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

First record of three shark species, Odontaspis ferox, Mustelus albipinnis and Centrophorus squamosus, from pagos Islands the Gala
~ a-marrero1, johanna s. zimmerhackel2, jens mayorga3 and alex hearn4 david acun
1 2

n Cient ca Charles Darwin (ECCD), Avenida Charles Darwin s/n, Puerto Ayora, Santa Cruz, Islas Gala pagos, Ecuador, Estacio t Hamburg, Olbersweg 24, 22767 Hamburg, Germany, Institut fu r Hydrobiologie und Fischereiwissenschaft (IHF), Universita 3 bal, Gala pagos, Ecuador, 4Turtle Island Restoration Network, Olema, CA, Universidad San Francisco de Quito, GAIAS, San Cristo USA

pagos Islands Two shark species, Odontaspis ferox and Centrophorus squamosus, are recorded for the rst time at the Gala based on one sighting reported by tourist divers and one specimen captured by a local sherman. The identication of a third shark species, Mustelus albipinnis, previously unidentied and recorded as Mustelus sp., is achieved based on several sightings at the same location.

pagos Islands, range distribution Keywords: Odontaspis ferox, Mustelus albipinnis, Centrophorus squamosus, Gala
Submitted 15 May 2013; accepted 5 June 2013

INTRODUCTION

pagos Islands and their surrounding Marine Reserve The Gala are situated in the eastern Pacic Ocean, 1000 km to the west of mainland Ecuador covering a total area of approximately 138,000 km2. The islands are located at the conuence of three major ocean currents that show a strong seasonal variation in intensity and direction (Chavez & Brusca, 1991). From May to November the cool Humboldt Current predominates establishing a cool, dry season known as garua, while from December to May its intensity decreases and it is the warm Panama Current that prevails, creating a warm rainy season (Banks, 2002). This complex environmental setting is reected in ve distinct biogeographical regions for subtidal reef communities (Edgar et al., 2002) and an extraordinarily important and diverse marine reserve, recognized by UNESCO as a World Heritage Site (Heylings et al., 2002). pagos sh biodiversity has been studied and reviewed Gala by different authors (McCosker & Rosenblatt, 1984; McCosker, 1987; Grove & Lavenberg, 1997). Long-term subtidal ecological monitoring programmes, local shers and scientic expeditions have been some of the most important sources of new information. In the last published review the pagos stood at number of shore-sh species recorded in Gala 550, including 30 shark species belonging to ten different families (McCosker & Rosenblatt, 2010).

Corresponding author: a-Marrero D. Acun Email: david.acuna@fcdarwin.org.ec

The smalltooth sandtiger shark Odontaspis ferox (Risso, 1810) is reported as having a circumglobal but fragmented distribution between 408N and 408S, mostly in warmtemperate and tropical waters (Compagno, 1984; Bonl, 1995). Odontaspis ferox is an offshore shark of deep waters, observed or caught mainly near the seabed, close to continental shelves and upper slopes at depths ranging from 10 to 883 m (Fergusson et al., 2008). The size of O. ferox at maturity differs in sexes, with ~275 cm for males and 365 cm total length (TL) for females, with the largest recorded size of a female measuring 450 cm TL (Compagno et al., 2005). Similar to Carcharias taurus, O. ferox is presumed to have a very low reproductive capacity, producing only two pups every two years and possibly practises uterine cannibalism in the form of oophagy (Compagno, 2001). This shark is shed with bottom gill-nets, line gear and bottom trawls principally in Japan, the Mediterranean Sea and southern Africa off Madagascar (Abe et al., 1968; Cooke, 1997). Although O. ferox is mostly discarded or utilized by-catch, there is a demand for its large oily liver in Japan (Abe et al., 1968). The likely low reproductive capacity and natural small numbers of this species makes it potentially sensitive to local extinction, even at small capture rates (Fergusson et al., 2008), and it is listed as Vulnerable in the International Union for Conservation of Nature and Natural Resources Red List of Threatened Species (IUCN RL). This pagos. is the rst record of O. ferox in the waters of Gala Mustelus albipinnis was described by Castro-Aguirre et al. (2005) as being one of two un-named species of smoothhound shark known to be found from the Gulf of California to Ecuador (P. Heemstra in Compagno, 1988 and in McCosker & Rosenblatt, 2010). Smoothhound sharks are generally found on the continental shelves in temperate and tropical
1

