Crayfish Plague (Aphanomyces astaci)

Quick Facts

🏥 Condition Name
Crayfish Plague (Aphanomyces astaci)
📋 Also Known As
Crayfish Fungus, Aphanomycosis, Crayfish Pestilence
📂 Category
Invertebrates
📁 Subcategory
Crustaceans - Crayfish
🦂 Affects
Exoskeleton, nervous system, and internal organs of freshwater crayfish
🏷️ Type
Fungal (Oomycete)
⚠️ Severity
Often fatal
💊 Treatable
No effective treatment available
🔄 Contagious
Highly contagious between crayfish
🧬 Hereditary
No
🦂 Common In
Native European, Australian, and Asian crayfish species; North American species are carriers

Crayfish plague (Aphanomyces astaci) - devastating to native species Overview

Crayfish plague, caused by the oomycete pathogen Aphanomyces astaci, represents one of the most devastating diseases affecting freshwater crayfish populations worldwide. This waterborne pathogen has been responsible for catastrophic declines in native crayfish populations across Europe, Australia, and parts of Asia since its introduction in the mid-nineteenth century. The disease operates with remarkable efficiency, capable of eliminating entire crayfish populations within weeks of introduction, making it a critical concern for both wild populations and captive crayfish keepers.

The pathogen primarily affects non-North American crayfish species with particular severity. European species such as the noble crayfish (Astacus astacus), white-clawed crayfish (Austropotamobius pallipes), and stone crayfish (Austropotamobius torrentium) exhibit extreme susceptibility, with mortality rates approaching one hundred percent in naive populations. Australian species including the redclaw crayfish (Cherax quadricarinatus) and yabby (Cherax destructor) similarly demonstrate high vulnerability when exposed to the pathogen. North American crayfish species, having coevolved with the pathogen, typically serve as asymptomatic carriers capable of spreading the disease without showing clinical signs.

The impact of crayfish plague on affected populations extends far beyond individual mortality. Complete population collapses can occur within days to weeks of pathogen introduction, fundamentally altering freshwater ecosystem dynamics. Crayfish serve as keystone species in many aquatic environments, and their loss triggers cascading effects throughout the food web. For hobbyists maintaining crayfish in captivity, the introduction of this pathogen can devastate entire collections and potentially spread to other systems sharing equipment or water sources.

Unfortunately, there is currently no effective treatment for crayfish plague once infection has occurred. The pathogen's rapid progression and the lack of effective antifungal treatments that are safe for invertebrates mean that prevention through strict biosecurity measures remains the only viable approach. Affected crayfish typically succumb within days of showing clinical signs, and euthanasia of infected individuals combined with thorough disinfection of affected systems represents the standard response to confirmed infections. Understanding this disease is essential for any crayfish keeper, particularly those maintaining species susceptible to infection.

Causes of Crayfish plague (Aphanomyces astaci) - devastating to native species

The primary cause of crayfish plague is infection by Aphanomyces astaci, an oomycete organism often colloquially referred to as a water mold despite being more closely related to brown algae than true fungi. This pathogen produces motile zoospores that actively seek out crayfish hosts using chemical signals released by the crustaceans. Once zoospores locate a suitable host, they encyst on the exoskeleton and germinate, penetrating through the cuticle to establish infection in the underlying tissues. The pathogen then spreads throughout the host's body, ultimately causing death through a combination of tissue destruction and overwhelming systemic infection.

Environmental factors play a significant role in disease transmission and severity. Water temperature significantly influences zoospore activity and survival, with optimal transmission occurring between fifteen and twenty degrees Celsius. Warmer temperatures accelerate disease progression but may reduce zoospore viability outside the host, while cooler temperatures extend zoospore survival in the water column but slow infection development. Water quality parameters including pH, hardness, and organic content also affect pathogen survival and infectivity, though the organism demonstrates remarkable adaptability across a range of conditions typically suitable for crayfish habitation.

Husbandry-related causes represent the primary route of introduction for captive crayfish keepers. The most common source of infection is the introduction of carrier crayfish, particularly North American species such as signal crayfish (Pacifastacus leniusculus), red swamp crayfish (Procambarus clarkii), or spinycheek crayfish (Faxonius limosus), into systems housing susceptible species. These carrier species shed zoospores continuously without showing disease signs, creating a persistent source of infection. Contaminated equipment, plants, substrate, or water transferred between systems can also introduce the pathogen, as can the introduction of fish or other aquatic organisms that have been housed with infected or carrier crayfish.

