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Home , Aetobatus laticeps, Aetomylaeus, Apristurus, Brown catshark, Bullhead shark, California butterfly ray

Regular paper

Updated checklist of marine cartilaginous from continental and insular (Tropical Eastern Pacific Ocean)

by

Marcos Douglas Calle-Morán (1) & Philippe Béarez* (2)

Abstract. – Ecuador, located on the central-eastern edge of the Pacific Ocean, has a high level of biological diversity, including no less than 800 of marine . The ecological and economic importance, as well as the vulnerability of cartilaginous fish along the coasts of Ecuador and the Galapagos Islands, justify the need to produce an updated list of this group. Marine fish are generally listed separately and no updated national list has been produced recently. The objective of this work is to present an updated list of the continental and insu- lar chondrichthyans of Ecuador based on field observations in the fishing ports of Santa Rosa de Salinas (Santa Elena province), Puerto López and Manta (Manabí province), alongside a review of the specialized literature. This group of fish includes 12 orders, 39 families, 65 genera, and 119 species. Of these, 64 are , 48 are rays, and 7 are .

© SFI Submitted: 17 Nov. 2019 Résumé. – Liste actualisée des poissons marins cartilagineux de l’Équateur continental et insulaire (océan Paci- Accepted: 30 Apr. 2020 fique tropical est). Editor: R. Causse L’Équateur, situé sur la bordure centre-est de l’océan Pacifique, présente un niveau élevé de diversité bio- logique et notamment pas moins de 800 espèces de poissons marins. L’importance écologique et économique, ainsi que la vulnérabilité des poissons cartilagineux le long des côtes de l’Équateur et des îles Galápagos justi- Key words fient la nécessité de produire une liste actualisée de ce groupe. Les poissons marins sont généralement répertoriés Sharks séparément dans ces deux territoires et aucune liste nationale n’a été produite récemment. L’objectif de ce travail Rays est de présenter une liste actualisée des chondrichtyens continentaux et insulaires de l’Équateur, basée sur des Chimaeras observations de terrain dans les ports de pêche de Santa Rosa de Salinas (province de Santa Elena), Puerto López et Manta (province de Manabí) et sur une revue de la littérature spécialisée. Dans cette zone du Pacifique, le Mainland Ecuador groupe des chondrichthyens comprend 12 ordres, 39 familles, 65 genres et 119 espèces. Parmi celles-ci, 64 sont Galápagos Islands des requins, 48 des raies et 7 des chimères.

INTRODUCTION Until now, the number of species for continental Ecuador was considered to be 101: 61 sharks, 33 rays, and 7 chimae- Ecuador is a small country on the Pacific side of South ras (Martínez-Ortiz and García-Domínguez, 2013), while for America, whose coast stretches approximately 1250 km the Galápagos Islands it was 56: 34 sharks, 20 rays, and at between 1°28’N and 3°23’S. The Galápagos Islands are least 2 chimaeras (Hearn et al., 2014; McCosker and Rosen- located on the equator in the Pacific Ocean, around 1000 km blatt, 2010). Of these chondrichthyans, 43 are distributed west of Guayaquil. The coasts of mainland Ecuador and the across both areas. Galápagos Islands are connected by the Carnegie Ridge, an This group of fish is important because of its ecological underwater ridge with depths up to 2500 m. Water circula- role in the health and stability of marine ecosystems. Many tion along the Ecuadorian coast is directly influenced by the species of sharks are top predators of the marine ecosystems tropical currents coming from the Panamic basin and the and therefore regulate the population of their prey (Stevens northern branch of the Humboldt Current, which brings cold et al., 2000). In addition to providing ecological services, waters from the south. The fact that Ecuador is located on an elasmobranchs also provide food and have a commercial oceanic front zone, with the presence of upwellings, explains value because their generate foreign exchange and the mixed fauna, such as tropical species from the north and both direct and indirect employment in the country. During temperate species from the south. This oceanographic con- the period from September 2007 to December 2011, nearly figuration makes Ecuador an area of particularly rich marine 113,000 tons of sharks and rays were landed in Ecuadorian biodiversity. fishing ports (Subsecretaría de Recursos Pesqueros, 2012), Of the 1,464 chondrichthyan species compiled by Fricke including 30 species of sharks and 4 species of rays. How- et al. (2020), 598 are sharks, 810 rays and 56 chimaeras. ever, due to their biological characteristics (slow growth,