~ a-marrero et al . david acun

waters around the world, yet M. mento and Triakis maculata pagos Islands are both also reported as present in the Gala (McCosker & Rosenblatt, 2010). As a recently-described species, little is known about its biology and ecology. It is thought to be benthopelagic, with a reported depth distrirez-Jime nez et al., 2005). Both bution of 30 281 m (Pe males and females mature at 90 100 cm TL, and the largest reported size was a 118 cm TL female caught in the Gulf of rez-Jime nez et al., 2005). Its reproduction California (Pe mode is by placental viviparity, with brood sizes ranging rez-Jime nez et al., 2005). This species from 3 to 23 pups (Pe is caught in the Gulf of California, where smoothhounds may represent 79% of the total catch by the local artisanal rquez-Farias, 2000), but due to lack of shark shery (Ma proper identication until the recent description of the species, its status has not been assessed. This shark was prepagos as Mustelus sp. (McCosker & viously recorded in Gala Rosenblatt, 2010) but has now been identied to species level. The leaf-scale gulper shark, Centrophorus squamosus (Bonnaterre, 1788), is a deep-water squaloid species which is found in the eastern Atlantic Ocean, the Indian Ocean and the western Pacic Ocean, where it has been recorded from Japan to New Zealand (Girard & Du Buit, 1999; Compagno ssimo et al., 2012). This benthopelagic et al., 2005; Ver shark inhabits the continental slopes at depths between 230 and 2400 m (Compagno & Niem, 1998). Size at maturity is greater in females (124 cm TL) than in males (86 cm TL) (Girard & Du Buit, 1999). It is a lecithotrophic viviparous species, according to the recent classication proposed by Hamlett et al. (2005). Centrophorus squamosus is an important component of deep-water longlining and bottom trawling sheries in the eastern Atlantic Ocean due to the value of its

liver and esh (Severino et al., 2009). Catches have decreased signicantly during the last decade and this shark is listed as Vulnerable in the IUCN RL (White, 2003). This rst record pagos Islands extends its known of C. squamosus in the Gala range into the eastern Pacic. This paper contributes to the development of the Species Checklist of the Charles Darwin Foundationan attempt to pagos establish a complete biodiversity catalogue for the Gala Islands (Bungartz et al., 2009). It also highlights the contribution that citizen science can make, in the form of photographs and specimens provided by local residents, shers and tourists.

MATERIALS AND METHODS

Each shark species was observed in a different biogeographical region: Wolf Island in the far north (Odontaspis ferox), San bal Island in the centre-south (Mustelus albipinnis) Cristo and Fernandina Island in the west (Centrophorus squamosus). Shark length was estimated visually for O. ferox and M. albipinnis, but measured in the case of C. squamosus. Shark Bay (1823.04 N 91848.46 W) is located at the east of Wolf Island (Figure 1). It is a rocky reef exposed to the predominant south-east current with a shallow narrow platform followed by a steep drop-off to about 600 m. The seabed here is mainly composed of big boulders with small sandy patches and abundant coral formations. Photograph and video footage were taken during the only sighting recorded so far. Tongo Reef (0854.19 S 89837.51 W) is a shallow at rocky platform that extends to about 300 m off the coast at the west bal Island (Figure 1). The border of this corner of San Cristo

pagos Archipelago with the islands of Wolf, San Cristo bal, Fernandina and Isabela indicated; (B) map of San Cristo bal Island showing Fig. 1. (A) Map of the Gala the location of the Mustelus albipinnis new record at Tongo Reef; (C) map of Wolf Island showing the location of the Odontaspis ferox new record at Shark Bay; (D) map of Fernandina and Isabela Islands showing the location of the Centrophorus squamosus new record south to Punta Mangle.

 pagos islands first record of three shark species from the gala

platform drops suddenly to depths over 50 m. The site of the encounters is less than 100 m from the coastline. Photograph and video footage were taken during the rst sighting. On 14 January 2012, one specimen of Centrophorus squamosus was caught off the east coast of Fernandina Island (0833.38 S 91824.23 W). The shark was captured at approximately six miles south of Punta Mangle, at the south-eastern side of Fernandina Island (Figure 1) using a 200 hundred hooks longline set near the bottom, at 600 m depth. The bait used was black-striped salema (Xenocys jessiae). This information was provided by the sherman and obtained using a depth-sounder. The shark was brought to the Charles Darwin Research Station, where it was identied a. and examined by biologist David Acun

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

RESULTS

n than to ventral n. The identication is additionally supported by previous personal sightings by the author (D.A.), who observed this species several times in the Mar de las Calmas Marine Reserve, Hierro Island, Canary Islands, Spain.

