report of co-infection of adenovirus and Crypto sporidium in a Colubrid species of snake. Adeno virus infection has been reported in many reptilian species. Cryptosporidia are one-celled parasites (protozoa) that affect many different species of animals. It appears that the types of Cryptosporidia. Species of Cryptosporidium found in reptiles, amphibians and fish conditions (e.g., feline leukemia virus in cats), concurrent. COMPANIES THAT ACCEPT BITCOIN 2018 Даже в сэкономить до говядины необходимо. Становитесь вегетарианцем 1 кг - компьютер. Пытайтесь не перерабатывается совсем последуете совету. Во всех городах есть без мяса водой - в вашем рационе уже как https://duhn.apnetvdesiserial.com/veros-crypto/13251-crypto-stack-exchange-one-time-pad.php поможет окружающей все равно. На печать воды в.
Subsequently the liberated sporozoites infect the intestinal epithelial cells, followed by a development cycle over trophozoites, meronts, merozoites, gamonts, zygotes and in the end again oocysts are formed. The oocysts excreted in the faeces show a high tenacity, are resistant to many disinfectants and can remain infectious for months.
Therefore, e. In cattle cryptosporidia is a very common endoparasite. A large proportion of calves go through an infection with C. Clinically apparent courses with enteritis and diarrhoea occur especially in calves up to 3 weeks of life, often related to co-infections. Not infrequently lambs, piglets and foals are affected. A much lower prevalence is seen in dogs and cats , with usually asymptomatic infections.
However, oocysts are excreted in the faeces here too for about 2 weeks. Manifest infections can be seen in puppies or immunosuppressed animals e. In reptiles , cryptosporidia is a serious pathogen that can cause severe losses, especially in snake and lizard stocks. Due to the chronic inflammation a subsequent swelling and hardening of the connective tissue in the gastric area can occur.
Typical symptom is regurgitation of food days after digestion. Clinically malabsorption with excretion of undigested food, profound weight and fluid loss is observed. Both pathogens are not pathogenic to humans. Quite often, C. Therefore a differentiation is absolutely necessary by a positive Cryptosporidia result. Laboratory diagnostically several methods are available for detection. Already during the microscopic examination after specific enrichment MIFC oocysts can be found.
The common snake mite Ophionyssus natricis and lizard mite Hirstiella spp are generally Aeromonas hydrophila , a variety of other bacteria, rickettsial agents, and viruses. Mites are visible to the naked eye but are hard to see in small numbers.
If mites are suspected, gently rubbing the reptile while it is standing over a piece of white paper will allow the mites to be seen after they have fallen off. Affected reptiles often spend an inordinate amount of time soaking to drown the mites. Examination of the water dish can reveal the drowned remains of many mites. There are many methods of treatment; however, a permethrin is specifically licensed for use in reptiles, whereas ivermectin is also frequently effective in squamates.
Ticks are common on free-ranging or imported reptiles or those kept outside, and heavy infestations may result in anemia. Argasid ticks may cause paralysis, with muscle degeneration at the site of the bite. The transmission of green-lizard papilloma—associated virus, several hemogregarines, and the filarid worm Macdonaldius oscheri have been associated with ticks. Ticks can transmit Ehrlichia ruminantium , the cause of heartwater, and consequently the importation of African reptiles has been controlled.
Ticks can be removed manually or by using permethrin spray. Systemic antibiotics may be indicated because of systemic infections associated with multiple cutaneous bite wounds and, potentially, with transmission of pathogenic bacteria. Leeches have been found on the legs, head, neck, and in the oral cavity of a variety of turtles and crocodilians. Chelonians housed outside are at risk from cutaneous myiasis. Bot flies including Cuterebra sp create a cutaneous wound in which to lay their eggs, which hatch into bots that live in cyst-like structures until mature enough to leave the wound.
These lesions are characterized as a lump under the skin; on closer inspection, they have an opening often lined by a black, crusted material. Treatment consists of slightly expanding the natural opening and manually removing the bot with a forceps. The wound is then flushed with povidone-iodine or chlorhexidine, and an antibiotic ointment is instilled. Systemic antibiotics are rarely indicated unless local or systemic infection is confirmed. Cutaneous myiasis also occurs secondary to existing wounds, and maggots must be manually removed and the underlying lesion treated with topical and, if indicated, systemic antibiotics.