Risk factors for severe disease include species susceptibility, with non-North American crayfish demonstrating dramatically higher mortality than their North American counterparts. Individual immune status, influenced by factors such as recent molting, nutritional condition, and stress levels, affects disease progression and survival time. Wild-caught crayfish may carry the pathogen without the keeper's knowledge, while captive-bred individuals from reputable sources typically present lower risk. Population density influences transmission rates, with higher densities facilitating more rapid spread through increased zoospore concentrations and contact opportunities.

The disease mechanism involves progressive invasion of host tissues by fungal hyphae following initial zoospore attachment and germination. The pathogen secretes enzymes that break down chitin and other components of the exoskeleton, allowing penetration into soft tissues. Once established internally, the infection spreads through the hemolymph (the invertebrate equivalent of blood), reaching vital organs and the nervous system. The host's immune system attempts to respond through melanization and hemocyte encapsulation, but susceptible species cannot mount an effective defense. Death typically results from a combination of tissue destruction, metabolic disruption, and secondary infections facilitated by compromised immune function and exoskeletal integrity.

Symptoms & Warning Signs

Early warning signs of crayfish plague infection often manifest as subtle behavioral changes that may be easily overlooked by inexperienced keepers. Affected crayfish frequently demonstrate reduced appetite and decreased interest in food items that would normally elicit immediate feeding responses. Activity levels may decrease, with infected individuals spending more time in sheltered areas and showing less exploratory behavior than usual. Some crayfish exhibit unusual daytime activity when they would normally remain hidden, suggesting disruption of normal behavioral patterns. Early-stage infected crayfish may appear restless or demonstrate repetitive movements, potentially reflecting neurological involvement or discomfort.

Physical symptoms become more apparent as the infection progresses through the host's tissues. White or pale patches may appear on the exoskeleton, particularly around joints and the ventral surface where the cuticle is thinner. These discolored areas represent sites of active fungal growth and tissue damage beneath the shell. The exoskeleton may develop a chalky or dull appearance in contrast to the normal glossy surface of healthy crayfish. Careful examination may reveal small dark spots or lesions where melanization responses have occurred, representing the crayfish's immune system attempting to contain the infection. The walking legs and claws may show particular involvement, with visible deterioration or discoloration of these appendages.

Behavioral changes intensify as the disease advances and systemic involvement increases. Infected crayfish often exhibit pronounced lethargy, remaining motionless for extended periods and showing minimal response to stimuli that would normally provoke defensive or escape behaviors. Coordination problems become evident, with affected individuals demonstrating difficulty walking, climbing, or maintaining normal posture. The characteristic defensive posture with raised claws may be absent or weakly executed, and the crayfish may be unable to right itself if turned over. Complete food refusal is typical in advanced cases, with the crayfish showing no interest in even preferred food items.

Molting-related symptoms present particular concern, as infected crayfish may attempt to molt but fail to complete the process successfully. Incomplete molts leave the crayfish vulnerable and often prove fatal, with individuals becoming trapped in their old exoskeleton. The stress of molting combined with active infection frequently accelerates disease progression and hastens death. Pre-molt crayfish that contract the infection may never successfully complete their next molt, while post-molt individuals with soft new exoskeletons are extremely vulnerable to rapid pathogen invasion.

Symptom progression in crayfish plague is notably rapid compared to many other invertebrate diseases. Initial behavioral changes may be present for only one to three days before more obvious physical symptoms appear. Once physical symptoms become visible, death typically follows within three to seven days in highly susceptible species. Some individuals may survive slightly longer, but recovery from established infection is exceptionally rare in susceptible species. The speed of progression means that by the time keepers notice obvious symptoms, the disease has usually advanced significantly.

Critical and emergency symptoms indicating imminent death include complete paralysis of the limbs while the crayfish remains alive, inability to maintain any normal posture, and visible fungal growth on external surfaces. Milky or cloudy hemolymph visible through thinner portions of the exoskeleton indicates severe systemic infection. Affected crayfish may exhibit terminal behaviors including spastic movements, limb autotomy (self-amputation of limbs), and loss of responsiveness to all stimuli. At this stage, death typically occurs within hours, and other crayfish in the system should be considered exposed and at high risk of developing the disease.

