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fibropapillomatosis of sea turtles

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fibropapillomatosis of sea turtles

Summary

  • Last modified
  • 03 January 2018
  • Datasheet Type(s)
  • Animal Disease
  • Preferred Scientific Name
  • fibropapillomatosis of sea turtles
  • Overview
  • Fibropapillomatosis (FP) is a disease of sea turtles that results in the production of tumours, both external and internal (i.e., visceral), that are considered benign, but may obstruct crucial functions, such as swim...

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Pictures

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PictureTitleCaptionCopyright
Hawaiian green turtle (Chelonia mydas) severely afflicted with Fibropapillomatosis (FP) tumours. The mouth tumors, which are unique to Hawaiian greens, can occur inside the mouth and throat, impairing both breathing and feeding.  Eye tumors impair vision and can blind the turtle. The large tumors around the flippers can impair swimming. Although FP tumors are benign, they can easily be a significant factor in a turtle's demise and result in death.
TitleHawaiian green turtle severely afflicted with Fibropapillomatosis (FP)
CaptionHawaiian green turtle (Chelonia mydas) severely afflicted with Fibropapillomatosis (FP) tumours. The mouth tumors, which are unique to Hawaiian greens, can occur inside the mouth and throat, impairing both breathing and feeding. Eye tumors impair vision and can blind the turtle. The large tumors around the flippers can impair swimming. Although FP tumors are benign, they can easily be a significant factor in a turtle's demise and result in death.
Copyright©Peter Bennett & Ursula Keuper-Bennett-1992 - CC BY 3.0
Hawaiian green turtle (Chelonia mydas) severely afflicted with Fibropapillomatosis (FP) tumours. The mouth tumors, which are unique to Hawaiian greens, can occur inside the mouth and throat, impairing both breathing and feeding.  Eye tumors impair vision and can blind the turtle. The large tumors around the flippers can impair swimming. Although FP tumors are benign, they can easily be a significant factor in a turtle's demise and result in death.
Hawaiian green turtle severely afflicted with Fibropapillomatosis (FP)Hawaiian green turtle (Chelonia mydas) severely afflicted with Fibropapillomatosis (FP) tumours. The mouth tumors, which are unique to Hawaiian greens, can occur inside the mouth and throat, impairing both breathing and feeding. Eye tumors impair vision and can blind the turtle. The large tumors around the flippers can impair swimming. Although FP tumors are benign, they can easily be a significant factor in a turtle's demise and result in death.©Peter Bennett & Ursula Keuper-Bennett-1992 - CC BY 3.0
Green sea turtle (Chelonia mydas) with significant Fibropapilloma (FP) tumors. Basking on a beach, north of Hale'iwa, Hawaii, USA.  November, 2011.
TitleGreen sea turtle with significant Fibropapilloma (FP) tumors
CaptionGreen sea turtle (Chelonia mydas) with significant Fibropapilloma (FP) tumors. Basking on a beach, north of Hale'iwa, Hawaii, USA. November, 2011.
Copyright©Andrew Danielson-2011 - CC BY-SA 3.0
Green sea turtle (Chelonia mydas) with significant Fibropapilloma (FP) tumors. Basking on a beach, north of Hale'iwa, Hawaii, USA.  November, 2011.
Green sea turtle with significant Fibropapilloma (FP) tumorsGreen sea turtle (Chelonia mydas) with significant Fibropapilloma (FP) tumors. Basking on a beach, north of Hale'iwa, Hawaii, USA. November, 2011.©Andrew Danielson-2011 - CC BY-SA 3.0

Identity

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Preferred Scientific Name

  • fibropapillomatosis of sea turtles

International Common Names

  • English: FP; green turtle fibropapillomatosis; marine turtle fibropapillomatosis; sea turtle virus

