|Scientific Name:||Balistes capriscus|
|Species Authority:||Gmelin, 1789|
Balistes buniva Lacepède, 1803
Balistes carolinensis Gmelin, 1789
Balistes forcipatus Gmelin, 1789
Balistes fuliginosus DeKay, 1842
Balistes moribundus Cope, 1871
Balistes powellii Cope, 1870
Balistes spilotopterygius Walbaum, 1792
Balistes taeniopterus Poey, 1860
|Taxonomic Source(s):||Eschmeyer, W.N. and Fricke, R. (eds). 2015. Catalog of Fishes: genera, species, references. Updated 2 September 2015. Available at: http://researcharchive.calacademy.org/research/ichthyology/catalog/fishcatmain.asp. (Accessed: 2 September 2015).|
|Red List Category & Criteria:||Vulnerable A2bd ver 3.1|
|Assessor(s):||Liu, J., Zapfe, G., Shao, K.-T., Leis, J.L., Matsuura, K., Hardy, G., Liu, M. & Tyler, J.|
|Facilitator/Compiler(s):||Carpenter, K.E., Comeros-Raynal, M., Harwell, H. & Sanciangco, J.|
Balistes capriscus is widely distributed where it occurs over hard bottoms, reefs, and ledges. Juveniles are associated with Sargassum. It is valued by both commercial and recreational fisheries. A stock assessment in the Gulf of Mexico indicates that the current rate of removal is not sustainable: a steady decline in landings since the peak in 1990, current landings (850,000 pounds in 1998) are below the minimum sustainable yield (MSY) range, estimated biomass levels are low, and estimated exploitation rates are high. This evidence suggests that the stock is overfished and that catches should at least be held constant if not reduced to bring the population back to sustainable levels. Spawning stock biomass in the northern Gulf of Mexico has declined by 43-58% over three generation lengths. In Brazil, declines in landings have been reported from Sao Paulo, Rio de Grande Sul, and Espirito Santo, the latter state reporting commercial extinction of this species due to over exploitation. Capture production for triggerfishes in the eastern central Atlantic show pronounced declines in catch. This species once dominated the Gulf of Guinea ecosystem, especially from Ghana to Sierra Leone, for nearly 20 years. However, annual catches of triggerfish declined drastically; this is especially noticeable in Ghana, where landings have very low since 1989.
The species is managed by the South Atlantic Fishery Management Council and under the Gulf of Mexico 1981 Reef Fish Fishery Management Plan (FMP) and subsequent amendments, conservation measures include size limits (12" TL), catch limits (five fish/person bag limit, 20 fish snapper-grouper aggregate), and gear restrictions. A rebuilding plan was created by the Gulf of Mexico Fishery Management Council and implemented by the National Marine Fisheries Service in August 2008, but there are currently no indications of recovery.
In over 25% of its range (Gulf of Mexico, Gulf of Guinea and Brazil), it is estimated that populations have declined by 63-68% over three generation lengths. We infer that population declines of at least 30% over three generations have occurred throughout the remainder of its range (~50%) except for the Atlantic seaboard of North America where it is only harvested recreationally. Globally, it is estimated that this species has declined by at least 30% over the past three generation lengths. Therefore, Balistes capriscus is listed as Vulnerable under A2bd.
