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Chelonid herpesvirus 5

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Datasheet

Chelonid herpesvirus 5

Summary

  • Last modified
  • 14 July 2018
  • Datasheet Type(s)
  • Invasive Species
  • Preferred Scientific Name
  • Chelonid herpesvirus 5
  • Taxonomic Tree
  • Domain: Virus
  •   Unknown: "ssDNA viruses"
  •     Unknown: "DNA viruses"
  •       Order: Herpesvirales
  •         Family: Herpesviridae
  • Summary of Invasiveness
  • Chelonid herpesvirus 5, also known as fibropapilloma-associated marine turtle herpesvirus, is a herpesvirus that is presumed to be the cause of fibropapillomatosis in sea turtles, although Koch’s postulates for provin...

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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

  • Chelonid herpesvirus 5

Other Scientific Names

  • Chelonid fibropapilloma-associated herpesvirus
  • fibropapilloma-associated marine turtle herpesvirus

Summary of Invasiveness

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Chelonid herpesvirus 5, also known as fibropapilloma-associated marine turtle herpesvirus, is a herpesvirus that is presumed to be the cause of fibropapillomatosis in sea turtles, 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. The disease takes the form of internal and external tumours, which although not malignant can obstruct crucial functions. It was first reported in Florida in the 1930s, and although the mode of transmission is unclear, it now has a worldwide circumtropical distribution, with infection rates greater than 70% in some regions. It most commonly infects green turtles (Chelonia mydas), classified by the IUCN as Endangered, but it has also been reported in the other six sea turtle species, which (except for one for which there is insufficient data) are all classified as Vulnerable, Endangered or Critically Endangered.

Taxonomic Tree

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  • Domain: Virus
  •     Unknown: "ssDNA viruses"
  •         Unknown: "DNA viruses"
  •             Order: Herpesvirales
  •                 Family: Herpesviridae
  •                     Subfamily: Alphaherpesvirinae
  •                         Species: Chelonid herpesvirus 5

Notes on Taxonomy and Nomenclature

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Fibropapillomatosis (FP) of sea turtles (also referred to as green turtle fibropapillomatosis [GTFP] or marine turtle fibropapillomatosis [MTFP]) is a debilitating disease characterized by multiple benign cutaneous, and occasional visceral, fibropapillomas.  Because the disease was able to be transmitted to green turtles (Chelonia mydas) using filtered cell-free tumour homogenates (Herbst et al., 1995), and viral particles consistent with a herpesvirus were observed by electron microscopy in cutaneous fibropapilloma tissues (Jacobson et al., 1991), a viral aetiology was strongly suspected.  In 1998, three closely related herpesviruses were identified by consensus polymerase chain reaction and sequencing in green turtles from Florida and Hawaii and an olive ridley turtle (Lepidochelys olivacea) (Quackenbush et al., 1998).  These viruses were initially named GTHV-Fl, GTHV-Ha, and ORTHV, respectively, and represented a unique phylogenetic clade among the herpesviruses.  Subsequent studies suggested that the genetic differences among these viruses represented only minor sequence variation that reflected intratypic variants of a single FP-associated Chelonian herpesvirus (Lackovich et al., 1999).  This virus was initially called fibropapilloma-associated marine turtle herpesvirus (FPTHV) (Greenblatt et al., 2005), but more recently has also been referred to as the Chelonid herpesvirus 5 (ChHV5) (Ackermann et al., 2012) or Chelonid fibropapilloma-associated herpesvirus (CFPHV) (Alfaro-Nunez et al., 2014).  Based on genomic analysis, this virus has been taxonomically assigned to the subfamily Alphaherpesvirinae in the Family Herpesviridae and Order Herpesvirales (Davison et al., 2009).

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. However, the virus has been documented only in a small number of locations (T. Work, USGS National Wildlife Health Center Honolulu Field Station, Honolulu, Hawaii, USA, personal communication, 2015). The Distribution table in this datasheet lists only these locations; for the distribution of the disease, see the 'fibropapillomatosis of sea turtles' datasheet.