Overview

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Fibropapillomatosis (FP) is a disease of sea turtles that results in the production of tumours, both external and internal (i.e., visceral), that are considered benign, but may obstruct crucial functions, such as swimming, feeding, sight, and buoyancy, and can lead to death. It is presumed to be caused by Chelonid herpesvirus 5, although Koch’s postulates for proving the causal relationship have not yet been fulfilled due to the inability of the virus to grow in cell culture and the difficulties of experimentally infecting protected endangered species. It occurs most commonly in the green turtle (Chelonia mydas), which is currently listed as Endangered by the IUCN, but it has also been reported in the other six species of sea turtle, which (except for one for which there is insufficient data) are all classified as Vulnerable, Endangered or Critically Endangered.  It was first reported in the 1930s in green turtles in Florida; it now has a worldwide, circumtropical distribution and is considered a pandemic, with infection rates above 70% in some regions.

Host Animals

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Animal nameContextLife stageSystem
Caretta caretta (loggerhead sea turtle)Wild hostAquatic: All Stages
Chelonia mydas (green sea turtle)Domesticated host, Wild hostAquatic: All Stages
Dermochelys coriaceaWild hostAquatic: All Stages
Eretmochelys imbricata (hawksbill turtle)Wild hostAquatic: All Stages
Lepidochelys kempiiWild hostAquatic: All Stages
Lepidochelys olivacea (olive ridley sea turtle)Wild hostAquatic: All Stages
Natator depressusWild hostAquatic: All Stages

Hosts/Species Affected

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Sea turtles, along with other turtles and tortoises, are part of the order Testudines.  The seven living species include the green (Chelonia mydas), loggerhead (Caretta caretta), Kemp’s ridley (Lepidochelys kempii), hawksbill (Eretmochelys imbricata), flatback (Natator depressus), olive ridley (Lepidochelys olivacea), and the leatherback (Dermochelys coriacea).  All species are in the family Cheloniidae, except the leatherback which belongs to the family Dermochelyidae.  Fibropapillomatosis was first reported over 75 years ago in green turtles from Key West, Florida, USA (Smith and Coates, 1938).  Initially, it was thought that it was confined to green turtles, but in recent years it has been found in all other species; however, prevalence is still greatest in C. mydas (Aguirre and Lutz, 2004; Foley et al., 2005).

Distribution

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Marine turtle fibropapillomatosis was first reported in the 1930s in green turtles (Chelonia mydas) in Key West, Florida, USA (Smith and Coates, 1938). C. mydas is globally distributed in tropical and subtropical oceans, normally between latitudes 40° N and 40° S (Hirth, 1997), nests in more than 80 countries and is believed to inhabit coastal waters of over 140 countries (Groombridge and Luxmoore, 1989). Fibropapillomatosis now has a worldwide, circumtropical distribution (Aguirre and Lutz, 2004) in C. mydas and other hosts, and has been reported from every major ocean basin in which green turtles are found, particularly in near-shore habitats.

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Sea Areas

Atlantic, Eastern CentralPresentDuarte et al., 2012; Girard et al., 2013
Atlantic, SouthwestPresentRodenbusch et al., 2014
Atlantic, Western CentralPresentWilliams et al., 1994
Indian Ocean, EasternPresentHerbst, 1994
Indian Ocean, WesternPresentLeroux et al., 2010
Pacific, Eastern CentralPresentBalazs, 1991
Pacific, NorthwestPresentHerbst, 1994
Pacific, Western CentralPresentAdnyana et al., 1997

Asia

IndiaPresentSilas et al., 1984
IndonesiaPresentPresent based on regional distribution.
-Nusa TenggaraWidespreadAdnyana et al., 1997
JapanPresentHerbst, 1994Bonin Islands
MalaysiaPresentHerbst, 1994
PhilippinesPresentNalo-Ochona, 2000; Lucero et al., 2012
Sri LankaPresentHerbst, 1994

Africa

CongoPresentGirard et al., 2013
Equatorial GuineaPresentFormia et al., 2007
GabonPresentFormia et al., 2007
GambiaPresent, few occurrencesBarnett et al., 2004
MadagascarPresentLeroux et al., 2010
Sao Tome and PrincipePresentDuarte et al., 2012