Diagnosis

Visual examination provides the first indicators of potential crayfish plague infection, though definitive diagnosis requires laboratory confirmation. Keepers should carefully observe affected crayfish for the characteristic physical signs including white patches on the exoskeleton, particularly in joint areas and the ventral surface. The presence of visible fungal growth on external surfaces, though occurring only in advanced cases, strongly suggests oomycete infection. Examination of deceased crayfish may reveal extensive internal damage, with the hepatopancreas and muscle tissue appearing abnormally pale or showing signs of liquefaction in severe cases. However, visual signs alone cannot confirm crayfish plague specifically, as other fungal and bacterial infections may produce similar appearances.

Behavioral observation plays a crucial role in early detection and should be conducted regularly by all crayfish keepers. Noting changes in activity levels, feeding behavior, and responses to stimuli helps identify potentially infected individuals before physical symptoms become apparent. Keeping detailed records of normal behavior for each crayfish allows for more accurate detection of subtle changes that might indicate early infection. The characteristic combination of lethargy, loss of coordination, and failure to assume defensive postures provides strong behavioral evidence of disease, though these signs may also accompany other health problems. Separating suspected individuals for closer observation while implementing precautionary biosecurity measures represents prudent practice.

Environmental parameter assessment forms an essential component of the diagnostic process, helping differentiate infectious disease from environmental stress or water quality issues. Testing water temperature, pH, ammonia, nitrite, nitrate, and general hardness establishes whether environmental factors might explain observed symptoms or could be contributing to disease susceptibility. Reviewing recent changes to the system, including new introductions, equipment sharing, or water source modifications, may identify potential pathogen entry points. While environmental factors cannot cause crayfish plague, stress from suboptimal conditions can accelerate disease progression and complicate diagnosis.

Differential diagnosis requires distinguishing crayfish plague from other conditions producing similar symptoms. Bacterial shell disease, other fungal infections including those caused by Saprolegnia species, and various environmental stressors can mimic some aspects of crayfish plague presentation. The rapid progression and high mortality rate characteristic of crayfish plague in susceptible species helps distinguish it from slower-developing conditions. Laboratory diagnosis through polymerase chain reaction testing of tissue samples provides definitive confirmation of Aphanomyces astaci infection. Given the serious implications of a positive diagnosis, keepers experiencing unexplained mortality in susceptible crayfish species should consider submitting samples to appropriate diagnostic laboratories, particularly if there is any possibility of exposure to North American crayfish or contaminated water sources.

Treatment Options

Environmental management constitutes the primary response to suspected or confirmed crayfish plague, as no effective medical treatment currently exists for infected individuals. Immediately isolating suspected infected crayfish prevents further transmission to healthy individuals in the system. Maintaining optimal water quality parameters supports whatever immune response the crayfish can mount, though this rarely affects outcomes in susceptible species. Temperature management may slow disease progression, with cooler temperatures potentially extending survival time, though this does not alter the ultimate outcome for most infected individuals. Ensuring proper oxygenation and minimizing additional stressors removes complications that might accelerate disease progression.

Supportive care for infected crayfish remains limited in effectiveness but may be attempted for valuable individuals or those showing early symptoms. Providing easily accessible hiding places reduces stress from perceived predation risk. Offering highly palatable foods may encourage continued feeding in early-stage infected individuals, supporting energy reserves needed for any immune response. Some keepers report anecdotal success with salt baths using aquarium salt at concentrations of one to two tablespoons per gallon for short-duration treatments, though scientific evidence supporting efficacy against Aphanomyces astaci is lacking. Maintaining pristine water quality through more frequent water changes may reduce environmental pathogen load.

Medical treatment options for crayfish plague remain essentially nonexistent despite extensive research efforts. Antifungal medications effective against oomycetes in other contexts have not demonstrated reliable efficacy in treating infected crayfish. Malachite green, historically used for fungal infections in aquatic organisms, is not effective against Aphanomyces astaci and poses significant toxicity risks to invertebrates. Copper-based treatments, effective against some aquatic pathogens, are absolutely contraindicated as copper is lethal to crayfish and other crustaceans at concentrations far below therapeutic levels. Research into potential treatments continues, but no breakthrough therapeutic has emerged as of current knowledge.

