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Journal of Fish Biology (2014) doi:10.1111/jfb.12517, available online at wileyonlinelibrary.com Not all fish are equal: functional biodiversity of cartilaginous fishes (Elasmobranchii and Holocephali) in Chile C. Bustamante*†‡, C. Vargas-Caro*† and M. B. Bennett* *School of Biomedical Sciences, The University of Queensland, St Lucia, Qld 4072, Australia and †Programa de Conservación de Tiburones (Chile), Valdivia, Chile (Received 27 November 2013, Accepted 29 July 2014) A review of the primary literature on the cartilaginous fishes (sharks, skates, rays and chimaeras), together with new information suggests that 106 species occur in Chilean waters, comprising 58 sharks, 30 skates, 13 rays and five chimaeras. The presence of 93 species was confirmed, although 30 species were encountered rarely, through validated catch records and sightings made in artisanal and com- mercial fisheries and on specific research cruises. Overall, only 63 species appear to havearange distribution that normally includes Chilean waters. Actual reliable records of occurrence are lacking for 13 species. Chile has a cartilaginous fish fauna that is relatively impoverished compared with the global species inventory, but conservative compared with countries in South America with warm-temperate waters. The region of highest species richness occurs in the mid-Chilean latitudes of c. 30–40∘ S. This region represents a transition zone with a mix of species related to both the warm-temperate Peru- vian province to the north and cold-temperate Magellan province to the south. This study provides clarification of species occurrence and the functional biodiversity of Chile’s cartilaginous fish fauna. © 2014 The Fisheries Society of the British Isles Key words: checklist; chimaera; shark; skate; south-east Pacific Ocean; species richness. INTRODUCTION The influence of sharks, rays, skates (class Elasmobranchii) and chimaeras (class Holocephali) in ecosystems is under increasing threat owing to continuing population declines of many species as a result of overfishing and by-catch mortality (Stevens et al., 2000; Ferretti et al., 2010). Knowledge of national and regional elasmobranch and holocephalan biodiversity is necessary to formulate management plans and con- servation actions (Compagno, 1999; CPPS, 2009). Species richness is one of the most commonly used indices for assessing biodiversity in ecological studies (Whittaker, 1972; Magurran, 2003; Sahney et al., 2010), but in some cases even this basic infor- mation lacks scientific rigour and leads to interpretation errors when management and conservation measures are implemented (Novacek, 2001; Mejía-Falla et al., 2007; Rozzi et al., 2008). This paper reviews Chile’s national elasmobranch and holocephalan species inven- tory. Existing records, new descriptions and recent taxonomic changes at the species ‡Author to whom correspondence should be addressed. Tel.: +61 (7) 33467975 email: c.bustamantediaz@ uq.edu.au 1 © 2014 The Fisheries Society of the British Isles 2 C. BUSTAMANTE ET AL. level are examined through application of specific criteria, to assess the functional diversity and distributions of these cartilaginous fishes within the south-eastern Pacific Ocean. Species are highlighted where there are taxonomic issues, or records are rare or of uncertain validity. The resulting validated species inventory contributes to an under- standing of the diversity of cartilaginous fishes off the coast of South America that should aid future management and conservation planning. MATERIALS AND METHODS Historic and recent literature was reviewed for any reference to cartilaginous fishes (Class Elasmobranchii and Holocephali, sensu Eschmeyer & Fong, 2011) in Chilean waters that would complement the species inventories of Pequeño (1989, 1997) and Lamilla & Bustamante (2005). Additionally, an extensive survey comprising 10 localities along the Chilean coast was con- ducted in 2009 to assess cartilaginous fish by-catch in coastal artisanal (medium and small scale) fisheries and semi-industrial oceanic fleets. Descriptions of the variables surveyed and adetailed sampling methodology are provided by Lamilla et al. (2010). On-board operations and landings were observed in commercial fisheries with a history of interaction; where interaction is defined as an ‘historical or recent record of catch and landing of at least one cartilaginous fish during a commercial trip’ (Lamilla et al., 2010). For each landing site and on-board survey, all carti- laginous fishes caught and landed were recorded and identified to species. Localities alongthe Chilean coast [Fig. 1(a)] and the sampling effort for landing site visits and on-board surveys are provided in Table I. Table II presents information about