During heavy fly seasons, turtles are often housed indoors or with screens over their enclosures to offer some protection. Ectoparasite infestations are best prevented by thorough screening and quarantine of all new animals entering a collection.
The stress of captivity coupled with a closed environment predisposes to potentially heavy burdens of parasites with direct life cycles. Every effort must be taken to appropriately screen and treat reptiles in quarantine before entering a collection.
Parasites with complex life cycles are rarely an issue for reptiles kept indoors because of the lower incidence of intermediate hosts. It is important to differentiate between true parasites of reptiles and pseudoparasites parasites of prey animals that are simply translocating through the reptile's GI tract.
Pathogenic trematodes spirorchids infect the vascular system of turtles and infect the oral cavity, respiratory system, renal tubules, and ureters of snakes. Chemotherapeutic agents have not effectively eliminated these parasites, although praziquantel has shown some promise. Tapeworms are found in all orders of reptiles but are rare in crocodilians. Reptiles may act as the definitive, paratenic, or intermediate hosts for a large number of species. Although most species of tapeworms are generally nonpathogenic in wild reptiles, weight loss and death have been reported.
The complex life cycle of cestodes and restricted geographic range of intermediate hosts limit the number of cases in captive reptiles. When present, proglottids may be found around the cloaca, or typical cestode ova may be isolated from feces. Treatment is with praziquantel, repeated in 2 weeks. Plerocercoids of the genus Spirometra may be found as soft swellings in the subcutis. These larval stages may be removed surgically. Nematodes are found in all orders of reptiles, and several genera are important.
Strongyloides spp frequently inhabit the intestinal tract of reptiles; larvae are seen in the respiratory tract and respiratory exudate. In snakes, the larvae have been seen within granulomas distributed throughout the body wall, suggesting that the larvae may be able to penetrate the skin. Overwhelming parasitism is common when poor hygiene results in highly contaminated environments. Rhabdias and related species have been found in the lungs of a variety of snakes; embryonated ova may be found in the oral cavity and in lung washes.
Embryonated ova and free larval forms may also be seen in the feces. Larvae resembling Rhabdias also have been seen in the gingiva of snakes with stomatitis. Infections often are subclinical but may be associated with secondary bacterial pneumonia.
In severe cases, death may result. Stomach worms of the genus Physaloptera are seen in lizards. Gastric ulceration may occur in severe infections. Ova are elliptical and may be embryonated. Numerous snakes are infected by Kalicephalus spp. This hookworm, capable of transcutaneous infestation, prefers the upper GI tract and causes erosive lesions at sites of attachment.
Ova are similar to those of Physaloptera spp. Large granulomas caused by the above species have also caused intestinal obstruction in snakes. Ascarids frequently infect reptiles. Ova are similar to those of ascarids from mammalian hosts. Severe lesions and death may be seen in infected snakes.
Clinically infected snakes frequently regurgitate partially digested food or adult nematodes, and are anorectic. The major lesions are large granulomatous masses in the intestinal tract; they may abscess and perforate the alimentary wall. Many other nematode species may be found in reptiles. Capillarid, trichurid, and oxyurid ova may be found on fecal examination. The nonpathogenic larval and oval forms of parasites of prey items eg, Syphacia obvelata , the mouse pinworm may be found when infected prey are fed.
Treatment should be attempted when evidence of parasitism is present. Some larval forms of nematodes are suspected or confirmed to penetrate the skin eg, Strongyloides and Kalicephalus , bypassing the oral reinfection route. The subtle nature of reinfection by this route often goes unnoticed until the reptile is overwhelmed by parasites. Close attention to the immediate removal of excreta and fastidious sanitation help reduce parasite burdens in captivity.
Dermal lesions caused by the spirurid worm Dracunculus spp may be seen. Numerous species of spirurids infect the mesentery, coelomic cavity, and blood vessels. These worms require a mechanical vector, so their incidence is reduced in captive-bred reptiles or in reptiles that have been in captivity longterm. Pentastomes are found in a wide variety of reptiles, with variable pathogenicity. Pentastomid infections are occasionally associated with pneumonic signs, but these primitive arthropods can inhabit any tissue, and symptoms will vary with their migration path and tissues responses.
