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First record of three shark species,Odontaspis ferox, Mustelus albipinnisand Centrophorus squamosus, from

the Galapagos Islandsdavid acun~ a-marrero

1, johanna s. zimmerhackel

2, jens mayorga

3and alex hearn

4

1Estacion Cientfica Charles Darwin (ECCD), Avenida Charles Darwin s/n, Puerto Ayora, Santa Cruz, Islas Gala pagos, Ecuador,2Institut fur Hydrobiologie und Fischereiwissenschaft (IHF), Universitat Hamburg, Olbersweg 24, 22767 Hamburg, Germany,3Universidad San Francisco de Quito, GAIAS, San Cristobal, Galapagos, Ecuador, 4Turtle Island Restoration Network, Olema, CA,USA

Two shark species, Odontaspis feroxandCentrophorus squamosus, are recorded for the first time at the Galapagos Islandsbased on one sighting reported by tourist divers and one specimen captured by a local fisherman. The identification of a third

shark species, Mustelus albipinnis, previously unidentified and recorded as Mustelus sp., is achieved based on several sightingsat the same location.

Keywords: Odontaspis ferox, Mustelus albipinnis, Centrophorus squamosus, Galapagos Islands, range distribution

Submitted 15 May 2013; accepted 5 June 2013

I N T R O D U C T I O N

The Galapagos Islands and their surrounding Marine Reserveare situated in the eastern Pacific Ocean, 1000 km to the westof mainland Ecuador covering a total area of approximately

138,000 km2

. The islands are located at the confluence ofthree major ocean currents that show a strong seasonal vari-ation in intensity and direction (Chavez & Brusca, 1991).From May to November the cool Humboldt Current predo-minates establishing a cool, dry season known as garua,while from December to May its intensity decreases and it isthe warm Panama Current that prevails, creating a warmrainy season (Banks, 2002). This complex environmentalsetting is reflected in five distinct biogeographical regionsfor subtidal reef communities (Edgar et al., 2002) and anextraordinarily important and diverse marine reserve, recog-nized by UNESCO as a World Heritage Site (Heylings et al.,2002).

Galapagos fish biodiversity has been studied and reviewedby different authors (McCosker & Rosenblatt, 1984;McCosker, 1987; Grove & Lavenberg, 1997). Long-term sub-tidal ecological monitoring programmes, local fishers andscientific expeditions have been some of the most importantsources of new information. In the last published review thenumber of shore-fish species recorded in Galapagos stood at550, including 30 shark species belonging to ten differentfamilies (McCosker & Rosenblatt, 2010).

The smalltooth sandtiger shark Odontaspis ferox (Risso,1810) is reported as having a circumglobal but fragmenteddistribution between 408N and 408S, mostly in warm-temperate and tropical waters (Compagno, 1984; Bonfil,1995). Odontaspis ferox is an offshore shark of deep waters,

observed or caught mainly near the seabed, close to continen-tal shelves and upper slopes at depths ranging from 10 to883 m (Fergusson et al., 2008). The size ofO. feroxat maturitydiffers in sexes, with ~275 cm for males and 365 cm totallength (TL) for females, with the largest recorded size of afemale measuring 450 cm TL (Compagno et al., 2005).Similar to Carcharias taurus, O. ferox is presumed to have avery low reproductive capacity, producing only two pupsevery two years and possibly practises uterine cannibalismin the form of oophagy (Compagno, 2001). This shark isfished with bottom gill-nets, line gear and bottom trawls prin-cipally in Japan, the Mediterranean Sea and southern Africaoff Madagascar (Abe et al., 1968; Cooke, 1997). Although O.

ferox is mostly discarded or utilized by-catch, there is ademand for its large oily liver in Japan (Abe et al., 1968).The likely low reproductive capacity and natural smallnumbers of this species makes it potentially sensitive tolocal extinction, even at small capture rates (Fergussonet al., 2008), and it is listed as Vulnerable in theInternational Union for Conservation of Nature and NaturalResources Red List of Threatened Species (IUCN RL). Thisis the first record ofO. ferox in the waters of Galapagos.