each fishery evaluated during this research. The 10 localities span c.33∘ of latitude and represent four zones (north, central, south and aus- tral south) that are independent of bathymetric reference and the biographic provinces described by Camus (2001). North includes localities between Iquique (20⋅2∘ S) and Caldera (27⋅0∘ S), central localities between Coquimbo (29⋅9∘ S) and Valparaiso (33⋅0∘ S), south localities between Concepción (36⋅9∘ S) and Puerto Montt (41⋅5∘ S) and austral south includes localities between Aysén (45⋅4∘ S) and Punta Arenas (53⋅1∘ S). The oceanic islands zone is treated separately and includes species reported around insular territories within the Chilean exclusive economic zone, such as Nazca and Sala y Gómez submarine ridges, Juan Fernández Islands, Desventuradas Islands and Easter Island [Fig. 1(b)]. To assess cartilaginous fishes of Chile on a meaningful basis, specific criteria are usedto ensure that only those species reported and confirmed within circumscribed geographical areas within the political boundaries are considered (Mejía-Falla et al., 2007). For this purpose, con- firmed species are defined as those that meet the following criteria: (1) identification hasbeen validated scientifically, and there are no reasonable doubts about a species’ identity, (2) thereare reports of its presence, capture and sale from fishing or scientific expeditions and (3) records do not indicate a haphazard or anecdotal encounter of a species that has a low natural abundance. The modern classification of extant Elasmobranchii and Holocephali and comparative values of global biodiversity were extracted from Eschmeyer & Fong (2011) and Ebert et al. (2013). RESULTS A total of 106 cartilaginous species occur in Chilean waters (58 sharks, 30 skates, 13 rays and five chimaeras), represented in 58 genera (37 shark, nine skate, nine ray and three chimaera genera) and 29 families (16 shark, six skate, four ray and three chimaera families). This represents an increase in the reported biodiversity of cartilaginous fishes in Chile with 14 species added to the known ichthyofauna, these are brown lanternshark Etmopterus unicolor (Engelhardt 1912), pocket shark Mollisquama parini Dolganov 1984, frog shark Somniosus longus (Tanaka 1912), © 2014 The Fisheries Society of the British Isles, Journal of Fish Biology 2014, doi:10.1111/jfb.12517 BIODIVERSITY OF CARTILAGINOUS FISHES IN CHILE 3 Fig. 1. Map of South America showing the location of study area (inset boxes), indicating (a) main localities in Chile, zones and biogeographic provinces on the latitudinal gradient and (b) oceanic islands zone. Contour lines ( ) indicate 2000 m isobath on (b). Biogeographic provinces modified from Camus (2001). smalltooth sand tiger shark Odontaspis ferox (Risso 1810), longfin mako Isurus paucus Guitart 1966, crocodile shark Pseudocarcharias kamoharai (Matsubara 1936), silky shark Carcharhinus falciformis (Müller & Henle 1839), scalloped hammerhead Sphyrna lewini (Griffith & Smith 1834), Chilean guitarfish Tarsistes philippii Jordan 1919, joined-fins skate Bathyraja cousseauae Díaz de Astarloa & Mabragaña 2004, dark-belly skate Bathyraja meridionalis Stehmann 1987, cuphead skate Bathyraja scaphiops (Norman 1937), spinetail mobula Mobula japanica (Müller & Henle 1841) and eastern Pacific black ghostshark Hydrolagus melanophasma James, Ebert, Long & Didier 2009. While this total of 106 species is indicative of overall diversity, it is not a good index of functional biodiversity as many species identifications are based on anecdotal or single verified observations, and as such, information about their distributions is poor or absent. A total of 6450 specimens of sharks, rays, skates and chimaeras were examined from 138 landing and 286 on-board surveys, which confirmed the presence of 40 cartilagi- nous fish species. Bibliographic records provide a further 53 species that meetthe confirmed species criterion resulting in 93 confirmed species, of which 30 speciesare © 2014 The Fisheries Society of the British Isles, Journal of Fish Biology 2014, doi:10.1111/jfb.12517 4 C. BUSTAMANTE ET AL. Table I. Localities of landing sites and bases for on-board surveys with an indication of relative observer effort. Days-at-sea observed during on-board surveys are indicated in parentheses Locality Zone Landing site visits On-board surveys Iquique North 26 33 (46) Antofagasta North 0 9 (15) Caldera North 7 30 (69) Coquimbo Central 2 48 (68) Valparaíso Central 1 39 (45) Concepción South 70 24 (36) Valdivia South 18 76 (88) Puerto Montt South 10 21 (52) Aysén Austral south 0 6 (27) Punta Arenas Austral south 4 0 relatively rarely encountered. The situation for the other 13 species reported to occur in Chilean waters is less clear. The species reported in Chilean waters and