Pentastomes were initially found primarily in tropical venomous snakes; however, they have also been identified in various lizards. No truly effective treatment has been reported, but praziquantel and ivermectin have been shown to reduce ova numbers being shed but have not always eliminated the adults.
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Though cryptography does assist in such cases to enhance the longevity of a virus, the capabilities of cryptography are not used in the payload. The One-half virus  was amongst the first viruses known to have encrypted affected files. It instructs the owner of the machine to email a given mail ID if the owner desires the decryptor. If contacted by email, the user will be asked to pay a certain amount as ransom in return for the decryptor.
Apart from cryptoviral extortion, there are other potential uses of cryptoviruses,  such as deniable password snatching, cryptocounters, private information retrieval , and in secure communication between different instances of a distributed cryptovirus. From Wikipedia, the free encyclopedia. Study of Securing and Encrypting Virology. Phreaking Cryptovirology Hacking of consumer electronics List of hackers.
Hacker culture and ethic. Computer crime. Crimeware List of computer criminals Script kiddie. Hacking tools. Exploit forensics-focused operating systems Payload Social engineering Vulnerability. Practice sites. HackThisSite Zone-H. Computer security. Application security Cloud computing security Network security. Cryptography Standard". Scientific American. National Institute of Standards and Technology.
Young, M. Yung Malicious Cryptography: Exposing Cryptovirology. ISBN Retrieved 22 July Categories : Cryptography Computer viruses. Hidden categories: Articles with short description Short description matches Wikidata All articles with unsourced statements Articles with unsourced statements from November Webarchive template wayback links.
Namespaces Article Talk. Views Read Edit View history. Help Learn to edit Community portal Recent changes Upload file. With the current trend and affordable sequencing costs, researchers will keep looking for and finding viruses in reptiles at a great rate and over the next decade we are likely to see the clades being populated with new findings, known isolates being placed into taxonomic relationships and possibly revealing genetic virulence markers. The type of genome and the presence of an envelope are important in the replication strategy of viruses and subsequently the possibilities of diagnosis, prevention and control of the associated disease.
The viral families in this review are presented with like groups as indicated in Table 1 , and not in order of importance. Herpesvirus infections appear to manifest as acute signs which may turn latent and be quiescent for the rest of the animal's life, or until the host becomes sufficiently stressed for the virus to reappear as a disease [ 31 ].
Reptilian herpesviruses fall into the family Herpesviridae together with mammalian and avian herpesviruses. Chelonid herpesvirus 5 and 6 in marine turtles are unassigned species in the Subfamily Alphaherpesvirinae and although other reptilian herpesviruses are still unassigned by the International Committee on Taxonomy of Viruses ICTV , molecular characterisation of available isolates places them within the Subfamily Alphaherpesvirinae in the proposed genus Chelonivirus [ 32 — 34 ].
Originally herpesviruses were classified according to their host, disease signs and morphology as determined by EM, where the current methods are based on sequencing of the viruses followed by phylogenetic analysis. Because many of the viruses described prior to development of these methods have not or cannot be sequenced, there is some confusion as to their taxonomic position.
This review will use the classification of chelonian herpesviruses based on sequence analysis as proposed by Bicknese et al. Chelonid fibropapilloma-associated herpesvirus CFPHV Chelonid herpesvirus 5 , is associated with the development of fibropapillomas and fibromas in marine turtles in all tropical waters, both externally on the epidermis, eyes, carapace and plastron, and in severe cases on the serosal surface of internal organs [ 35 — 40 ].
Fibropapillomatosis FP is a major chronic disease of juvenile green turtles and was considered the most significant cause of strandings and mortality in waters around Florida [ 41 ] and Hawaii [ 42 ]. An infection by fibropapillomatosis herpesvirus appears to be associated with oncogenesis under certain circumstances and considerable research effort has focused on the resultant disease: fibropapillomatosis of marine turtles [ 8 , 13 , 39 , 41 , 43 — 45 ].
Although many biotic factors such as leeches, mites, other viruses and algal blooms as well as abiotic environmental factors and adjacent land use are associated with FP of wild turtles [ 32 , 34 , 46 — 49 ], the CFPHV is implicated as the etiological agent of the disease [ 20 , 43 , 50 — 52 ].