Quarantine protocols represent essential practice for preventing disease spread rather than treating infection. Any crayfish system experiencing unexplained mortality should be immediately isolated from other systems, with all equipment dedicated exclusively to the affected tank. New crayfish acquisitions should undergo minimum thirty-day quarantine before introduction to established systems, with longer periods recommended for species known to be potential carriers. Strict separation between North American crayfish species and susceptible non-North American species must be maintained, ideally in completely separate locations with no possibility of equipment or water sharing.

Treatment monitoring in cases where supportive care is attempted involves close observation for any improvement or stabilization of condition. Continued feeding and maintenance of activity levels may indicate disease resistance, though apparent improvement is rare in susceptible species. Monitoring other crayfish in the affected system for early symptoms allows rapid response if transmission has occurred. Water quality testing should continue throughout any treatment attempt to ensure optimal environmental conditions are maintained.

Recognizing when treatment is not viable is essential for preventing unnecessary suffering. Crayfish showing advanced symptoms including paralysis, visible fungal growth on external surfaces, or complete loss of responsiveness should be humanely euthanized rather than allowed to experience prolonged decline. Clove oil overdose provides a humane euthanasia method for crayfish, with immersion in a solution of water with ten to fifteen drops of clove oil per liter inducing rapid unconsciousness followed by death. Following euthanasia of infected individuals, thorough disinfection of the affected system using bleach solution or complete teardown prevents pathogen persistence and protects any future inhabitants.

Recovery & Prognosis

Recovery from crayfish plague infection is exceptionally rare in susceptible species, with survival rates approaching zero in naive populations of European, Australian, and Asian crayfish. The few documented cases of survival in susceptible species typically involve individuals that received minimal pathogen exposure or possessed unusual individual resistance. Recovery timeline in these rare cases extends over several weeks to months, during which the crayfish remains vulnerable to reinfection and secondary complications. North American crayfish that develop clinical disease may recover, as their coevolved immune systems can sometimes control the infection, though they typically become chronic carriers capable of transmitting the pathogen to susceptible species.

Post-treatment care for any crayfish that survives apparent infection focuses on supporting complete recovery while maintaining strict isolation. Recovered individuals should remain in quarantine for extended periods to confirm disease resolution and monitor for recurrence. Optimal nutrition supports immune function and tissue repair, with high-quality varied diet including protein sources and calcium for exoskeleton maintenance. Water quality must remain pristine, with regular testing and maintenance to prevent additional stress on recovering individuals. Any signs of relapse require immediate reassessment and likely indicate that recovery was not complete.

Prognosis factors influencing survival include species identity as the primary determinant, with North American species showing dramatically better outcomes than other geographic groups. Individual genetic variation in immune response affects disease resistance, explaining rare survival in predominantly susceptible populations. Stage at which intervention occurs impacts outcomes, though even very early detection rarely changes prognosis in highly susceptible species. General health status prior to infection, including nutritional condition and absence of concurrent health issues, may marginally influence survival but cannot overcome species-level susceptibility.

Long-term considerations for keepers following a crayfish plague outbreak extend well beyond the immediate health crisis. The affected system must be thoroughly disinfected before reuse, with all porous materials such as driftwood and substrate disposed of or sterilized. Equipment used with infected crayfish should either be discarded or treated with bleach solution followed by thorough rinsing and drying. Future stocking decisions should account for the demonstrated presence of the pathogen, with consideration of whether to continue keeping susceptible species or transition to North American species that can coexist with endemic pathogen presence. Stricter biosecurity protocols should be implemented to prevent future introductions.

Prevention

Proper husbandry forms the foundation of crayfish plague prevention, beginning with species selection appropriate to the keeper's situation and biosecurity capabilities. Keepers should carefully consider whether to maintain North American species, which may carry the pathogen asymptomatically, or susceptible species that face catastrophic mortality if exposed. Mixing these groups creates unacceptable disease risk and should never be attempted. Source verification for new crayfish acquisitions provides some protection, with captive-bred individuals from established disease-free colonies presenting lower risk than wild-caught specimens or those of unknown provenance. Maintaining detailed records of acquisition sources enables traceability if disease occurs.