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, SouthwestPresentRodenbusch et al., 2014
Atlantic, Western CentralPresentEne et al., 2005
Pacific, Eastern CentralPresentHerbst et al., 2004
Pacific, SouthwestPresentQuackenbush et al., 2001

North America

MexicoPresentQuackenbush et al., 2001
USAPresentPresent based on regional distribution.
-FloridaWidespreadEne et al., 2005
-HawaiiPresentHerbst et al., 2004
-North CarolinaPresentHerbst et al., 2004
-TexasPresentTristan et al., 2010

Central America and Caribbean

BarbadosPresentQuackenbush et al., 2001
Costa RicaPresentQuackenbush et al., 2001
Puerto RicoPresent

South America

BrazilWidespreadRodenbusch et al., 2014

Oceania

AustraliaPresentPresent based on regional distribution.
-QueenslandPresentQuackenbush et al., 2001

History of Introduction and Spread

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In 1938, Smith and Coates at the New York Aquarium first described fibropapillomas in a captive green turtle (Chelonia mydas) that had been captured near Key West, Florida, 2 years previously (Smith and Coates, 1938).  Subsequently, they also observed fibropapillomas in three free-ranging green turtles captured off of Key West (Smith and Coates, 1938).  In the same year, Lucké also described similar tumours from a green turtle caught off of Cape Sable, Florida (Lucké, 1938).  In 1948, Schlumberger and Lucké subsequently described internal fibropapillomas in the lungs of a green turtle from Florida (Schlumberger and Lucké, 1948).  A decade later, Hendrickson noted the occasional occurrence of fibrous masses on nesting females in Malaya and Sarawak (Hendrickson, 1958).  The first confirmed case of fibropapillomatosis in Hawaii occurred the same year in a juvenile green turtle captured in Kaneohe Bay according to Balazs (1991); according to local fishermen, it was rare to nonexistent prior to this (Balazs, 1991). According to Williams and Bunkley-Williams (1996), turtles with fibropapillomas were first reported in Hawaii in the late 1940s. Aguirre and Lutz (2004) said that prevalence at Kaneohe Bay had reached 42-65%, but there has been a steady decline in the disease in Hawaii since its peak in the mid-1990s (Chaloupka et al., 2009). In 1980 at a turtle farm in Grand Cayman, an outbreak occurred that apparently began in wild-caught green turtles and subsequently developed over several years in the farmed turtles (Jacobson, 1981).  The first cases of the disease in green turtles in the Indian River Lagoon on the east coast of Florida were reported in 1982 (Ehrhart et al., 1986).  Netting surveys of the Lagoon going back to 1977 did not detect any green turtles with the disease.  In addition, late 19th Century accounts of the Florida east coast green turtle fishery and reports on Indian River Lagoon turtles published between 1978 and 1983 failed to yield any record of green turtle fibropapillomatosis prior to these first cases in 1982 (Herbst, 1994).  Continued monitoring at this site after 1982 revealed an approximately 50% prevalence (Herbst, 1994).

Risk of Introduction

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The distribution of fibropapillomatosis is worldwide, but primarily circumtropical and associated with coastal ecosystems with high human population densities and agricultural run-off.  As the Earth’s climate changes and there are changes in marine turtle and human populations, there is a possibility that the geographical distribution of the disease will expand.

Pathogen Characteristics

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The Chelonid herpesvirus 5 (ChHV5) is strongly associated with, and the presumed aetiological agent of, marine turtle fibropapillomatosis (Ackermann et al., 2012; Greenblatt et al., 2005; Lackovich et al., 1999).  Little is known about this virus, however, due to its inability thus far to be cultivated in cell culture. This inability, together with the difficulties in conducting experimental infection studies in a legally protected endangered species, mean that Koch’s postulates for proving the causal relationship between the virus and clinical disease (i.e., fibropapilloma formation) have remained unfulfilled.

Nevertheless, there is a strong association between the presence of a herpesvirus and fibropapillomatosis in sea turtles.  Some of this evidence includes disease transmission using filtered cell-free tumour homogenates (Herbst et al., 1995), observation of viral particles consistent with a herpesvirus using electron microscopy in cutaneous fibropapilloma tissues (Jacobson et al., 1991), and identification of closely related novel herpesviruses in affected tissues of green turtles (Lepidochelys olivacea) and an olive ridley turtle (Lepidochelys olivacea) from Florida and Hawaii using consensus polymerase chain reaction and sequencing (Quackenbush et al., 1998).  In fact, DNA polymerase sequences of this herpesvirus have been detected by PCR in every tested fibropapilloma and fibroma reported to date (Lackovich et al., 1999; Lu et al., 2000a; Quackenbush et al., 2001; Quackenbush et al., 1998).

Whole genome sequencing of ChHV5 DNA extracted from a glottis tumour of a Hawaiian green turtle with fibropapillomatosis determined that the virus had a type D genome with gene order typical for the varicellovirus genus within the alphaherpesvirinae (Ackermann et al., 2012).  However, four genes that are atypical for alphaherpesvirus genomes were also identified.  These included two members of the C-type lectin-like domain superfamily (F-lec1 and F-lec2), an ortholog of the mouse cytomegalovirus M04 (F-M04), and a viral sialyltransferase (F-sial).  Because RNA transcripts of F-sial and F-M04, but not F-lec1 and F-lec2, were detected in tumour tissues of Hawaiian fibropapillomatosis cases, it is thought that F-sial and F-M04 may play a role in the pathogenesis of the disease (Ackermann et al., 2012).