|Range Description:||Balistes capriscus is widespread in the Atlantic Ocean. In the western Atlantic, it is known from Nova Scotia (Canada) south along the U.S. coast, Bermuda, the Bahamas, throughout the Gulf of Mexico and Caribbean Sea, and along South America to Argentina, including Trinidade Island (Simon et al. 2013). In the eastern Atlantic, it is known from Ireland south into the Mediterranean Sea, the Azores, the Canary Islands, the Islands of Madeira, and along West Africa to Angola, including offshore oceanic islands in this region. Its depth range is 0-110 m.|
Native:Albania; Algeria; Angola (Angola); Anguilla; Antigua and Barbuda; Argentina; Aruba; Bahamas; Barbados; Belgium; Belize; Benin; Bermuda; Bonaire, Sint Eustatius and Saba (Saba, Sint Eustatius); Brazil (Trindade); Bulgaria; Cameroon; Canada; Cape Verde; Cayman Islands; Colombia; Congo; Congo, The Democratic Republic of the; Costa Rica; Côte d'Ivoire; Croatia; Cuba; Cyprus; Dominica; Dominican Republic; Egypt; Equatorial Guinea; France; French Guiana; Gabon; Gambia; Georgia; Ghana; Gibraltar; Greece; Grenada; Guadeloupe; Guatemala; Guernsey; Guinea; Guinea-Bissau; Guyana; Haiti; Honduras; Ireland; Israel; Italy; Jamaica; Jersey; Lebanon; Liberia; Libya; Malta; Martinique; Mauritania; Mexico; Monaco; Montenegro; Montserrat; Morocco; Netherlands; Nicaragua; Nigeria; Norway; Panama; Portugal (Azores, Madeira); Puerto Rico; Romania; Russian Federation; Saint Helena, Ascension and Tristan da Cunha (Ascension, Saint Helena (main island)); Saint Kitts and Nevis; Saint Lucia; Saint Martin (French part); Saint Vincent and the Grenadines; Sao Tomé and Principe; Senegal; Serbia (Serbia, Serbia); Sierra Leone; Sint Maarten (Dutch part); Slovenia; Spain (Canary Is.); Suriname; Sweden; Syrian Arab Republic; Togo; Trinidad and Tobago; Tunisia; Turkey; Turks and Caicos Islands; Ukraine; United Kingdom; United States (Georgia); Uruguay; Venezuela, Bolivarian Republic of; Virgin Islands, British; Virgin Islands, U.S.; Western Sahara
|FAO Marine Fishing Areas:|
Atlantic – western central; Atlantic – southwest; Atlantic – southeast; Atlantic – northwest; Atlantic – northeast; Atlantic – eastern central; Mediterranean and Black Sea; Pacific – eastern central
|Range Map:||Click here to open the map viewer and explore range.|
|Population:||U.S. Gulf of Mexico|
The Gray Triggerfish is managed as a single stock in the Gulf of Mexico. Analysis of mitochondrial DNA sequences from five localities across this species’ exploited range in U.S. waters revealed low-levels of heterogeneity, which is consistent with the management of this species as a single stock (Antoni et al. 2011). Microsatellite markers have recently been developed for finer-resolution studies of population structure (Antoni and Saillant 2012, Sorenson 2013).
Projections of possible future condition of the stock under different fishing scenarios were made using parameter outputs from the ASPIC (A Stock Production Model Incorporating Covariates) models one and two using 1998 as the last year in the assessment and 1999 as the first year of management. To determine whether or not the stock could be rebuilt (B/BMSY =1.0) within a ten year time frame, projections were carried out to the years 1999-2008. These projections indicate that the stock may be overfished, that overfishing is still occurring, and that catches should at least be held constant or preferably reduced to bring the stock back to healthy levels. The production model analyses utilized to project the stock trajectory indicated that current fishing mortality rates were not sustainable (Valle et al. 2001). Although management efforts are in place for the Gulf of Mexico population (see the Conservation Actions section for detailed management measures), no indications of population recovery were evident as of 2014. The next benchmark assessment for B. capriscus is scheduled for 2015.
An aggregated production model indicated that there were uncertainties in the ASPIC models used, due to sensitivity of the model. Robust conclusions from the model results cannot be determined, as the data were not adequate to resolve the status of the stock; however, nearly every run conducted resulted in the stock being overfished and experiencing overfishing. It is noted that this model should be explored further (Nowlis and Saul 2005).
A benchmark assessment was performed in 2006, and updated in 2011 using the same statistical catch-at-age-model (SSASPM). Results suggest the stock is overfished and experiencing overfishing. The stock was approaching an overfishing condition in 2005, but was not yet overfished. A rebuilding plan was established 2008 to rebuild the stock within 10 years, but there is currently no evidence of substantial improvements in stock status. The ratio of SSBcurrent/SSBMSY ranged from 0.24-0.39. Using the Gulf-wide shrimp effort model, which was evaluated as containing the least amount of uncertainty, SSB in 2010 was estimated at 6.90E+11, which is roughly 60% below the SSBMSY proxy of 1.78E+12. Fcurrent in 2010 was estimated at 0.35, which is above FMSY of 0.34. Estimates of fishing mortality ranged from 0.53 to 0.35 depending on the model used in the 2011 update (in 2006 fishing mortality was estimated at 0.435). The overall trend in SSB from 1963 (when the stock was assumed to be unfished) to 2010 is one of decline – current SSB is roughly 15-20% of original, un-fished biomass. The preferred model from the U.S. Gulf of Mexico stock assessment estimated that spawning stock biomass declined by 43-58% during the past three generation lengths (12-14 years) (see table A-24 in SEDAR 2011). The continuity model estimated that spawning stock biomass declines were slightly higher, 62-69% over three generation lengths (see Table A-1 in SEDAR 2011). In addition, models suggested recruitment has also decreased; depending on model specifications, the declines range from 51-68% to 84-87% in the last 12-24 years (see Tables A-24 and A-1 in SEDAR 2011, respectively).