Mustelus albipinnis was described by Castro-Aguirre et al.(2005) as being one of two un-named species of smoothhoundshark known to be found from the Gulf of California toEcuador (P. Heemstra in Compagno, 1988 and in McCosker& Rosenblatt, 2010). Smoothhound sharks are generally

found on the continental shelves in temperate and tropical

Corresponding author:

D. Acuna-Marrero

Email: [email protected]

1

Marine Biodiversity Records, page 1 of 6. # Marine Biological Association of the United Kingdom, 2013

doi:10.1017/S1755267213000596; Vol. 6; e87; 2013 Published online
mailto:[email protected]:[email protected]


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waters around the world, yet M. mento and Triakis maculataare both also reported as present in the Galapagos Islands(McCosker & Rosenblatt, 2010). As a recently-describedspecies, little is known about its biology and ecology. It isthought to be benthopelagic, with a reported depth distri-bution of 30281 m (Perez-Jimenez et al., 2005). Bothmales and females mature at 90100 cm TL, and the largestreported size was a 118 cm TL female caught in the Gulf of

California (Perez-Jimenez et al., 2005). Its reproductionmode is by placental viviparity, with brood sizes rangingfrom 3 to 23 pups (Perez-Jimenez et al., 2005). This speciesis caught in the Gulf of California, where smoothhoundsmay represent 79% of the total catch by the local artisanalshark fishery (Marquez-Farias, 2000), but due to lack ofproper identification until the recent description of thespecies, its status has not been assessed. This shark was pre-viously recorded in Galapagos as Mustelus sp. (McCosker &Rosenblatt, 2010) but has now been identified to species level.

The leaf-scale gulper shark, Centrophorus squamosus(Bonnaterre, 1788), is a deep-water squaloid species which isfound in the eastern Atlantic Ocean, the Indian Ocean andthe western Pacific Ocean, where it has been recorded fromJapan to New Zealand (Girard & Du Buit, 1999; Compagnoet al., 2005; Verssimo et al., 2012). This benthopelagicshark inhabits the continental slopes at depths between 230and 2400 m (Compagno & Niem, 1998). Size at maturity isgreater in females (124 cm TL) than in males (86 cm TL)(Girard & Du Buit, 1999). It is a lecithotrophic viviparousspecies, according to the recent classification proposed byHamlett et al. (2005). Centrophorus squamosus is an impor-tant component of deep-water longlining and bottom trawlingfisheries in the eastern Atlantic Ocean due to the value of its

liver and flesh (Severino et al., 2009). Catches have decreasedsignificantly during the last decade and this shark is listed asVulnerable in the IUCN RL (White, 2003). This first recordof C. squamosus in the Galapagos Islands extends its knownrange into the eastern Pacific.

This paper contributes to the development of the SpeciesChecklist of the Charles Darwin Foundationan attempt toestablish a complete biodiversity catalogue for the Galapagos

Islands (Bungartz et al., 2009). It also highlights the contri-bution that citizen science can make, in the form of photo-graphs and specimens provided by local residents, fishersand tourists.

M A T E R I A L S A N D M E T H O D S

Each shark species was observed in a different biogeographicalregion: Wolf Island in the far north (Odontaspis ferox), SanCristobal Island in the centre-south (Mustelus albipinnis)and Fernandina Island in the west (Centrophorus squamosus).Shark length was estimated visually for O. ferox and M. albi-

pinnis, but measured in the case of C. squamosus.Shark Bay (1823.04N 91848.46W) is located at the east of

Wolf Island (Figure 1). It is a rocky reef exposed to the predo-minant south-east current with a shallow narrow platform fol-lowed by a steep drop-off to about 600 m. The seabed here ismainly composed of big boulders with small sandy patchesand abundant coral formations. Photograph and videofootage were taken during the only sighting recorded so far.