North America

MexicoPresentQuackenbush et al., 1998
USAPresentPresent based on regional distribution.
-CaliforniaPresent, few occurrences1990McDonald and Dutton, 1990San Diego Bay
-FloridaWidespread1938Smith and Coates, 1938; Aguirre and Lutz, 2004; Ene et al., 2005
-HawaiiWidespreadlate 1940sBalazs, 1991; Williams and Bunkley-Williams, 1996; Chaloupka et al., 2009Disease rare until 1980s; in one well-studied population prevalence then increased rapidly until mid-1990s and has since declined again
-North CarolinaPresent, few occurrencesHarms et al., 2008
-TexasPresentTristan et al., 2010

Central America and Caribbean

Antigua and BarbudaPresentWilliams et al., 1994
BahamasPresentWilliams et al., 1994
BarbadosPresent1982-1983Gamache and Horrocks, 1992
BelizePresentHerbst, 1994
British Virgin IslandsPresentOvering, 1996
Cayman IslandsPresentHerbst, 1994
Costa RicaPresentHerbst, 1994
CubaPresentMoncada and Prieto, 2000
Dominican RepublicPresentWilliams et al., 1994
Netherlands AntillesPresentWilliams et al., 1994
NicaraguaPresentLagueux et al., 1998
PanamaPresentHerbst, 1994
Puerto RicoPresent1987Teas, 1991
Saint LuciaPresentJn Pierre, 2008
Turks and Caicos IslandsPresentRichardson et al., 2009
United States Virgin IslandsPresent1971Eliazar et al., 2000

South America

BrazilWidespreadRodenbusch et al., 2014
ColombiaPresentHerbst, 1994
UruguayPresentPastorino et al., 2008
VenezuelaPresentHerbst, 1994

Oceania

AustraliaPresentPresent based on regional distribution.
-QueenslandPresent1989Limpus and Miller, 1994Shoal Water Bay, Repulse Bay, Moreton Bay
-Western AustraliaPresent, few occurrences1996Raidal and Prince, 1996
Caroline IslandsPresentKolinski, 1994

Pathology

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Cutaneous fibropapillomas of green turtles (Chelonia mydas) are single to multiple raised masses, ranging from 0.1 cm to greater than 30 cm in diameter, that are often ulcerated and necrotic.  They are usually found on the soft skin, but may be found anywhere on the turtle’s body including the flippers, neck, chin, inguinal and axillary regions, and tail (Herbst, 1994).  The conjunctiva is a common site for the tumours (ocular fibropapillomas), and masses may grow over the cornea resulting in impairment of vision.  A significant percentage of animals with cutaneous fibropapillomas also have internal nodules.  For example, at a turtle hospital in Florida, approximately 17% of the green turtles with severe cutaneous fibropapillomatosis were also found to have internal tumours (Herbst, 1994), and in a survey of green turtles with FP from the Hawaiian Islands from 1993-2003, 39% had internal tumours (Work et al., 2004).  The organs most commonly affected appear to be the lungs and kidneys, but tumours also occur in other organs including the heart, gastrointestinal tract, and liver (Herbst, 1994; Work et al., 2004).  Green turtles with FP from Hawaii appear unique in that many have oral tumours, particularly of the glottis (Aguirre et al., 2002).

Cutaneous fibropapillomas can become large enough to interfere with locomotion and are easily entangled in fishing line.  Visceral fibromas can eventually disrupt normal organ functions and lead to death (Work et al., 2004).  Cardiac and kidney dysfunction, respiratory compromise, and gastrointestinal obstruction have all been identified as causes of death (Herbst, 1994).  Many green turtles with multiple cutaneous fibropapillomas become severely debilitated.  Clinical chemistry results of affected turtles confirm a general pattern of debilitation with hypoproteinaemia, electrolyte imbalances, uraemia, and elevation in liver enzymes (Herbst, 1994).  Capture and release studies show that the clinical course of the disease can be variable and prolonged and that some individuals may experience tumour regression and spontaneously recover from the disease (Chaloupka et al., 2009; Herbst, 1994; Machado Guimaraes et al., 2013).