Projections of SSB for Gray Triggerfish in the Gulf of Mexico were generated using the 'Gulf-wide Shrimp Effort' model. Under each of three scenarios for which SSB was projected, SSB was expected to increase. Recovery periods under the no fishing scenario ranged from 3 to 4.5 years. However, there is concern that the trend of steadily declining recruitment and the low recruitment levels in recent years were not reflected in the model, which uses the estimated spawner-recruit relationship, and thus projections of recovery may be overly optimistic. The Allowable Biological Catch advice for 2012 and 2013 was based on the assumption that recruitment was at recent (2005-2009) levels, rather than based on the spawner-recruit relationship (SEDAR 2011); this was thought a more precautionary approach to Grey Triggerfish harvest.
Recreational landings in numbers of fish by state (Texas, Mississippi, Alabama, Louisiana and Florida) and fishing mode were estimated for the period 1986-1998, landings peaked in 1990, followed by a steady decline. The majority of annual landings since 1986 have been reported from the West Coast of Florida, followed by Louisiana. Landings from the charter boat mode have dominated recreational landings most years. Landings in weight by year and state were estimated for the commercial sector for the period 1986-1998 with a trend similar to the recreational sector, but with a peak in 1993 followed by a steady decline. The greatest proportion of commercial landings has been reported for the west coast of Florida, followed by Louisiana.
Southern Gulf of Mexico: Balistes capriscus is a minor commercial species along the Mexican coast in the Gulf of Mexico, where it is primarily utilized in small-scale artisanal fisheries (a more important commercial fishery for this species is along the Mexican Caribbean coast). Anecdotal evidence suggests that populations have declined in this region, but no quantitative data are available at this time.
U.S. Atlantic coast
This species is not targeted by fisheries in this region (Kellison et al. 2012). Mean annual catch per unit effort (CPUE) in the Southeast Area Monitoring and Assessment Program during the fall and summer trawl surveys from 1987 to 2010 indicate no clear trend (NOAA unpub. data). Catch per unit effort (CPUE) was greater in the 1970s (2.5% of catch) than in 2005-2006 (0.4% of catch) at specific capture site (210 Rock) in Onslow Bay, North Carolina (Ruderhausen et al. 2008). It is a dominant component within Sargassum off North Carolina (Casazza and Ross 2008).
In Brazil, it has been intensively caught in the industrial fleet over the past two decades. Total landings decreased by 10.5%, with landings from 1996-2005 at 29.64 t and from 2001-2005 at 26.54 t (Ataliba et al. 2009). It has been exploited off the state of São Paolo since the 1960s. In the past, it was included in the miscellaneous fish category, but in the last 15 years, it has become a very important fishery resource and is currently one of the most commonly caught commercial species (Bernardes 1988, Castro 2000). It was previously one of the most important species in the fisheries of Espirito Santo State, but overexploitation has since caused it to become commercially extinct (Freitas Netto and Di Beneditto 2010, Freitas Netto et al. in press). The use of gill nets may have caused the declines in population (Freitas Netto 2004). About 46% of the catches were composed of immature fishes, which increases the likelihood for growth overfishing to occur (Bernardes and Dias 2000, Bernardes 2001).
Landings by state changed substantially from 1986-2005. The majority of the landings came from Sao Paulo and Rio de Janeiro, with generally lower landings from Santa Catarina and Rio de Grande Sul. Landings have declined by approximately 95% over 20 years in Sao Paulo and Rio de Grande Sul, while Santa Catarina has no trend, and landings increased by approximately 200% in Rio de Janeiro (Ataliba et al. 2009). However, increased landings in Brazil may be due to declines in more traditional fisheries (Ataliba et al. 2009) and therefore, may not clearly reflect the status of the population.