Tongo Reef (0854.19S 89837.51W) is a shallow flat rockyplatform that extends to about 300 m off the coast at the westcorner of San Cristobal Island (Figure 1). The border of this

Fig. 1. (A) Map of the Galapagos Archipelago with the islands of Wolf, San Cristo bal, Fernandina and Isabela indicated; (B) map of San Cristo bal Island showingthe location of the Mustelus albipinnis new record at Tongo Reef; (C) map of Wolf Island showing the location of the Odontaspis feroxnew record at Shark Bay; (D)

map of Fernandina and Isabela Islands showing the location of the Centrophorus squamosus new record south to Punta Mangle.

2 d avi d a cun~ a-marrero e t a l.


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platform drops suddenly to depths over 50 m. The site of theencounters is less than 100 m from the coastline. Photographand video footage were taken during the first sighting.

On 14 January 2012, one specimen of Centrophorus squa-mosus was caught off the east coast of Fernandina Island(0833.38S 91824.23W). The shark was captured at approxi-mately six miles south of Punta Mangle, at the south-easternside of Fernandina Island (Figure 1) using a 200 hundred

hooks longline set near the bottom, at 600 m depth. Thebait used was black-striped salema (Xenocys jessiae). Thisinformation was provided by the fisherman and obtainedusing a depth-sounder. The shark was brought to theCharles Darwin Research Station, where it was identifiedand examined by biologist David Acuna.

R E S U L T S

systematicsOrder LAMNIFORMES

Family ODONTASPIDIDAE Muller & Henle, 1839Genus Odontaspis Risso, 1826Odontaspis ferox Risso, 1810

One unknown shark was observed in the late afternoon on 22November 2010 by dive tourists Martin Krammer and MartinSchlifski at Sharks Bay, Wolf Island, Galapagos (1823.04N91848.46W). Both divers videotaped and photographed theshark because of its uncommon appearance (Figures 2 & 3).The individual was a female of ~400 cm TL, swimmingfrom deep waters towards the surface along the rockybottom with sandy patches in a protected area. The encounterlasted approximately two minutes and the shark was swim-ming slowly away when divers approached it. The divers

reported that it may have been pregnant, given its distendedabdomen. Photographs and videos were shared with diveguides Lenin Barrera and Ruben Intriago, and given to theCharles Darwin Foundation (CDF) where scientists JohannaZimmerhackel, Cesar Penaherrera and David Acuna ident-ified the specimen as the smalltooth sandtiger sharkOdontaspis ferox due to the following features: (i) anal finpresent; (ii) two dorsal fins and five gill slits; (iii) eyes situatedover mouth; (iv) caudal fin asymmetrical without keel; (v)subterminal mouth with large bladelike anterior teeth; and(vi) long conical snout and first dorsal fin closer to pelvic

fin than to ventral fin. The identification is additionally sup-ported by previous personal sightings by the author (D.A.),who observed this species several times in the Mar de lasCalmas Marine Reserve, Hierro Island, Canary Islands, Spain.

Order CARCHARHINIFORMES

Family TRIAKIDAE Gray, 1851Genus Mustelus Linck, 1790

Mustelus albipinnis Castro-Aguirre, Antuna-Mendiola,Gonzalez-Acosta & de la Cruz-Aguero, 2005

Local student Jens Mayorga reported seeing an unusual sharkwhile SCUBA diving off the coast of San Cristobal, undertak-ing a study of shark presence at Tongo Reef, where fishers reg-ularly clean their catch. He reported three encounters: the firsttook place on 11 September 2010; the second on 11 September2011; and the third on 10 April 2012. Each encounter was witha single, apparently female individual, which he estimated tobe between 150 and 200 cm TL (significantly larger than the

reported maximum length for this species), swimmingslowly in approximately 10 m depth (significantly shallowerthan the shallowest depth reported for this species). Allthree encounters took place around dusk, and the firstoccurred ten minutes after a fishing boat started to clean itscatch at the site. The first encounter lasted over twentyminutes, with the shark showing signs of curiosity towardsthe diver, who at that time was alone. It slowly circled thediver several times over this period. During the remainingtwo encounters, when the diver was accompanied by abuddy, the shark appeared more active, and only circled thedivers once. Photographs (Figure 4) and video stills (takenduring the first encounter) were sent to John McCosker(California Academy of Science) and Phil Heemstra (South

African Institute for Aquatic Biodiversity), both of whomidentified the shark as Mustelus albipinnis. Mustelus albipinniscan be distinguished visually by the white outline of its dorsal,

Fig. 2. Individual of Odontaspis ferox observed at Shark Bay, Wolf Island(photograph extracted from video footage filmed by Martin Krammer and

Martin Schlifski during a diving trip to the Galapagos northern islands).