systematics
Order LAMNIFORMES Family ODONTASPIDIDAE Mu ller & Henle, 1839 Genus Odontaspis Risso, 1826 Odontaspis ferox Risso, 1810 One unknown shark was observed in the late afternoon on 22 November 2010 by dive tourists Martin Krammer and Martin pagos (1823.04 N Schlifski at Sharks Bay, Wolf Island, Gala 91848.46 W). Both divers videotaped and photographed the shark because of its uncommon appearance (Figures 2 & 3). The individual was a female of ~400 cm TL, swimming from deep waters towards the surface along the rocky bottom with sandy patches in a protected area. The encounter lasted approximately two minutes and the shark was swimming slowly away when divers approached it. The divers reported that it may have been pregnant, given its distended abdomen. Photographs and videos were shared with dive guides Lenin Barrera and Ruben Intriago, and given to the Charles Darwin Foundation (CDF) where scientists Johanna aherrera and David Acun a identZimmerhackel, Cesar Pen ied the specimen as the smalltooth sandtiger shark Odontaspis ferox due to the following features: (i) anal n present; (ii) two dorsal ns and ve gill slits; (iii) eyes situated over mouth; (iv) caudal n asymmetrical without keel; (v) subterminal mouth with large bladelike anterior teeth; and (vi) long conical snout and rst dorsal n closer to pelvic Order CARCHARHINIFORMES Family TRIAKIDAE Gray, 1851 Genus Mustelus Linck, 1790 Mustelus albipinnis Castro-Aguirre, Antuna-Mendiola, lez-Acosta & de la Cruz-Agu Gonza ero, 2005 Local student Jens Mayorga reported seeing an unusual shark bal, undertakwhile SCUBA diving off the coast of San Cristo ing a study of shark presence at Tongo Reef, where shers regularly clean their catch. He reported three encounters: the rst took place on 11 September 2010; the second on 11 September 2011; and the third on 10 April 2012. Each encounter was with a single, apparently female individual, which he estimated to be between 150 and 200 cm TL (signicantly larger than the reported maximum length for this species), swimming slowly in approximately 10 m depth (signicantly shallower than the shallowest depth reported for this species). All three encounters took place around dusk, and the rst occurred ten minutes after a shing boat started to clean its catch at the site. The rst encounter lasted over twenty minutes, with the shark showing signs of curiosity towards the diver, who at that time was alone. It slowly circled the diver several times over this period. During the remaining two encounters, when the diver was accompanied by a buddy, the shark appeared more active, and only circled the divers once. Photographs (Figure 4) and video stills (taken during the rst encounter) were sent to John McCosker (California Academy of Science) and Phil Heemstra (South African Institute for Aquatic Biodiversity), both of whom identied the shark as Mustelus albipinnis. Mustelus albipinnis can 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 lmed by Martin Krammer and pagos northern islands). Martin Schlifski during a diving trip to the Gala

Fig. 4. Mustelus albipinnis swimming in shallow water at Tongo Reef, San bal Island (photograph extracted from video footage lmed by Jens Cristo Mayorga).

~ a-marrero et al . david acun

pectoral, pelvic and anal ns, being the only known houndshark with this feature (Castro-Aguirre et al., 2005). Order SQUALIFORMES Compagno, 1973 Family CENTROPHORIDAE Bleeker, 1859 Genus Centrophorus Mu ller & Henle, 1837 Centrophorus squamosus Bonnaterre, 1788 Identication as Centrophorus squamosus was made by the following main features: (i) body cylindrical and low; (ii) ve pairs of gills and spiracles present; (iii) no anal n, n spines present on both dorsal ns and caudal n with subterminal notch; (iv) bladelike unicuspidate teeth in upper and lower jaws, with lowers much larger; and (v) rear tips of pectoral ns broadly angular and not strongly extended. A set of pictures of the shark were sent to Alberto Brito (University of La Laguna, Spain) who conrmed the species identication as Centrophorus squamosus (A. Brito, personal communication, 2012) (Figure 5). The individual examined was a mature female, measuring 95 cm TL, and weighting 5.2 kg. Five yolk balls were found within the uterus and embryos were not distinguished.
DISCUSSION

Odontaspis ferox and Centrophorus squamosus constitute two pagos Marine Reserve new records of shark species in the Gala and two new families not previously registered: Odontaspidae and Centrophoridae. The species listed until now as Mustelus sp. (McCosker & Rosenblatt, 2010) is ultimately identied as Mustelus albipinnis. Since the description of the smalltooth sandtiger shark Odontaspis ferox by Risso in 1810, there is an increasing occurrence of records of this species in tropical waters around the globe. The site of the encounter concurs with previous records, stating that O. ferox is known to occur at insular locations that shelve steeply into depths of several hundred metres (Fergusson et al., 2008). The observed female of ~400 cm TL was encountered in 10 m depth. The total length of the shark and its depth are within the range of previous records (Fergusson et al., 2008). However, the fact that the rst sighting of O. ferox took place at a frequently used dive spot, suggests that the presence of this species in this site and depth is fairly unusual. Although there is no ofcial report on pregnant females in shallow waters, there have