FAO capture production list annual Mediterranean and northeast Atlantic landings for this species from 1992-2009 as ranging from one tonne to 432 tonnes. Tunisia and Portugal have the highest catches from 1993-2009, with an average of 77 tonnes (1993-2009) and 66 tonnes (1999-2009), respectively (FAO 2014). Libya has the highest catch recorded in 2009 at 432 tonnes. Capture production for triggerfishes in the eastern central Atlantic show pronounced declines in catch in Ghana and Togo. Furthermore, declines in catches were also recorded for the Soviet Union from 1970-1989 in the divisions: Cape Verde, Sahara and Sherbro. In Ireland, there was an increase in triggerfish abundance between 1989-1991, with at least 184 specimens recorded, representing 53.6% of the total number of specimens documented (343) since the first record in 1845. Large numbers of specimens were reportedly sighted by SCUBA divers and captured on rod and line by anglers. It is unknown if the increase in triggerfish numbers represent a real change in the species overall abundance in Irish water or an increase in recording effort (Quigley et al. 1993).
Balistes capriscus appears to be one of the most abundant near-bottom pelagic species occurring in the Sierra Leone waters. Results of two surveys conducted by the UNIDO/GEF Gulf of Guinea LME Project and FAO/NORAD in Moree, Ghana indicate that this species is one of the top 20 species encountered, indicating possible recovery of the stock of this species in Ghana (Koranteng 2000). In Cote-d'Ivoire, this species was very abundant from 1983-1986 during the hot season, but then disappeared from 1993 (there was no survey conducted between 1987 and 1992). There was apparently no link of the abundance of B. capriscus to the usual physical and biological environmental parameters. This was the opposite of the pattern that was observed in Ghana by Koranteng (1998), where species abundance was higher in colder regime and declined when waters warmed up (Joanny and Menard 2002).
In Ghana, from ~1973 and following a decline in the sardinella fishery in the western Gulf of Guinea, there was an increase in landings of this species (as Balistes carolinensis) in the Ghanian demersal fishery. In 1979-1980, the demersal component of triggerfish assessed in bottom trawl surveys was put at 99,000 t (or 57.2 kg ha-1), or 62% of the total demersal fish biomass in Ghanian coastal waters (Koranteng and Pauly 2002). In May 1981-March 1982, the estimated biomass was between 314,000 and 500,000 t (Stromme et al. 1982, Ofori-Adu 1987, 1994, Aggrey-Fynn 2007). Triggerfish catch drastically declined from 18,610 tonnes in 1986 to 2 tonnes in 1995 in Ghanaian waters, and recent report from CECAF capture production (FAO 2008) show that the annual catches of triggerfish show two separate peaks between 1972 and 2003. The first catch peak (12,563 tons) occurred in 1979, and the second peak (17,250 tons) in 1986/87. Before 1979, triggerfish catch levels were from 2 830 tons (1972) to 9,138 tons (1977). In between 1979 and 1986/87, triggerfish catch fell to ‘regular or stable’ annual levels (5,000–8,000 tons/year). After 1987, triggerfish catch experienced a ‘sharp’ decline in 1988 to 2,862 tons which continued to decline, indicating that inshore triggerfish catch generally increased with inshore effort. Inshore catch decreased dramatically from 1979-80 at relatively same effort and continued to decrease from 1980-84, catch increased in 1984-1986 at increased effort and decreased progressively with effort from 1987-91 (Aggrey-Fynn 2007). The triggerfish biomass was highest in 1985 and decreased afterwards up to 1991. The biomass decrease was quite significant from 1987-89 at virtually the same effort; and 1989-91 there was virtually no biomass of the triggerfish resource in Ghanaian coastal waters (Koranteng 1998, Aggrey-Fynn 2007).