Environmental control measures support overall health and disease resistance while minimizing conditions that might favor pathogen survival. Maintaining appropriate water temperature within species-specific optimal ranges supports immune function. Water quality management including regular testing and maintenance prevents stress that could increase disease susceptibility. Adequate space and hiding places reduce aggression-related stress and injuries that might provide pathogen entry points. Avoiding overcrowding reduces both stress and potential transmission rates should the pathogen be introduced.

Quarantine protocols represent the single most important preventive measure for captive crayfish keepers. All new acquisitions should undergo minimum thirty-day quarantine in completely separate systems with dedicated equipment before any possibility of contact with established collections. Extended quarantine periods of sixty to ninety days provide additional security, particularly for high-value collections or when acquiring crayfish from unknown sources. Quarantine systems should be maintained in separate rooms or locations from main collections where possible, with strict protocols preventing any water, equipment, or material transfer between quarantine and established systems.

Stress reduction supports immune function and overall disease resistance in crayfish populations. Providing appropriate environmental enrichment including hiding places, suitable substrate, and climbing structures reduces chronic stress. Maintaining consistent environmental parameters without sudden changes in temperature, lighting, or water chemistry prevents acute stress responses. Minimizing handling and disturbance reduces stress while also reducing opportunities for pathogen introduction through contaminated equipment. Appropriate feeding schedules and high-quality nutrition support optimal health and immune function.

Preventive monitoring enables early detection of potential disease introduction before catastrophic losses occur. Regular observation of all crayfish for behavioral changes or physical abnormalities allows rapid identification of problems. Maintaining baseline records of normal behavior for individuals enables detection of subtle changes that might indicate early infection. Any unexplained mortality should trigger immediate investigation and implementation of precautionary isolation measures. Periodic review of biosecurity protocols identifies potential weaknesses before they result in disease introduction. Staying informed about crayfish plague occurrence in the broader aquarium hobby and wild populations enables appropriate risk assessment for new acquisitions.

Living With & Managing Crayfish plague (Aphanomyces astaci) - devastating to native species

Enclosure maintenance for crayfish focuses on preventing pathogen introduction while providing optimal living conditions. Regular cleaning removes organic debris that could support pathogen survival outside of hosts, though excessive cleaning that disrupts biological filtration should be avoided. Equipment including nets, siphons, and maintenance tools should be dedicated to individual systems or thoroughly disinfected between uses. Any items added to enclosures, including decorations, plants, and substrate, should be quarantined or sterilized before introduction. Water source management ensures that incoming water does not introduce pathogens, with municipal water supplies generally safer than natural water sources that might contain wild crayfish or their associated pathogens.

Environmental parameters require careful maintenance within species-appropriate ranges to support health and disease resistance. Temperature should remain stable within the optimal range for the species kept, typically between eighteen and twenty-four degrees Celsius for most commonly kept crayfish species. Water hardness adequate for exoskeleton development and maintenance should be provided, with calcium supplementation if natural hardness is insufficient. pH stability prevents stress from fluctuations, with most crayfish preferring slightly alkaline conditions between seven and eight. Dissolved oxygen levels must remain adequate, particularly at warmer temperatures when oxygen solubility decreases.

Feeding and nutrition support immune function and overall health that may provide marginal benefits in resisting infection. Varied diet including high-quality commercial crayfish or crustacean foods provides balanced nutrition. Supplementation with blanched vegetables such as spinach, zucchini, and peas provides fiber and additional nutrients. Protein sources including fish, shrimp, and bloodworms support growth and tissue maintenance. Calcium supplementation through cuttlebone, mineral blocks, or calcium-rich foods supports exoskeleton health, which provides the first barrier against pathogen invasion. Avoiding overfeeding prevents water quality degradation and associated stress.

Handling considerations balance necessary husbandry tasks against stress and contamination risks. Minimizing handling reduces stress while also limiting opportunities for pathogen transfer between systems on keeper hands or equipment. When handling is necessary, wet hands or nets should be used to prevent damage to the exoskeleton. Hand washing between working with different systems prevents potential pathogen transfer. Moving crayfish between systems should be avoided unless absolutely necessary, and transferred individuals should undergo quarantine appropriate to the biosecurity status of source and destination systems.