Fig. 3. Detail of the long conical snout and large bladelike teeth of theOdontaspis ferox observed at Shark Bay, Wolf Island (photograph extractedfrom video footage filmed by Martin Krammer and Martin Schlifski).

Fig. 4. Mustelus albipinnis swimming in shallow water at Tongo Reef, SanCristobal Island (photograph extracted from video footage filmed by Jens

Mayorga).

first record of three shark species from the gala p ag os i sl an ds 3


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pectoral, pelvic and anal fins, being the only known hound-shark with this feature (Castro-Aguirre et al., 2005).

Order SQUALIFORMES Compagno, 1973Family CENTROPHORIDAE Bleeker, 1859Genus Centrophorus Muller & Henle, 1837Centrophorus squamosus Bonnaterre, 1788

Identification as Centrophorus squamosus was made by thefollowing main features: (i) body cylindrical and low; (ii)five pairs of gills and spiracles present; (iii) no anal fin, finspines present on both dorsal fins and caudal fin with sub-terminal notch; (iv) bladelike unicuspidate teeth in upperand lower jaws, with lowers much larger; and (v) rear tips ofpectoral fins broadly angular and not strongly extended. Aset of pictures of the shark were sent to Alberto Brito(University of La Laguna, Spain) who confirmed the speciesidentification as Centrophorus squamosus (A. Brito, personalcommunication, 2012) (Figure 5). The individual examinedwas a mature female, measuring 95 cm TL, and weighting5.2 kg. Five yolk balls were found within the uterus and

embryos were not distinguished.

D I S C U S S I O N

Odontaspis feroxand Centrophorus squamosus constitute twonew records of shark species in the Galapagos Marine Reserveand two new families not previously registered: Odontaspidaeand Centrophoridae. The species listed until now as Mustelussp. (McCosker & Rosenblatt, 2010) is ultimately identified as

Mustelus albipinnis.Since the description of the smalltooth sandtiger shark

Odontaspis ferox by Risso in 1810, there is an increasingoccurrence of records of this species in tropical waters

around the globe. The site of the encounter concurs with pre-vious records, stating that O. feroxis known to occur at insularlocations that shelve steeply into depths of several hundredmetres (Fergusson et al., 2008). The observed female of~400 cm TL was encountered in 10 m depth. The totallength of the shark and its depth are within the range of pre-vious records (Fergusson et al., 2008). However, the fact thatthe first sighting of O. ferox took place at a frequently useddive spot, suggests that the presence of this species in thissite and depth is fairly unusual. Although there is no officialreport on pregnant females in shallow waters, there have

been reasonably frequent sightings of pregnant individualsin depths from 0 to 10 m in protected bays around theCanary Islands for more than a decade (between July andSeptember from 1999 to 2010) (D. Acuna, personal communi-cation, 2012). Even though there is no certainty that thefemale shark observed in the Galapagos was pregnant, theseobservations point to the hypothesis that O. ferox may givebirth in shallow waters. While this is the first confirmed

record of this species in the waters of Galapagos, O. feroxhas been recorded in other insular locations of the TropicalEastern Pacific, such as Malpelo Island, Colombia, andrecently there are unconfirmed but reliable sightings of O.

ferox from the Cocos Island National Park (Fergusson et al.,2008) strengthening the assumption that this species ispresent around the whole Galapagos volcanic province.Thus, the sighting of the specimen in the Galapagos is reason-able and expected. Individuals of O. ferox show a regionalcolour variation (Seigel & Compagno, 1986). The observedindividual from Galapagos was characterized by pigmentationsimilar to the smalltooth sandtiger sharks from Malpelo,Cocos and New South Wales. Those show moderatelyspotted and brownish-grey pigmentation, while specimensfrom the Mediterranean tend to have uneven, black ovalblotches (Fergusson et al., 2008).