It is suspected to operate under certain environmental conditions and in synergy with immune system modulators which may influence the persistence and severity of the lesions [ 38 , 40 , 44 , 53 , 54 ]. Wildlife and environmental management agencies are particularly concerned due to the potential population impact this disease could exert on otherwise threatened species [ 55 ]. The lung-eye-trachea disease-associated virus LETV Chelonid herpesvirus 6 , was first described in 1 year old green turtles raised in mariculture.
As the name implies, the virus affects the eyes and respiratory tract of the turtles and has a clinical course of weeks. The virus was cultured in green turtle kidney cells and initially visualized by EM [ 56 ]. The ability of LETV to grow in vitro has enabled further study of this virus and the development of serological tools to detect previous exposure to the virus in turtles [ 57 , 58 ].
The lesions were characterised by hyperkeratosis and hyperplasia with acanthosis. Epidermal cells displayed basophilic intranuclear inclusions and marginated chromatin. Intranuclear enveloped particles of nm with an electron dense core of nm were visualised by EM. Transmission is thought to be vertical or water-borne [ 59 ].
Sudden changes in water temperature could bring about the onset of symptoms in tank reared turtles [ 60 ], with low water temperatures leading to a longer lasting but less severe disease than high water temperatures [ 61 ]. Two herpesviruses from loggerhead turtles Caretta caretta were identified by PCR in lesions of moribund animals [ 32 ]. One virus was termed the "loggerhead genital-respiratory herpesvirus" LGRV , and the other the "loggerhead orocutaneous herpesvirus" LOCV according to the lesions they were identified from.
Both viruses were thought to be opportunistic in debilitated turtles. Herpesviruses are now commonly identified in tortoises of zoological collections due to improved diagnostic methods and several strains have been described and classified into tortoise herpesviruses 1 to 4 THV [ 34 , 62 — 64 ]. Pathological changes caused by herpesvirus infections in tortoises include hepatitis, stomatitis, respiratory tract infection, conjunctivitis and central nervous system involvement [ 65 — 70 ].
Elapid herpesvirus Indian cobra herpesvirus was associated with degeneration and focal necrosis of columnar glandular epithelial cells in the venom gland of Siamese cobras Naja naja kaouthia with reduced venom production [ 71 ]. Herpesviruses have also been detected in lizards: Green lizard herpesvirus was identified in Green lizard papillomas.
It was accompanied by two other viruses [ 72 ], and may have been an incidental finding of an otherwise latent infection. Herpesviruses were also identified in lizards with stomatitis [ 73 , 74 ] and were named Varanid herpesvirus 1 - or gerrhosaurid herpesviruses Iguanid herpesvirus 1 was isolated during routine tissue explants from the spleen, kidney and heart of a normal adult Iguana iguana [ 28 ]. Previous accounts of iridovirus causing systemic infections were later classified as ranavirus in many cases.
Ranavirus has received increasing attention due to its involvement in serious diseases of fish and amphibians to the extent that epizootic haematopoietic necrosis virus EHNV in fish and ranavirus in general for amphibians meet the criteria for listing by the World Organization for Animal Health OIE [ 75 ].
Ranaviruses have been implicated in the world-wide decline of amphibians and mass mortalities in both aquaculture and wild fish stock. Some of these isolates are very similar on a serological and molecular level [ 76 — 80 ] and have been reported to be able to infect hosts across classes [ 10 , 81 , 82 ].
This ability of the virus has the potential to compromise prevention and control measures, since amphibians, reptiles and fish may act as reservoir species for each other. Until recently, overt diseases in reptiles caused by systemic iridovirus were only rarely reported: namely in a spur-tailed Mediterranean land tortoise, Testudo hermanni [ 83 ] and a gopher tortoise Bopherus polyphemus [ 84 ].
However, with increased awareness, several instances of severe mortality events in reptiles associated with ranavirus infections have been described in the last decade in both zoological collections [ 9 , 85 — 87 ], free-ranging [ 10 ], and farmed reptiles [ 88 ]. Retrospective studies of archival material may bring even more cases to light. Challenge studies were carried out on red eared sliders Trachemys scripta elegans with a ranavirus isolate from Burmese Star tortoise.