Diagnosis

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Diagnosis of cutaneous fibropapillomatosis is made by gross pathological and histological examination of the tumours.  Visceral fibropapillomas can be detected by radiography, MRI, and/or endoscopy.  Several molecular polymerase chain reaction assays have been developed for the detection of Chelonid herpesvirus DNA from tumour tissues (Lu et al., 2000a; Lu et al., 2003; Quackenbush et al., 2001).  Recently, a sensitive PCR assay was developed and validated for detecting the virus in clinically healthy turtles, presumably representing latent infection (Alfaro-Nunez and Gilbert, 2014).  A serodiagnostic enzyme-linked immunosorbent assay to detect virus-specific antibodies has been developed; however, antibodies were detected only in samples collected after cutaneous fibropapillomas appeared (Herbst et al., 2008).  Virus isolation is not currently possible as the virus has remained refractory to in vitro cultivation (Work et al., 2009).

Disease Course

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Experimental infection studies of fibropapillomatosis (FP) have been hampered by the inability to grow in culture the virus presumed to be responsible for the disease, and by the protected status of sea turtles.  However, Herbst et al. (1995) successfully demonstrated experimental transmission of FP using cell-free extracts of tumours.  Fibropapillomas developed in all 12 turtles receiving tumour extracts and were first detected between 15 and 43 weeks post-inoculation.  This presumably reflects the incubation period of the causative virus.  Initiation of tumour growth was positively correlated with increasing water temperature.  Common sites for fibropapillomas are the flippers, neck, chin, inguinal and axillary regions, and the tail base.  Tumours can also appear on the conjunctiva and grow over the cornea affecting vision.  Hawaiian green turtles (Chelonia mydas) can have oral tumours, particularly in the glottis (Aguirre et al., 2002).  As many as 39% of turtles with external tumours also have internal tumours, most commonly in the lungs, kidneys, and heart (Herbst, 1994; Work et al., 2004).  In Hawaii, tumour regression has been observed in some turtles over an approximately 10-year period (Chaloupka et al., 2009), and this has also been observed in affected turtles from southeastern Brazil (Machado Guimaraes et al., 2013).  However, tumour regression appears to be uncommon. The immunosuppression associated with FP (Work and Balasz, 1999; Work et al., 2001) makes turtles prone to secondary bacterial infections (Work et al., 2003).

Epidemiology

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Marine turtle fibropapillomatosis (FP) was first reported in the 1930s in green turtles (Chelonia mydas) in Key West, Florida, USA (Smith and Coates, 1938).  Prevalence increased greatly in the decades that followed its initial description, especially in specific well-monitored areas such as Indian River Lagoon, Florida and Kaneohe Bay, Hawaii.  It now has a worldwide, circumtropical distribution and is considered a pandemic, with infection rates above 70% in certain regions (Aguirre and Lutz, 2004), while rates as high as 92% have been reported from Kaneohe Bay, Hawaii (Balazs, 1991).  Indeed it has been reported from every major ocean basin in which green turtles are found.  In general, there appears to be an association of high FP prevalence with near-shore marine habitats; however, large differences in prevalence among populations over short distances < 1 km) can also be observed (Herbst, 1994).  While sea turtle FP has been strongly linked to the presence of Chelonid herpesvirus 5, epidemiological studies and field observations support the notion that the prevalence of the disease is associated with heavily polluted coastal areas, areas of high human density, agricultural runoff, and/or biotoxin-producing algae (Aguirre and Lutz, 2004).  While still unclear, it is likely that these unfavourable environmental conditions predispose the turtles to viral infection (or viral reactivation); according to Ene et al. (2005), it is likely that turtles acquire the virus after returning to neritic habitat.  Encouragingly, however, recent decades have seen a steady decline in the number of green turtles with FP in Hawaii, with a prevalence rate of 9.4% recorded in 2007 in Pala’au State Park on the Hawaiian island of Molokai (Chaloupka et al., 2009).  Investigators have even observed regression of advanced FP in some turtles over an approximately 10-year period (Chaloupka et al., 2009), and this has also been observed in affected turtles from southeastern Brazil (Machado Guimaraes et al., 2013).  The reasons for this decline are purely speculative, but plausible explanations would include the development of herd immunity and/or removal of a tumour-inducing (or co-factor) environmental insult in the near shore habitat (Chaloupka et al., 2009). Unfortunately, unlike Hawaii, prevalence of FP in sea turtles elsewhere has continued to increase, or at best remained stable.