Balistes capriscus is an important economic resource in the Gulf of Guinea. It was recorded to have reached maximum abundance in the late 1970s to early 1980s in the Canary Current (Caverivière 1982, Stromme et al. 1982). This species was the most abundant demersal/semi-pelagic species in the 1980s (Mensah and Quaatey 2002). At the time that this species dominated in the Gulf of Guinea ecosystem, it corresponded to a period of low temperature and high salinity, and a decrease in the total density of sciaenids, some lutjanids were also displaced by the expanding triggerfish population with the latter occupying the niche of the snappers (Koranteng 1998, Koranteng 2001, Koranteng and Pauly 2002). In 1982-1990, the Balistes capriscus stock declined and accounted for only 4% of the total catch of the species and the other groups in the sparid species assemblage in the Gulf of Guinea (Koranteng 2001a). This species dominated the Gulf of Guinea ecosystem, especially from Ghana to Sierra Leone, for nearly 20 years, displacing Brachydeuterus auritus as the most abundant species off Ghana. For reasons that have yet to be determined, triggerfish declined in abundance from ~1988, and only a few specimens were caught in surveys conducted in the 1990s (Koranteng and Pauly 2002). There are reported to be two separate stocks in the Gulf of Guinea with no apparent interchange- the eastern stock which occurred off Ghana and the western stock off Guinea Bissau and Guinea (Stromme et al. 1982, Stromme 1983, Mensah and Quaatey 2002); both have almost disappeared (Aggrey-Fynn 2007). To address this gap, catch and biological data were obtained for this species was obtained from trawl surveys conducted in 2005 off the waters of Benin, Togo, Ghana and the Ivory Coast, results of the study indicate that the distributions of the Grey Triggerfish had not changed despite the decline documented for nearly two decades. Moreover, the depth distribution, modal size classes and growth were comparable to previous studies before the decline (Aggrey-Fynn 2009).
Grey Triggerfish exploitation ratio (0.2727) indicate that the triggerfish resource is underexploited as compared to the exploitation ratio of 0.67 derived from 1980 growth results of Gray Triggerfish (Aggrey-Fynn 2007). Thus, the triggerfish resource might not have been overexploited before their disappearance. Though current exploitation rates show an under-exploited fishery, the Balistes fishery has not revived in Ghana. Environmental forcing, specifically the intense cooling along the coast of Ghana in recent years and triggerfish migration from cold coastal waters to warm offshore waters, and the unusual minimum mean temperatures experienced in 1979 and 1987 that have not been recorded again in coastal waters of Ghana (Aggrey-Fynn 2007) may be impacting the populations.
|Current Population Trend:||Decreasing|
|Habitat and Ecology:||Balistes capriscus inhabits hard bottoms, reefs, and ledges. It is also associated with artificial reef structures (Frazer and Lindberg 1994, Vose and Nelson 1994, Wilson et al. 1995, Kurz 1995, Simmons 2008). It has a wide bathymetric distribution (Ofori-Adu and Koranteng 1993), usually between 15–55 m depth in coastal waters, but the preferred depth range is 30–50 m (ATLANTNIRO 1980). Adults drift along the bottom either alone or in small groups, while the juveniles drift at the surface with Sargassum. High abundances of small fish (1.3 - 10.6 cm standard length) have been recorded from Sargassum mats in the northwestern Gulf of Mexico in the pelagic zone from May to August (Wells and Rooker 2004). It occurs mainly in the thermocline at an average depth of 47 m, with temperature and oxygen values below the values found in shallow waters in clay and silt substrate (Bianchi 1992a). In the Gulf of Mexico, adults inhabit reef areas (natural and artificial reefs, low or high-relief reefs) between 10-106 m (Smith 1976, Johnson and Saloman 1984, Ingram 2001, K. Rademacher pers. comm.). Age-0 gray triggerfish recruited to benthic artificial reefs from September to December in the northern Gulf of Mexico (Simmons 2008). This species has a protracted pelagic stage and recruits to benthic reef structure as relatively large, older juveniles from September to December. It spends 4-7 months in the pelagic environment before recruiting to benthic habitat (Simmons and Szedlmayer 2011). |