Fig. 5. Specimen of Centrophorus squamosus captured south of Fernandina Island being examined at the Charles Darwin Research Station (photograph: a). David Acun

been reasonably frequent sightings of pregnant individuals in depths from 0 to 10 m in protected bays around the Canary Islands for more than a decade (between July and a, personal communiSeptember from 1999 to 2010) (D. Acun cation, 2012). Even though there is no certainty that the pagos was pregnant, these female shark observed in the Gala observations point to the hypothesis that O. ferox may give birth in shallow waters. While this is the rst conrmed pagos, O. ferox record of this species in the waters of Gala has been recorded in other insular locations of the Tropical Eastern Pacic, such as Malpelo Island, Colombia, and recently there are unconrmed but reliable sightings of O. ferox from the Cocos Island National Park (Fergusson et al., 2008) strengthening the assumption that this species is pagos volcanic province. present around the whole Gala pagos is reasonThus, the sighting of the specimen in the Gala able and expected. Individuals of O. ferox show a regional colour variation (Seigel & Compagno, 1986). The observed pagos was characterized by pigmentation individual from Gala similar to the smalltooth sandtiger sharks from Malpelo, Cocos and New South Wales. Those show moderately spotted and brownish-grey pigmentation, while specimens from the Mediterranean tend to have uneven, black oval blotches (Fergusson et al., 2008). In the case of Mustelus albipinnis it is likely, given the history of reports for this species, and the three encounters over an 18-month period, that this species is resident at the pagos Islands, but that it is rarely found due to its Gala reported preferred habitat in deeper waters and the absence of a trawl shery or other human activities. It seems likely that the organic residues thrown regularly to the sea by shermen at the site of the sightings may have caused the shark to ascend to shallow waters where the encounters occurred. The estimated size of the specimen is signicantly greater than the rez-Jime nez et al., maximum size registered for this species (Pe 2005). This difference in sizes may be explained by the recent description of the species and the limited number of individuals (N 42). Centrophorus squamosus is an important component of deep-water sheries in the north-eastern Atlantic Ocean, being captured at depths between 500 and 1300 m with bottom trawl, longline and xed bottom nets (Compagno, n et al., 2006; Figueiredo et al., 2008; Severino o 1984; Ban et al., 2009). Sex-ratio changes signicantly depending on the catch depths, with males dominant at shallower depths and females in deeper waters (Clarke et al., 2001). There is evidence that this species displays complex migratory patterns associated with its reproduction (Figueiredo et al., 2008). Size at 50% maturity was 125 cm TL for females, with the smallest mature female at 110 cm TL (Girard & Du Buit, n et al., 2006; Figueiredo et al., 2008). While o 1999; Ban depth of capture and shing gear match what has been reported for C. squamosus, the specimen captured in pagos, a gravid female measuring 95 cm TL, is signiGala cantly smaller than the smallest mature female registered in the north-eastern Atlantic. A different size range for the eastern Pacic or the lower latitude of the specimen captured pagos in comparison with the catches analysed in the in Gala north-eastern Atlantic could explain these differences, but more data are needed to verify these hypotheses. The presence of a gravid female at low latitudes concurs with the latitudinal gradient observed in the north-eastern Atlantic for C. squamosus females, with an increase of pregnant specimens further

 pagos islands first record of three shark species from the gala

n et al., 2006). This rst record of C. squamosus in o south (Ban the eastern Pacic Ocean extends its previously known distribution range from the Atlantic, Indo-Pacic and western Pacic to circumglobal.