The mechanisms of FP transmission are unclear.  However, it is possible that the virus could be transmitted to uninfected individuals by direct contact with infected animals (Work et al., 2014) or by contact with substrates harboring the virus.  One recent study showed that a small percentage of FP tumours had lytic viral replication, a prerequisite for viral shedding.  Thus, it is possible that FP is maintained by superspreaders, a few highly infectious individuals that are able to transmit the virus via viral shedding (Work et al., 2014).  There is also evidence implicating the possible role of a mechanical vector in the transmission of FP.  Using quantitative PCR, Greenblatt et al. (2004) tested several ecto- and endoparasites of Hawaiian green turtles Chelonia mydas, including marine leeches, blood flukes, bladder parasites, barnacles, and amphipods of the skin and oral cavity.  Only marine leeches of the genus Ozobranchus were found to carry high viral DNA loads, with some samples containing as much as 1 x 107 DNA copies per leech, implicating them as a possible mechanical vector for the virus. Molecular evidence of the virus has also been found in a cleaner fish, suggesting that they too may act as vectors (Lu et al, 2000b).

Impact: Environmental

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Although fibropapillomatosis (FP) in sea turtles may have little environmental impact in and of itself, it is clear that there is a strong link between this disease and the environmental health of the coastal habitat (Santos et al., 2010, 2011).  Sea turtles are often considered sentinels of ecosystem health (Aguirre and Lutz, 2004).  In addition to infectious agents, they are particularly vulnerable to a variety of environmental insults such as high water temperature, pollutants, and marine biotoxins, all of which can impair their immune functions, making them more susceptible to a wide range of pathogens.  In fact, it has been suggested that the epidemiology of FP could serve as an effective tool to monitor ecosystem health in near-shore marine habitats (Aguirre and Lutz, 2004). 

Unfortunately, all sea turtle species are in danger of becoming extinct in the near future.  According to the IUCN Red List of Threatened Species, the hawksbill (Eretmochelys imbricata) and Kemp’s ridley (Lepidochelys kempii) turtles are listed as ‘critically endangered’; the loggerhead (Caretta caretta) and green (Chelonia mydas) turtles are listed as ‘endangered’; and the olive ridley (Lepidochelys olivacea) and leatherback (Dermochelys coriacea) turtles are listed as ‘vulnerable’ (IUCN, 2014).  The flatback turtle (Natator depressus) is listed as ‘data deficient’, meaning that its conservation status is unclear due to lack of data; it was previously listed as ‘vulnerable’ (IUCN, 2014).  There are many factors that contribute to the population decline of these animals.  Some of these factors include harvesting for food, the illegal sea turtle shell trade, ingestion and entanglement of marine debris, artificial lighting, beach erosion, invasive species predation, marine pollution including oil spills, and warming of the oceans due to climate change (Sea Turtle Conservancy, 2015).  FP and other diseases further add to the threats.

Having said this, studies in Hawaii to date have failed to show a link between disease and effect on somatic growth rates (Chaloupka and Balazs, 2005), or adverse effects on population growth (Chaloupka et al., 2007).

Zoonoses and Food Safety

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Fibropapillomatosis is not zoonotic.  However, the presence of the disease may suggest an ecological imbalance in the near-shore coastal environment that may directly or indirectly affect the human population.

