next pandemic.
Conclusion
Flies have been humanity’s unwelcome companions for millennia. Yet, their role in disease transmission has been underestimated, overshadowed by more visible threats like mosquitoes. Dr. Stoffolano’s research urges us to reconsider this oversight. From ancient plagues to modern pandemics, flies have shaped human history in ways we are only beginning to understand.
The key to mitigating their impact lies in recognizing their biology and behavior. By studying their crop, tracking their movements, and addressing the environmental factors that fuel their spread, we can disrupt the cycle of disease. In a world grappling with climate change and antibiotic resistance, this knowledge is not just academic—it’s a matter of survival.
Frequently Asked Questions (FAQs) and Concepts
Synanthropic Flies: Flies that live in close association with humans, such as houseflies (Musca domestica), blowflies (Calliphoridae), and flesh flies (Sarcophagidae). These insects thrive in human environments like garbage dumps, kitchens, and farms. Their importance lies in their ability to spread diseases by transferring pathogens from waste, carcasses, or bodily fluids to food, water, or humans. For example, houseflies contaminated 33% of diarrheal cases in a Thai village by spreading bacteria like Shigella.
Diverticulated Crop: A pouch-like organ in a fly’s digestive system that stores food and pathogens before digestion. Unlike the acidic midgut, the crop’s neutral pH allows pathogens to survive and multiply. This organ is critical because it acts as a reservoir for microbes, enabling flies to transmit diseases over distances. For instance, anthrax spores (Bacillus anthracis) can remain viable in the crop for four days, later regurgitated onto plants or food.
Pathogen: A microorganism (bacteria, virus, fungus, or parasite) that causes disease. Pathogens like Salmonella (food poisoning) and *SARS-CoV-2* (COVID-19) are significant because they threaten human and animal health. Flies spread pathogens mechanically—by carrying them on their bodies—or biologically, through their crop and feces.
Mechanical Transmission: The passive transfer of pathogens on a fly’s body (legs, wings, or mouthparts) without the pathogen multiplying. This process is important because it allows rapid contamination of surfaces. For example, houseflies spread E. coli from animal feces to salads or fruits in markets.
Regurgitation: The process where flies vomit digestive enzymes to liquefy food, leaving behind pathogen-filled droplets. This is crucial for disease spread, as regurgitated fluids can contaminate food. In Kruger National Park, blowflies regurgitated anthrax spores onto plants, infecting grazing animals.
Defecation: The expulsion of undigested material from the fly’s midgut, often containing live pathogens. Defecation spots on food or surfaces spread diseases like cholera. Female flies defecate more frequently when developing eggs, increasing contamination risks.
Horizontal Gene Transfer: The exchange of genetic material (e.g., antibiotic resistance genes) between bacteria, even across species. This process in the fly crop creates “superbugs.” For example, houseflies transferred tetracycline resistance between E. coli strains at a rate of 10⁻² (transfers per day) in lab studies.
Antibiotic Resistance: The ability of bacteria to survive drugs designed to kill them. Flies spread resistant bacteria from hospitals or farms to humans. In Bangladesh, 75% of houseflies carried multidrug-resistant E. coli, complicating treatment of infections.
Zoonotic Diseases: Infections that jump from animals to humans, like anthrax or Nipah virus. Flies bridge species by feeding on both wildlife and humans. For example, flies transferred Bacillus cereus Biovar anthracis from monkeys to humans in Uganda.
Bacillus anthracis: The bacterium causing anthrax, a deadly disease affecting livestock and humans. Flies spread its spores through regurgitation. In Zambia, flies feeding on a hippopotamus carcass contributed to 511 human anthrax cases.
SARS-CoV-2: The virus causing COVID-19. Lab studies show houseflies can mechanically carry the virus, raising concerns about their role in outbreaks in crowded, unsanitary areas.
E. coli: A bacteria species, some strains of which cause severe food poisoning. Flies spread E. coli from feces to food. A U.S. study found flies contaminated 65% of spinach samples with this pathogen.
Shigella: Bacteria causing dysentery (severe diarrhea with blood). Flies spread Shigella in unsanitary conditions. In Thailand, they were linked to 33% of diarrheal cases in villages.
Salmonella: A foodborne pathogen causing fever and stomach cramps. Flies transfer Salmonella from animal waste to human food, especially in farms or markets.
Chlamydia trachomatis: A bacterium causing eye infections (trachoma) and sexually transmitted diseases. Houseflies store it in their crop for 24 hours, transmitting it through contact with eye secretions.
Orf Virus: A virus causing painful sores in livestock. Flies can carry the virus in regurgitant for six hours, spreading it to animals or humans handling infected sheep or goats.
Nipah Virus: A deadly virus transmitted from bats to humans via contaminated food or fluids. Flies feeding on bat guano or half-eaten fruits may spread it, as seen in outbreaks in South Asia.
Non-Human Primates (NHPs): Animals like chimpanzees or gorillas. Flies follow NHPs to feed on their feces or wounds, spreading diseases like yaws. In Tanzania, flies transferred Treponema pallidum pertenue between baboons.
Sylvatic Anthrax: A form of anthrax circulating in wildlife. Flies spread it between animals and humans, as seen in Uganda where mangabey monkeys were linked to human cases.
Yaws: A tropical infection causing skin sores, spread by flies feeding on wounds. DNA studies found the yaws pathogen (Treponema pallidum pertenue) in fly crops in Tanzania.
Gravid Females: Female flies carrying eggs. They pose higher risks because they seek protein-rich (and pathogen-laden) food like carcasses. Gravid houseflies produce 6.5 times more defecation spots than males.
DNA Metabarcoding: A technique identifying pathogens in flies using genetic markers. It helps track disease sources. For example, scientists used it to link flies to Bacillus cereus Biovar anthracis in Uganda.
Wet Markets: Markets selling live animals, often in crowded conditions. These hubs attract flies, which spread pathogens between animals and humans. The 2003 SARS outbreak was linked to wildlife in wet markets.
Climate Change: Rising global temperatures increase fly activity. A 1°C temperature rise boosts fly breeding by 10% in tropical regions, escalating disease risks like malaria or dengue.
Urbanization: The growth of cities creating fly-friendly habitats (e.g., slums, garbage piles). In Malaysia, 1,037 flies were collected from urban sites, most being disease-carrying houseflies.
Reference:
Stoffolano Jr, J. G. (2022). Synanthropic flies—a review including how they obtain nutrients, along with pathogens, store them in the crop and mechanisms of transmission. Insects, 13(9), 776.