The von Bertalanffy growth parameters indicate that males attain a larger size than females (Ingram 2001). The instantaneous mortality rates were estimated as 0.26 for males and 0.27 for females (Bernardes 2002). Balistidae spp. form harems, build demersal nests, and demonstrate parental care of eggs (Fricke 1980, Blumer 1982, Gladstone 1994, Ishihara and Kuwamura 1996, Kuwamura 1997). This species is a multiple-batch spawner, starting as early as May and peaking during June and July, and with a 1:2 male to female ratio. Males and females both reach sexual maturity at 250 mm fork length (FL), but males are at age-1 and females are at age-2 (Wilson et al. 1995, Ingram 2001). Eggs are incubated in demersal nests within 12 to 58 hours, after which they enter the plankton (Thresher 1984). Between the sizes 1 to 8 cm (SL), it is associated with Sargassum (Ingram 2001). It is sexually dimorphic, with males being significantly larger than females (Ingram 2001, Moore 2001, Simmon 2008). In West Africa, spawning occurred from October to December with a peak in the warmer months (November and December). The onset of spawning was characterized by relatively short preceding period of average minimum sea-surface temperature of 22.6 degree C, normally occurring in July. First time spawners were 13.3-15.7 cm (FL), 50.0-70.5 g in weight and one year old (Ofori-Danson 1990). In Brazil, spawning occurs from November to February south of Sao Paolo (Bernardes and Dias 2000). In the Gulf of Mexico, the spawning season is estimated to be from July to October (Dooley 1972). It has a high fecundity (Ingram 2001, Simmons 2008). Batch fecundities in fish from the eastern Gulf of Mexico ranged from 213,912 to 1,172,854 oocytes from fish ranging in size from 267 to 388 mm FL, and relative batch fecundity had a mean of 13,809 oocytes per gram ovary and ranged from 6,318 to 24,188 oocytes per gram (Hood and Johnson 1997). One-year-old males (>250 mm fork length) and two-year-old females (>250 mm fork length) exhibit seasonal maturation cycles associated with spawning. The mean number of spawnings (± standard error) per spawning season was estimated to be 24.3 (± 4.1). Mean total annual fecundity (± standard error) was estimated to be 17,071,634 eggs year-1 (± 2,010,787).
The maximum size is 52 cm total length (Aggrey-Fynn 2009) and maximum age is at least 16 years (SEDAR 2005). Off southern Brazil, the longevity was estimated at 11 years for males and females. The mean annual mortality rate (based on number of fish at age) ranged from 0.32 to 0.53 (Johnson and Saloman 1984). Longevity is estimated at between 12 and 16 years old; using a maximum age based estimator with low prediction error (M = 4.899t−0.916: Then et al. 2014), the estimated adult natural mortality rate is 0.39-0.50. Age at first maturity is 1-2 years of age. Using the estimator for generation length of 1/adult mortality + age at first reproduction results in a generation length of 4 years; incorporating age-specific fecundity (estimated for the Gulf of Mexico, 170289e0.3159*Age: SEDAR 2011) resulted in a generation length of 8 years. Thus, the time window for calculating population declines over three generation lengths is 12-24 years.
|Generation Length (years):||4-8|
|Use and Trade:||Balistes capriscus is important to both commercial and recreational fisheries. Until recently, it was not considered a desirable catch by most fishers; however, the decline in other reef fish stocks (e.g., red snapper and groupers) has probably caused an increased targeting of this and other under-utilized species (Valle et al. 2001). It is an important economic resource in the Gulf of Guinea. It can be taken in bottom trawls and traps, as well as on handlines and is generally marketed fresh (Matsuura 2002).|
|Major Threat(s):||There have been declines in abundance of this species in the Gulf of Guinea, Gulf of Mexico and Brazil due to overexploitation.|
In U.S. waters, this species is managed under the South Atlantic Fishery Management Council and under the Gulf of Mexico 1981 Reef Fish Fishery Management Plan (FMP) and subsequent amendments. In 1984, prohibitions on roller trawls and fish traps may have reduced catch of this species. In 1999, a minimum size limit and a 20-fish aggregate bag limit for reef fishes was established. A commercial quota was set to 80,000 pounds for 2008, 93,000 for 2009, and 106,000 pounds for 2010. Following the 2011 stock assessment, the Gulf of Mexico Fishery Management Council established measures that included reductions in total allowable catch to end overfishing and rebuild the stock by 2017. The implementation of Amendment 37 established a closed season from June 1- July 31, and reduced the bag limit to two Gray Triggerfish per person.
It is not known to be well-managed where it is exploited elsewhere in its range.
|Errata reason:||Correction in assessor name, Jing, L to Liu, J.|
|Citation:||Liu, J., Zapfe, G., Shao, K.-T., Leis, J.L., Matsuura, K., Hardy, G., Liu, M. & Tyler, J. 2015. Balistes capriscus. (errata version published in 2016) The IUCN Red List of Threatened Species 2015: e.T193736A97662794.Downloaded on 18 August 2017.|
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