Disease Treatment

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Surgical removal of the tumours is the most common treatment of affected turtles.  Unfortunately, surgical excision is associated with a high rate of disease recurrence and secondary bacterial infections.  Recently alternative therapies have been attempted.  For example, Sellera et al. (2014) successfully used photodynamic therapy to remove lesions.  For this procedure, the animal is given a systemic or local administration of a non-toxic photosensitizer - in this case, methylene blue.  Then, the photosensitized target tissue is irradiated by visible or near-infrared low-intensity light at a specific wavelength and time to ‘activate’ the methylene blue molecules.  In a trial of five affected green turtles (Chelonia mydas) that were treated twice at 15-day intervals, the treated lesions showed obvious tissue necrosis at 30 days post treatment, were totally or partially detached from the epidermis, and were easily removed using tweezers  (Sellera et al., 2014).  The potential for disease recurrence when this method is used is not known, however.  Another promising treatment option for fibropapillomatosis is electrochemotherapy (Brunner et al., 2014).  This technique uses a combination of chemotherapy, usually with bleomycin or cisplatin, and electroporation.  It consists of a series of short, high-voltage electric pulses that lead to increased membrane permeability and more efficient transport of antineoplastic drugs through the cellular membrane.  In a small study two green turtle fibropapillomas were treated in two sessions with a 33-day interval between sessions.  In both animals, complete regression of the lesions occurred without side effects or complications (Brunner et al., 2014).  Importantly, there was no sign of local recurrence, even one year after the end of treatment.

Prevention and Control

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Currently no prevention or control measures for wild populations of turtles are available, although the disease has been on a secular decline in Hawaii (Chaloupka et al., 2009), where turtles that strand with tumours are removed from the population (which is not the case in places where the disease persists).  Given the association with environmentally polluted, near-shore coastal regions, it would make sense to attempt to clean up these areas as much as practical (which would benefit coastal marine ecosystems as a whole).  In captivity, strict hygiene and quarantine procedures should be followed, including the use of separate water sources.  Newly acquired animals should be kept isolated for a minimum of 3 months and should undergo thorough physical examinations both before and after quarantine.  Preventive measures to reduce stress experienced by the animals in captivity may also help reduce mortality.  Reduction in the number of animals per tank, high water quality, and optimal water temperature will help to reduce the number of animals affected in captivity.

References

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Ackermann M; Koriabine M; Hartmann-Fritsch F; Jong PJ de; Lewis TD; Schetle N; Work TM; Dagenais J; Balazs GH; Leong JA, 2012. The genome of Chelonid herpesvirus 5 harbors atypical genes. PLoS One, 7(10):e46623.

Adnyana W; Ladds PW; Blair D, 1997. Observations of fibropapillomatosis in green turtles (Chelonia mydas) in Indonesia. Australian Veterinary Journal, 75(10):737-742.

Aguirre AA; Balazs GH; Spraker TR; Murakawa SKK; Zimmerman B, 2002. Pathology of oropharyngeal fibropapillomatosis in green turtles Chelonia mydas. Journal of Aquatic Animal Health, 14(4):298-304.

Aguirre AA; Lutz PL, 2004. Marine turtles as sentinels of ecosystem health: Is fibropapillomatosis an indicator? Ecohealth, 1:275-283.

Aguirre AA; Vasconcelos Perez J; Spraker TR; Hernandez Saldana P; Principe B; Albavera Padilla E; Lopez Reyes EM; Balazs GH, 2000. Studies of marine turtle fibropapillomatosis in Mexico: an international collaboration of research and training. In: Proceedings of the 20th Annual Symposium on Sea Turtle Biology and Conservation, Orlando, Florida, USA, 29 February-4 March 2000 [ed. by Mosier, A. \Foley, A. \Brost, B.]. Miami, Florida, USA: National Marine Fisheries Service, 50. [NOAA Technical Memorandum NMFS-SEFSC-477.] http://www.seaturtlesociety.com/proceedings/20turtle.pdf