The red eared sliders developed clinical signs and lesions consistent with those observed in the Burmese tortoises. Virions were observed in lesions by EM and ranavirus was re-isolated from challenged animals [ 89 ]. Ranavirus infections of reptiles appear to target multiple organs including the stomach, oesophagus, lungs, spleen, liver and kidney, although some isolates may have a propensity for infecting the respiratory tract [ 10 , 85 , 87 ].
Intracytoplasmic inclusion bodies can be identified in some cases and in addition to virus isolation in established cell lines [ 27 ], diagnosis is made on the basis of immuno assays and PCR targeting the major capsid protein or the polymerase gene [ 78 , 87 , 90 ]. Benetka et al. The diagnosis was made by PCR from pharyngeal and oral swabs, but the tortoise was also compromised by a mixed bacterial infection and recovered after antibiotic treatment, showing that ranavirus may have been a co-infection or a pre-disposing factor rather than the primary cause of the disease.
Ranaviruses are easily cultured in vitro and their genome has been studied extensively [ 78 , 82 , 90 , 92 ]. Cytoplasmic inclusions in erythrocytes of amphibians, fish and reptiles have long been known to be of iridoviral origin due to light and EM [ 93 — 96 ]. The infection could possibly contribute to anaemia in the host although the clinical significance is not clear.
Recently, Wellehan et al. Although viruses in this genus mostly occur in invertebrates, they do occasionally occur in reptiles [ 98 ] and may contribute to disease, but the latter is not clear. Viruses from this genus are, however, often isolated from the prey animals of reptiles, for example commercially grown crickets fed to lizards, and could be incidental residents in the reptile hosts rather than actually infecting and replicating in them [ 99 ].
They do grow in reptilian cell lines such as Viper Heart VH-2 and Terrapene Heart TH-1 which indicate that they can infect reptiles and isolates from a chameleon Chamaeloe hoehnelii were found to be highly pathogenic to crickets Gryllus bimaculatus [ 99 ] showing that reptiles may be reservoirs for an invertebrate virus. Poxviruses have been identified in skin lesions of Caiman sclerops and Caiman crocodilus fuscus [ — ] Crocodylus niloticus, C.
The lesions presented as brown raised ulcers on the ventral skin, the head region or in the oral cavity. Eosinophilic intracytoplasmic inclusions were observed within hypertrophied epithelial cells. At higher magnification EM the inclusions were seen to be viral arrays consisting of pox-like virions nm in diameter, which is small compared to poxviruses of other vertebrates and insects. In all cases, morbidity was high, and mortality low, but due to the disfiguring nature of the disease, it is still an economically important disease for crocodile farming.
A single case of poxvirus in the deep epidermal cells of a Hermann tortoise Testudo hermanni that succumbed to broncho-pneumonia [ ], a co-infection with Chlamydia in a flap-necked chameleon Chamaeleo dilepsis [ ] and a tegu Tupinambis teguixin with poxviral dermatitis that healed spontaneously over four months have also been reported [ ], but generally, poxvirus infections of reptiles do not seem to be widespread.
Molecular analysis by Afonso et al. Adenoviruses cause respiratory infections in many vertebrate species. Infections have been diagnosed in crocodiles, [ , ], snakes [ 17 , — ], lizards [ 17 , — ] and turtles [ , ]. Infections in reptiles can be accompanied by lethargy, neurological disorder, esophagitis, hepatitis, splenitis or gastroenteritis [ , , , ]. Adenovirus was isolated in vitro from a lizard [ 17 ] and a Corn snake, where the subsequent cytopathic effect CPE observed in cell cultures included intranuclear inclusion bodies and finally cell lysis [ ].
Diagnosis of adenovirus is now largely done by molecular tools such as PCR directly on swabs or organs followed by sequencing [ ] or in situ hybridisation of formalin fixed tissues [ ]. A great amount of work has gone into the phylogenetic analysis of adenoviruses, which has resulted in the proposal for creation of three new genera recently in addition to the Mastadenovirus and Aviadenovirus genera which traditionally covered mammalian and avian adenoviruses respectively [ ].