Long-term health monitoring establishes patterns that enable early detection of developing problems. Regular observation sessions during feeding provide opportunities to assess behavior and physical condition of all individuals. Maintaining records of molting frequency, growth rates, and any observed abnormalities creates baseline data for comparison. Photography of individuals over time documents subtle changes that might otherwise go unnoticed. Establishing relationships with veterinarians or diagnostic laboratories familiar with invertebrate health enables rapid response if professional consultation becomes necessary. Participating in crayfish keeper communities provides access to collective experience and early warning of disease outbreaks affecting the hobby.

Species at Risk for Crayfish plague (Aphanomyces astaci) - devastating to native species

High-risk species for crayfish plague include all non-North American freshwater crayfish, with European species demonstrating extreme susceptibility. The noble crayfish (Astacus astacus) faces near-total mortality when exposed to the pathogen, having evolved without exposure to Aphanomyces astaci. White-clawed crayfish (Austropotamobius pallipes) and stone crayfish (Austropotamobius torrentium) similarly lack effective defenses and experience catastrophic population crashes following pathogen introduction. Turkish crayfish (Pontastacus leptodactylus) shows somewhat variable susceptibility but remains highly vulnerable compared to North American species. These European species have experienced dramatic population declines in the wild due to crayfish plague spread, and captive populations face identical vulnerability.

Australian and Asian crayfish demonstrate high sensitivity comparable to their European counterparts. Australian species including redclaw crayfish (Cherax quadricarinatus), yabby (Cherax destructor), and marron (Cherax cainii) experience severe disease when exposed despite their geographic isolation from the pathogen's origin. Various Asian crayfish species similarly lack evolutionary exposure to Aphanomyces astaci and demonstrate high mortality rates upon infection. The global aquarium trade creates constant risk of introducing the pathogen to previously unexposed populations of these species, necessitating strict biosecurity for keepers maintaining them.

Life stage considerations affect disease expression and mortality across all susceptible species. Recently molted crayfish with soft exoskeletons experience accelerated disease progression due to reduced physical barriers to pathogen invasion. Juvenile crayfish may succumb more rapidly than adults due to smaller body mass and less developed immune systems. Gravid females carrying eggs face particular risk, as infection typically prevents successful brood development. Stressed individuals from any life stage demonstrate reduced disease resistance, with factors including recent transport, inadequate nutrition, or suboptimal environmental conditions increasing vulnerability. North American crayfish species capable of carrying the pathogen asymptomatically present minimal clinical risk individually but enormous biosecurity risk when housed anywhere that susceptible species might eventually contact their water, equipment, or the carrier animals themselves.

Related Conditions

Commonly co-occurring conditions with crayfish plague often reflect the underlying stress and immune compromise that may have facilitated initial infection. Bacterial shell disease frequently accompanies or follows fungal infections, as damage to the exoskeleton provides entry points for bacterial pathogens. Secondary bacterial infections may complicate the clinical picture and contribute to mortality in cases where the primary plague infection might have been survivable. Parasitic infections including various protozoans may become more severe in immunocompromised individuals fighting plague infection. Nutritional deficiencies resulting from prolonged anorexia during illness can cause additional complications including soft shell syndrome and failed molts.

Conditions producing symptoms similar to crayfish plague require careful differentiation to ensure appropriate response. Saprolegnia and other fungal infections can produce white fuzzy growth on crayfish but typically progress more slowly and may respond to treatment. Bacterial shell disease causes discoloration and erosion of the exoskeleton but usually advances gradually rather than with the rapid progression characteristic of plague. Environmental stress from poor water quality can produce lethargy and behavioral changes mimicking early plague symptoms. Unsuccessful molts may result in death or weakness that could be confused with infectious disease. Careful observation of symptom progression, mortality patterns, and recent history of potential exposures helps distinguish between these conditions.

Complications arising from crayfish plague infection include immediate secondary infections and longer-term consequences for surviving individuals and affected systems. Bacterial invasion of tissues damaged by fungal infection accelerates decline and may affect organ systems not directly impacted by the primary pathogen. Failed molts in infected individuals leave crayfish vulnerable to desiccation, predation, and additional infections. Population-level consequences include complete collapse of susceptible species groups, potential spread to connected systems through shared equipment or water, and persistent environmental contamination with pathogen zoospores. Systems that have experienced plague outbreaks may harbor the pathogen indefinitely if North American carrier species are present, requiring permanent biosecurity modifications or complete system decontamination before susceptible species can be safely reintroduced.