Albareda DA; Garner M; Prosdocimi L; Rodriguez Heredia S; Paola JL di; Loureiro J, 2008. Pathological studies in green sea turtles (Chelonia mydas) and loggerhead sea turtles (Caretta caretta) from the northern coastal area of Buenos Aires, Argentina. In: Proceedings of the Twenty-Seventh Annual Symposium on Sea Turtle Biology and Conservation, Myrtle Beach, South Carolina, USA, 22-28 February 2007 [ed. by Rees, A. F. \Frick, M. \Panagopoulou, A. \Williams, K.]. Miami, Florida, USA: National Marine Fisheries Service, 4. [NOAA Technical Memorandum NMFS-SEFSC-569.] http://www.seaturtlesociety.com/proceedings/27turtle.pdf

Alfaro-Núñez A; Bertelsen MF; Bojesen AM; Rasmussen I; Zepeda-Mendoza L; Olsen MT; Gilbert MTP, 2014. Global distribution of Chelonid fibropapilloma-associated herpesvirus among clinically healthy sea turtles. BMC Evolutionary Biology, 14(206):(25 October 2014). http://www.biomedcentral.com/content/pdf/s12862-014-0206-z.pdf

Alfaro-Núñez A; Gilbert MTP, 2014. Validation of a sensitive PCR assay for the detection of Chelonid fibropapilloma-associated herpesvirus in latent turtle infections. Journal of Virological Methods, 206:38-41. http://www.sciencedirect.com/science/journal/01660934

Avens L; Goshe LR; Harms CA; Anderson ET; Hall AG; Cluse WM; Godfrey MH; Braun-Mcneill J; Stacy B; Bailey R; Lamont MM, 2012. Population characteristics, age structure, and growth dynamics of neritic juvenile green turtles in the northeastern Gulf of Mexico. Marine Ecology, Progress Series, 458:213-229. http://www.int-res.com/articles/meps_oa/m458p213.pdf

Balazs GH, 1985. Status and ecology of marine turtles at Johnston Atoll. Atoll Research Bulletin, 285:1-46.

Balazs GH, 1991. Current status of fibropapillomas in the Hawaiian green turtle, Chelonia mydas. In: Research plan for marine turtle fibropapilloma: Results of a December 1990 Workshop [ed. by Balazs, G. H. \Pooley, S. G.]., USA: U.S. Department of Commerce, National Oceanographic and Atmosperic Administration, National Marine Fisheries Service, 47-57. [NOAA-TM-NMFS-SWFSC-156.] http://swfsc.noaa.gov/publications/TM/SWFSC/NOAA-TM-NMFS-SWFSC-156.PDF

Balazs GH; Dudley WC; Hallacher LE; Coney JP; Koga SK, 1994. Ecology and cultural significance of turtles at Punalu'u, Hawaii. In: Proceedings of the Fourteenth Annual Symposium on Sea Turtle Biology and Conservation, Hilton Head, South Carolina, USA, 1-5 March 1994 [ed. by Bjorndal, K. A. \Bolten, A. B. \Johnson, D. A. \Eliazar, P. J.]. Miami, Florida, USA: National Marine Fisheries Service, 10-13. [NOAA Technical Memorandum NMFS-SEFSC-351.] http://www.seaturtlesociety.com/proceedings/14turtle.pdf

Balazs GH; Miya RK; Finn MA, 1993. Aspects of green turtles in their feeding, resting, and cleaning areas off Waikiki beach. In: Proceedings of the Thirteenth Annual Symposium on Sea Turtle Biology and Conservation, Jekyll Island, Georgia, USA, 23-27 February 1993 [ed. by Schroeder, B. A. \Witherington, B.]. Miami, Florida, USA: National Marine Fisheries Service. [NOAA Technical Memorandum NMFS-SEFSC-341.] http://www.seaturtlesociety.com/proceedings/13turtle.pdf

Balazs GH; Murakawa SKK; Ellis DM; Aguirre AA, 2000. Manifestation of fibropapillomatosis and rates of growth of green turtles at Kaneohe Bay in the Hawaiian Islands. In: Proceedings of the Eighteenth Annual Symposium on Sea Turtle Biology and Conservation, Mazatlán, Sinaloa, México, 3-7 March 1998 [ed. by Abreu-Grobois, F. A. \Briseño-Dueñas, R. \Márquez-Millán, R. \Sarti-Martínez, L.]. Miami, Florida, USA: National Marine Fisheries Service, 112-113. [NOAA Technical Memorandum MFS-SEFSC-436.] http://www.seaturtlesociety.com/proceedings/18turtle.pdf