Centrophorus squamosus from west of Ireland and Scotland. Journal of the Marine Biological Association of the United Kingdom 81, 10191029. Compagno L.J.V. (1984) Hexanchiformes to Lamniformes; Carcharhiniformes. In Compagno J.L.V. (ed.) Sharks of the world. An annotated and illustrated catalogue of shark species known to date. Volume 4. Rome: FAO, pp. 1 249; pp. 251265. Compagno L.J.V. (1988) Sharks of the order Carcharhiniformes. Princeton, NJ: Princeton University Press. Compagno L.J.V. (2001) Bullhead, mackerel and carpet sharks (Heterodontiformes, Lamniformes and Orectolobiformes). In Compagno L.J.V. (ed.) Sharks of the world. An annotated and illustrated catalogue of shark species known to date. Volume 2. Rome: FAO, pp. 1269. Compagno L.J.V. and Niem V.H. (1998) Squalidae. In Carpenter K.E. and Niem V.H. (eds) The living marine resources of the Western Central Pacic. Volume 2. Cephalopods, crustaceans, holothurians and sharks. Rome: FAO, pp. 12131232. Compagno L.J.V., Fowler S. and Dando M. (2005) Sharks of the world. Princeton eld guides. Princeton, NJ: Princeton University Press. Cooke A.J. (1997) Survey of elasmobranch sheries and trade in Madagascar. In Marshall N.T. and Barnett R. (eds) The trade in sharks and shark products in the Western Indian and Southern Indian and South East Atlantic Oceans. Nairobi: TRAFFIC East/ Southern Africa, pp. 101130. nez C. and Banks S. (2002) a J.M., Calvopin a M., Mart Edgar G.J., Farin Comunidades submareales rocosas II: peces y macroinvertebrados viles. In Danulat E. and Edgar G.J. (eds) Reserva Marina de mo pagos, pagos. L nea Base de la Biodiversidad. Santa Cruz, Gala Gala n Charles Darwin/Servicio Parque Nacional Ecuador: Fundacio pagos, pp. 2235. Gala Fergusson I.K., Graham K.J. and Compagno L.J.V. (2008) Distribution, abundance and biology of the smalltooth sandtiger shark Odontaspis ferox (Risso, 1810) (Lamniformes: Odontaspididae). Environmental Biology of Fishes 81, 207228. Figueiredo I., Moura T., Neves A. and Gordo L.S. (2008) Reproductive strategy of leafscale gulper shark Centrophorus squamosus and the Portuguese dogsh Centroscymnus coelolepis on the Portuguese continental slope. Journal of Fish Biology 73, 206225. Girard M. and Du Buit M.H. (1999) Reproductive biology of two deepwater sharks from the British Isles, Centroscymnus coelolepis and Centrophorus squamosus (Chondrichthyes: Squalidae). Journal of the Marine Biological Association of the United Kingdom 79, 923931. pagos Islands. Grove J.S. and Lavenberg R.J. (1997) Fishes of the Gala Stanford, CA: Stanford University Press. Hamlett W.C., Kormanik G., Storrie M., Stevens B. and Walker T.I. (2005) Chondrichthyan parity, lecithotrophy and matrotrophy. In Hamlett W.C. (ed.) Reproductive biology and phylogeny of Chondrichthyes: sharks, batoids and chimaeras. Eneld, NH: Science Publishers, pp. 395 434. n e Heylings P., Bensted-Smith R. and Altamirano M. (2002) Zonicacio pagos. In Danulat E. and Edgar historia de la Reserva Marina de Gala pagos. L nea Base de la G.J. (eds) Reserva Marina de Gala pagos, Ecuador: Fundacio n Charles Biodiversidad. Santa Cruz, Gala pagos, pp. 1021. Darwin/Servicio Parque Nacional Gala rquez-Farias F. (2000) Tiburones del Golfo de California. In Ma xico: evaluacio n y manejo Sustentabilidad y pesca responsable en Me 19992000. Mexico: INP, SEMARNAP, pp. 237 257. pagos Islands. Oceanus 30, McCosker J.E. (1987) The shes of the Gala 2832.

ACKNOWLEDGEMENTS

We especially thank Franklin Arreaga, Lenin Barrera, Ruben Intriago, Martin Krammer and Martin Schlifski for providing the specimen and/or the information, photographs and video footage of the three species recorded in this paper. Without their actions and interest in conservation these three shark pagos. We thank Dr species would remain unknown in Gala Alberto Brito (Animal Biology Department, University of La Laguna), Dr John McCosker (Department of Aquatic Biology, California Academy of Sciences) and Dr Phillip Heemstra (South African Institute for Aquatic Biodiversity) for corroborating the species identication. Special thanks go to Shark Savers Germany. Their support of the shark research and conservation project is greatly appreciated. We thank Aggressor Fleet and Beluga Reisen for their efforts in putting us in contact with two of the observers. This publication is contribution number 2069 of the Charles Darwin pagos Islands. Foundation for the Gala

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Correspondence should be addressed to: a-Marrero D. Acun n Cient ca Charles Darwin (ECCD) Estacio Avenida Charles Darwin s/n, Puerto Ayora, Santa Cruz, Islas pagos, Ecuador Gala email: david.acuna@fcdarwin.org.ec