Host-specificity is well-known among the herpesviruses and alphaherpesviruses, in particular, are known to show extensive coevolution with their hosts (Davison, 2002; McGeoch et al., 2000).  Thus, it is not surprising that the FP-associated herpesvirus has also been co-evolving with its turtle hosts, and it has been estimated that the virus and host have co-evolved for at least 8.9 million years (Herbst et al., 2004).

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

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Host and vector organismsPossibly the marine leech Ozobranchus spp. and the cleaner fish Thalassoma duperrey Yes Greenblatt et al., 2004

Vectors and Intermediate Hosts

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VectorSourceReferenceGroupDistribution
OzobranchusGreenblatt et al., 2004. AnnelidHawaii
Thalassoma duperreyLu et al., 2000b. OtherHawaii

Environmental Impact

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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.  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.

Threatened Species

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Threatened SpeciesConservation StatusWhere ThreatenedMechanismReferencesNotes
Caretta caretta (loggerhead sea turtle)VU (IUCN red list: Vulnerable) VU (IUCN red list: Vulnerable); USA ESA listing as endangered species USA ESA listing as endangered speciesPathogenicIUCN, 2014
Chelonia mydas (green sea turtle)EN (IUCN red list: Endangered) EN (IUCN red list: Endangered); USA ESA listing as endangered species USA ESA listing as endangered speciesFloridaPathogenic; Pest and disease transmissionIUCN, 2014; National Marine Fisheries Service, 2007
Dermochelys coriaceaVU (IUCN red list: Vulnerable) VU (IUCN red list: Vulnerable)PathogenicIUCN, 2014
Eretmochelys imbricata (hawksbill turtle)CR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered); USA ESA listing as endangered species USA ESA listing as endangered speciesPathogenicIUCN, 2014
Lepidochelys kempiiCR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered)PathogenicIUCN, 2014
Lepidochelys olivacea (olive ridley sea turtle)VU (IUCN red list: Vulnerable) VU (IUCN red list: Vulnerable); USA ESA listing as endangered species USA ESA listing as endangered speciesPathogenicIUCN, 2014
Natator depressusDD (IUCN red list: Data deficient) DD (IUCN red list: Data deficient)PathogenicIUCN, 2014

Risk and Impact Factors

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  • Host damage
  • Reduced native biodiversity
  • Threat to/ loss of endangered species
  • Threat to/ loss of native species
Impact mechanisms
  • Pest and disease transmission
  • Pathogenic

Uses

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Given the link between the presence of fibropapillomatosis and the environmental health of the coastal habitat, it has been suggested that the epidemiology of the disease could serve as an effective tool to monitor ecosystem health in near-shore marine habitats (Aguirre and Lutz, 2004).

Gaps in Knowledge/Research Needs

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Although the presence of Chelonid herpesvirus 5 has been highly associated with fibropapillomatosis (FP), Koch’s postulates have not been fulfilled so far.  The virus has not been cultured in vitro and there is no proof of causality for FP.  Thus, a critical research need is to develop methods to grow the virus in in vitro culture.  Another big question is what role various co-factors play in the development of FP.  Are there other infectious agents involved?  How do various environmental factors (pollutants, water temperature, and/or marine biotoxins) contribute to the development of FP?  An additional area of inquiry is the link between development of FP and the immune system.  Immunosuppression is strongly correlated with FP (Aguirre and Lutz, 2004).  However, is this immunosuppression the cause or a consequence of FP?  Unfortunately, the availability of sufficient research funding to address these questions continues to be a major barrier to better understanding this disease.

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

Barnett LK, Emms C, Jallow A, Cham AM, Mortimer JA, 2004. The distribution and conservation status of marine turtles in The Gambia, West Africa: a first assessment. Oryx, 38(2):203 - 208.

Barragan A, Sarti L, 1994. A possible case of fibropapilloma in Kemp's Ridley turtle (Lepidochelys kempi). Marine Turtle Newsletter, 76:27.

Bresette MJ, Gorham JC, Peery BD, 2002. Initial assessment of sea turtles in the southern Indian River lagoon system, Ft. Pierce, Florida. 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, 271-273. [NOAA Technical Memorandum NMFS-SEFSC-477.] http://www.seaturtlesociety.com/proceedings/20turtle.pdf

Bresette MJ, Herren RM, Singewald DA, 2005. Comparison of fibropapilloma rates of green turtles (Chelonia mydas) from two different sites in St. Lucie County, Florida. 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, 125-126. [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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