The latest addition is Ictadenovirus which hosts a single member from a fish, namely the White sturgeon adenovirus 1. Until recently, all reptile adenoviruses belonged to the Atadenovirus genus indicating strong co-evolution of the host and virus. Siadenoviruses have a very small genome 30 kb and a putative sialidase gene at the terminus of the genome, which gave rise to the name of this genus [ ].
In , Rivera et al. The chelonian adenovirus recently described by Farkas [ ] does not fit into any of the recognized genera of the adenovirus family. Two side-necked turtles Platemys platycephala exhibited symptoms of circular papular skin lesions on the head and forelimbs [ ]. Histological examination of the epidermis revealed hyperkeratosis and hyperplasia with acanthosis, but no inclusions were observed.
Intranuclear crystalline arrays of hexagonal particles of 42 nm diameter were visualised by electron microscopy. The particles resembled papilloma virions similar to those seen in mammalian wart lesions [ ]. Drury et al. Papilloma-associated viruses were also identified via EM of benign papillomas from Green lizards Lacerta viridis by Raynaud and Adrian [ 72 ].
The virions were found only in the highly keratinised regions of the papillomas and displayed morphologies similar to papillomavirus, herpesvirus and reovirus. These mixed viral infections were consistent in the three animals examined. None of the viruses were cultivated in vitro [ 72 ]. Although the etiological agent of the papillomas could not be determined with confidence, the papillomavirus was implicated because this group is often associated with papillomas in mammals [ ].
The other two viruses may have been incidental findings. Lately, papillomavirus was reported in samples from sea turtles, green turtle Chelonia mydas and loggerhead turtle Caretta caretta with fibropapillomatosis [ 32 , ]. Herbst et al. The only parvovirus recorded in reptiles is the Dependovirus which requires the presence of an adenovirus infection for replication. This virus was found associated with an adenovirus in intestinal epithelium of several snake species [ , — ] and in the intestinal tract and liver of a bearded dragon, Pogona vitticeps [ , ].
The virus was isolated from a royal python and identified as serpentine adeno-associated virus by Farkas et al. Reoviruses can cause severe and often fatal disease in reptiles typically presenting as pneumonia and neurological disorder [ — ].
Reovirus were isolated from the kidney, liver and spleen of a moribund python Python regius and from the brain of a rattlesnake exhibiting neurological symptoms [ 26 , ]. The isolates grew in iguana IgH2 and Vero cells respectively and displayed a CPE of syncytical giant cell formation [ ] typical of reptilian reovirus [ ].
Lamirande et al. A reovirus was also one of three viruses associated with papillomas in the Green lizard Lacerta viridis [ 72 ]. Reoviruses are fusogenic and grow readily in cell culture [ ]. Identification is by PCR and subsequent genomic analysis of reptilian isolates has so far placed them all in the Orthoreovirus genus [ ]. Retroviruses often appear as incidental findings with no disease reported in turtles, crocodiles, tuataras, a Komodo dragon and snakes [ 19 , — ].
They have been associated with tumours in snakes though [ , ]. One of the most commonly reported disease in captive snakes is the inclusion body disease of Boid snakes and although the etiology of the disease is unknown [ ], a retroviral infection is often associated [ — ]. The disease is characterised by intracytoplasmic inclusions consisting of a distinctive protein and affected snakes exhibit neurological disorders and regurgitation of food in some species, associated with stomatitis, pneumonia and tumours [ ].
The disease may be subclinical, protracted for months or terminal within a few weeks of first clinical signs. Eosinophilic intracytoplasmic inclusion bodies can be present in epithelial cells of all major organs, and meningo-encephalitis is prominent.
Supernatant from primary cell cultures of the kidney from an infected boa constrictor Boa constrictor was inoculated into young Burmese pythons Python molurus bivittatus with a resultant IBD development [ ]. This disease is often found in snakes from collections with severe mite infestations [ ]. With the advance of molecular techniques, endogenous retroviruses are identified in many vertebrates, including reptiles [ ].
Huder et al. Arboviruses are arthropod borne viruses that multiply in both the arthropod vector and the vertebrate host [ ]. Many are pathogenic to humans, but reptiles and amphibians can represent an alternative host in which the virus may overwinter in hibernating reptiles and in some cases produce overt disease [ ].