In the case of Mustelus albipinnis it is likely, given thehistory of reports for this species, and the three encountersover an 18-month period, that this species is resident at theGalapagos Islands, but that it is rarely found due to itsreported preferred habitat in deeper waters and the absenceof a trawl fishery or other human activities. It seems likelythat the organic residues thrown regularly to the sea by fisher-men at the site of the sightings may have caused the shark toascend to shallow waters where the encounters occurred. Theestimated size of the specimen is significantly greater than themaximum size registered for this species (Perez-Jimenez et al.,

2005). This difference in sizes may be explained by the recentdescription of the species and the limited number of individ-uals (N 42).

Centrophorus squamosus is an important component ofdeep-water fisheries in the north-eastern Atlantic Ocean,being captured at depths between 500 and 1300 m withbottom trawl, longline and fixed bottom nets (Compagno,1984; Banon et al., 2006; Figueiredo et al., 2008; Severinoet al., 2009). Sex-ratio changes significantly depending onthe catch depths, with males dominant at shallower depthsand females in deeper waters (Clarke et al., 2001). There is evi-dence that this species displays complex migratory patternsassociated with its reproduction (Figueiredo et al., 2008).Size at 50% maturity was 125 cm TL for females, with thesmallest mature female at 110 cm TL (Girard & Du Buit,1999; Banon et al., 2006; Figueiredo et al., 2008). Whiledepth of capture and fishing gear match what has beenreported for C. squamosus, the specimen captured inGalapagos, a gravid female measuring 95 cm TL, is signifi-cantly smaller than the smallest mature female registered inthe north-eastern Atlantic. A different size range for theeastern Pacific or the lower latitude of the specimen capturedin Galapagos in comparison with the catches analysed in thenorth-eastern Atlantic could explain these differences, butmore data are needed to verify these hypotheses. The presenceof a gravid female at low latitudes concurs with the latitudinalgradient observed in the north-eastern Atlantic for C. squamo-

sus females, with an increase of pregnant specimens further

Fig. 5. Specimen of Centrophorus squamosus captured south of FernandinaIsland being examined at the Charles Darwin Research Station (photograph:

David Acuna).



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south (Banon et al., 2006). This first record ofC. squamosus inthe eastern Pacific Ocean extends its previously known distri-bution range from the Atlantic, Indo-Pacific and westernPacific to circumglobal.

A C K N O W L E D G E M E N T S

We especially thank Franklin Arreaga, Lenin Barrera, RubenIntriago, Martin Krammer and Martin Schlifski for providingthe specimen and/or the information, photographs and videofootage of the three species recorded in this paper. Withouttheir actions and interest in conservation these three sharkspecies would remain unknown in Galapagos. We thank DrAlberto Brito (Animal Biology Department, University of LaLaguna), Dr John McCosker (Department of AquaticBiology, California Academy of Sciences) and Dr PhillipHeemstra (South African Institute for Aquatic Biodiversity)for corroborating the species identification. Special thanksgo to Shark Savers Germany. Their support of the sharkresearch and conservation project is greatly appreciated. We

thank Aggressor Fleet and Beluga Reisen for their efforts inputting us in contact with two of the observers. This publi-cation is contribution number 2069 of the Charles DarwinFoundation for the Galapagos Islands.

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Correspondence should be addressed to:

D. Acuna-Marrero

Estacion Cientfica Charles Darwin (ECCD)Avenida Charles Darwin s/n, Puerto Ayora, Santa Cruz, IslasGalapagos, Ecuador

email: [email protected]

http://www.iucnredlist.org/http://www.iucnredlist.org/mailto:[email protected]:[email protected]://www.iucnredlist.org/http://www.iucnredlist.org/

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