Balazs GH; Pooley SG, 1991. Research Plan for Marine Turtle Fibropapilloma: Results of a December 1990 Workshop. USA: U.S. Department of Commerce, National Oceanographic and Atmosperic Administration, National Marine Fisheries Service, vi + 113 pp. [NOAA-TM-NMFS-SWFSC-156.] http://swfsc.noaa.gov/publications/TM/SWFSC/NOAA-TM-NMFS-SWFSC-156.PDF

Balazs GH; Rice M; Murakawa SKK; Watson G, 2000. Growth rates and residency of immature green turtles at Kiholo Bay, Hawaii. In: Proceedings of the Eighteenth Annual Symposium on Sea Turtle Biology and Conservation, Mazatlán, Sinaloa, México, 3-7 March 1998 [ed. by Abreu-Grobois, F. A. \Briseño-Dueñas, R. \Márquez-Millán, R. \Sarti-Martínez, L.]. Miami, Florida, USA: National Marine Fisheries Service, 283-285. [NOAA Technical Memorandum MFS-SEFSC-436.] http://www.seaturtlesociety.com/proceedings/18turtle.pdf

Balazs GH; Rice MR; Hoffman N; Murakawa SKK; Parker DM; Shallenberger RJ, 2005. Green turtle foraging and resting habitats at Midway Atoll: Significant findings over 25 years, 1975-2000. In: Proceedings of the Twenty-First Annual Symposium on Sea Turtle Biology and Conservation, Philadelphia, Pennsylvania, USA, 24-28 February 2001 [ed. by Coyne, M. S. \Clark, R.]. Miami, Florida, USA: National Marine Fisheries Service, 102-104. [NOAA Technical Memorandum NMFS-SEFSC-528.] http://www.seaturtlesociety.com/proceedings/21turtle.pdf

Ballestero J; Segura A, 1994. Observation on the incidence of five external lesion types in 506 Olive Ridley Lepidochelys olivacea (Eschscholtz) nesters in the Ostional Wildlife Refuge, Guanacaste, Costa Rica. In: Proceedings of the Fourteenth Annual Symposium on Sea Turtle Biology and Conservation, Hilton Head, South Carolina, USA, 1-5 March 1994 [ed. by Bjorndal, K. A. \Bolten, A. B. \Johnson, D. A. \Eliazar, P. J.]. Miami, Florida, USA: National Marine Fisheries Service, 14-16. [NOAA Technical Memorandum NMFS-SEFSC-351.] http://www.seaturtlesociety.com/proceedings/14turtle.pdf

Baptistotte C; Scalfoni JT; Gallo BMG; Santos AS dos; Castilhos JC de; Lima EHSM; Bellini C; Barata PCR, 2001. Prevalence of sea turtle fibropapillomatosis in Brazil. In: Proceedings of the Twenty-First Annual Symposium on Sea Turtle Biology and Conservation, Philadelphia, Pennsylvania, USA, 24-28 February 2001 [ed. by Coyne, M. S. \Clark, R.]. Miami, Florida, USA: National Marine Fisheries Service, 111-113. [NOAA Technical Memorandum NMFS-SEFSC-528.] http://www.seaturtlesociety.com/proceedings/21turtle.pdf

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Links to Websites

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WebsiteURLComment
Global Sea Turtle Networkhttp://www.seaturtle.org/
Sea Turtle Conservancyhttp://www.conserveturtles.org/

Organizations

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USA: Sea Turtle Conservancy (STC), 4424 NW 13th St, Suite B-11, Gainesville, FL 32609, http://www.seaturtle.org/

Contributors

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27/02/2015 Original text by:

Chris A. Whitehouse, U.S. Army Medical Research Institute of Infectious Diseases, 1425 Porter Street, Fort Detrick, Maryland 21702, USA

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