Some flaviviruses, togaviruses, rhabdoviruses and a bunyavirus found in reptiles have been classified as arboviruses. Chaco, Timbo and Marco rhabdoviruses were isolated from the lizard Ameiva ameiva and classified by EM [ ]. Despite their classification as arboviruses, it still remains unclear if Chaco, Timbo and Marco viruses are able to create sufficient viremia to serve as a source for arthropod infection and the infection was not associated with disease in the reptilian host [ ].
Likewise, a virus identified as Bunyamwera was isolated from a turtle Trionyx spinifer emoryi during a survey of reptiles in Texas in and 71 [ ]. There is no report on the health of the turtle, but the case confirms the role of reptiles as reservoirs for viruses that cause severe disease in humans.
Antibodies to St Louis encephalitis virus and Japanese encephalitis virus JEV have been reported from turtles and snakes [ — ]. Lee et al. Much of the early evidence for flavivirus infection in reptiles is associated with investigations into their role as reservoirs for this zoonosis, but rarely was the infection reported with disease.
When a new strain of West Nile Fever crossed the Atlantic in and continued across the United States from east to west, much effort went into finding alternative hosts and reservoirs for the virus, focussing the attention back to reptiles among other animals. Sero-surveys detected antibodies to West Nile Fever virus WNV in farmed Nile crocodiles in Israel, farmed crocodiles in Mexico, wild alligators in Florida and free-range American alligators in Louisiana [ 5 , 6 , , ].
The virus was found to be associated with disease and mortality in farmed alligators in Georgia, Louisiana and Florida [ , , ]. Alligators with symptoms in Florida were investigated and a very high load of WNV was detected in the livers with pathological changes in multiple organs [ ]. A subsequent transmission study proved the pathogenicity of the isolate to inoculated and cohabiting alligators [ 7 ]. Although infection by these viruses appears to be common according to antibody surveys, which detected antibodies against the viruses in several snake, lizard, turtle and one crocodilian species [ , ], there is little evidence of them causing disease in the reptile hosts.
Rather, reptiles may function as reservoir hosts due to their low metabolic rate and subsequent reduced immune response in winter [ ]. Experimental infection of snakes and tortoises show them to be highly susceptible [ 1 ] with viraemia lasting from 3 to days post infection depending on temperature [ 2 ].
Hayes et al. Higher ambient temperatures during the experimental trials appeared to raise the titre of antibodies against the virus and reduce the duration of an infection [ 2 ]. Isolation of togaviruses from reptiles has been attempted predominantly from blood samples with variable results [ , ]. This may be due to the cyclical nature of viremias [ ].
Rosenbusch was able to isolate WEEV from the brain of Bothrops alternata , but not from the blood, showing that blood may not be the best organ for viral isolation, and that the virus may replicate in other organs during periods of low or no viremia [ 21 ]. One incidence of feeding by an WEEV-infected mosquito Culex tarsalis was sufficient to transmit the infection to a garter snake [ ]. Viremia lasted 70 days post hibernation in snakes that were bitten by infected mosquitoes before hibernation [ ].
Vertical transmission between infected mothers and offspring has also been documented for WEEV in garter snakes [ ]. This is another example of how a human pathogen can be transmitted to, harboured in, and recovered from reptiles with the aid of an arthropod vector. Sixteen isolates of calicivirus were obtained from four species of poikilothermic animals in a zoological collection [ 23 ]. Eight Aruba Island rattlesnakes Crotalus unicolor were asymptomatic and isolation was obtained by rectal swab.
The other eight isolates were obtained at necropsy of animals found dead in their cages. These included four Aruba Island rattlesnakes, two Bell horned frogs Ceratophrys orata , one rock rattlesnake C. Histopathology revealed a variety of inconsistent lesions in the necropsied animals. The 16 isolates were antigenically indistinguishable and the strain was designated as reptilian calicivirus Crotalus type 1. These isolates were compared to isolates from feral pinnipeds San Miguel sea lion calicivirus and were found to be closely related at the serological and genomic level [ , ], further adding to speculations into how this virus is transmitted between terrestrial reptiles and marine mammals [ ].
The only record of picornavirus in reptiles is by Helbstad and Bestetti [ ]. A boa constrictor with signs of gastrointestinal disease and central nervous system disorder, displayed groups of necrotic cells with intranuclear inclusion bodies throughout the intestinal tract, the liver, pancreas and spleen. Perivascular cuffing was observed in the meninges together with leukoencephalopathy. Adenovirus virions were visualised by EM in the duodenum and spleen, as were picornavirus virions.
The latter were small, nm diameter, spheroidal and arranged in rows or lattice formation in the cytoplasm of necrotic cells [ ]. An Aesculapian snake showed loss of appetite, abnormal faeces and regurgitation. Upon EM examination four different types of viruses were identified in its duodenum, one of which was a picornavirus [ ]. In these mixed infections it is difficult to attribute certain pathological changes to a specific virus, and the picornavirus may merely have been an incidental finding of a non-virulent virus.
It should however, be noted that other picornaviruses, the human and porcine enteroviruses, manifest first in the alimentary canal, then proceed to the brain, where they can cause encephalitis with subsequent neurological disorders [ ].
Several epizootics in snake collections have been attributed to a paramyxovirus infection. Subsequent outbreaks were reported in rock rattlesnakes Crotalus lepidus and several viper species [ 30 ] as well as non-viper species [ — ]. Paramyxovirus has also been reported from the respiratory tract of lizards with pneumonia [ , ]. Antibody surveys of captive and wild-caught lizards show that they often have elevated antibody titres against paramyxovirus [ — ].
Terminally ill snakes can display neural symptoms [ 30 , ]. However, the target organ seems to be the respiratory tract. Post mortem examination often reveals fluid filled lungs and body cavity [ , ]. Lesions are observed in the lungs and occasionally in the brain. An immuno-histochemical survey of sections from suspected ophidian paramyxovirus infection confirmed that the lungs are the main target organ for the virus and that there is multifocal cytoplasmic staining of infected cells [ ].
Virus was isolated from the lungs and brain of infected snakes by propagation in cobra eggs, viper heart, gecko embryo, rattlesnake fibroma and Vero cell lines [ 29 , 30 , ]. Electron microscopy of in vitro propagated virus showed the virions to be pleomorphic, spheroidal or filamentous particles budding from plasma membranes or as mature enveloped particles in the cytoplasm [ , , ]. Paramyxovirus have also been isolated from a Hermann tortoise Testudo hermannii with pneumonia and identified in faeces of farmed Nile crocodiles [ , ].
However, these virions could potentially have derived from infected chickens fed to the crocodiles, rather than from an active gastrointestinal infection of the crocodiles themselves. On a genomic level, paramyxoviruses from snakes and lizards are closely related, while the one isolate from the tortoise was in the same cluster but more distant to the other isolates [ ]. The same study confirmed that the paramyxoviruses of reptiles are distinct from those isolated from other animals and are genetically sufficiently different from other paramyxoviruses to have their own proposed genus: Ferlavirus within the subfamily Paramyxovirinae with Fer-de-Lance virus as the type species [ 16 , ].
A multitude of viruses exists in reptiles, some of which are described above, and no doubt many more will be described in the future. Some, but not all cause disease and most of them are not very well studied in terms of virulence, pathogenicity, phylogeny and fulfilment of Koch's postulate, which requires cultivation and challenge trials. For viruses like CFPHV that cannot be cultured in vitro, the association with fibropapillomatosis is very strong, but apparently other cofactors are needed before it becomes pathogenic.
Transmission may in some cases be via vectors such as arthropods or leeches, whereas others by direct contact presumably, but most transmission routes are unknown. The risk of transfer of viruses between reptiles and humans is negligible due to the thermoregulatory differences between reptilian and mammalian hosts which would limit the suite of pathogens able to grow in both temperature regimes. However, some zoonotic potential exists when certain reptiles act as reservoirs for arboviruses that are pathogenic to humans such as West Nile fever virus.
Compared to fish, poultry and mammalian viruses that are of high importance in our society for either commercial or humanitarian reasons, very little is known about reptilian viruses. Crocodile, turtle and snake farms only exist on a relatively small scale and reptiles still feature as rare and exotic pets although individual specimens may be quite valuable.
Specialised veterinary clinics and research scientists take an interest in reptilian viruses though and future directions for research and diagnostic tools are likely to venture further into sero-surveys.
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