(1) Programa de Doctorado en Ciencias con mención en Recursos Acuáticos, Facultad de Ciencias del Mar, Universidad Autónoma de Sinaloa, Paseo Claussen S/N, CP 80000, Mazatlán, México. [[email protected]] (2) UMR 7209 AASPE, CNRS-MNHN, Muséum national d’Histoire naturelle, CP 56, 55 rue Buffon, 75005 Paris, France. [[email protected]] * Corresponding author

Cybium 2020, 44(3): 239-250. https://doi.org/10.26028/cybium/2020-443-004 Checklist of marine cartilaginous fishes from continental and insular Ecuador Ca l l e -Mo r á n & Bé a r e z late maturity, and low fecundity), chondrichthyans are vul- names in Spanish and English. Information is given about nerable to overfishing, as well as loss and marine distribution in Ecuador (i.e. mainland and/or Galápagos) and pollution (Dulvy et al., 2014). In addition to the targeted IUCN Red list category (NE for Not Evaluated, DD for Data fishing of sharks by surface longlines, has a major Deficient, LC for Least Concern, NT for Near Threatened, impact on the catch of demersal elasmobranchs, mainly VU for Vulnerable, EN for Endangered, CR for Critically for , dogfish, , rays, and Endangered). Species recorded during the survey are indi- (Moreno-Vera et al., 2018). cated with “pers. obs.” (personal observation); references The last chondrichthyans survey for mainland Ecuador from the literature are given for the others. was done in 2013 (Martínez-Ortiz and García-Domínguez, 2013); it was in 2014 for the Galápagos Islands (Hearn et al., 2014). This information has generally been compiled sepa- RESULTS rately; therefore, obtaining updated information and a more accurate list of species was deemed necessary. The objective A total of 119 chondrichthyan species were recorded for of this work was to produce an updated list of cartilaginous both the mainland and the islands of Ecuador. Of these 119 fish, with up-to-date nomenclature, for both continental and species, there were 64 sharks (54%), 48 rays (40%), and 7 insular Ecuador, in the Central Eastern Pacific Ocean. chimaeras (6%), distributed within 65 genera, 39 families, and 12 orders. A total of 52 species (44%) were recorded by the authors; the rest were obtained from the specialized lit- MATERIAL AND METHODS erature.

Between 2007 and 2018, the fishing ports of Manta, Subclass Puerto López (Manabí Province) and Santa Rosa (Santa Cohort Selachii Elena Province) were surveyed (Fig. 1). Manta was surveyed Order during April 2014, 2015, 2016, and 2017; Puerto López from Hexanchidae November 2007 to June 2008; and Santa Rosa from Febru- 1. griseus (Bonnaterre, 1788) – Cañabota ary 2008 to January 2009, from June 2012 to May 2013, gris – Bluntnose sixgill and from June 2017 to May 2018. It should be noted that Mainland. Ebert (2016). NT Manta is the main national fish landing port and Santa Rosa 2. cepedianus (Péron, 1807) – Cañabota is the second. For the Galápagos Islands, it was not possible gata – Broadnose sevengill shark to obtain information in the field, so the information used in Mainland. Pers. obs., Béarez (1996), Jiménez-Pra- this study was obtained from the specialized literature such do and Béarez (2004), Martínez-Ortiz and García- as Grove and Lavenberg (1997), McCosker and Rosen- Domínguez (2013). DD blatt (2010), McCosker et al. (2012), Acuña-Marrero et al. (2013), Hearn et al. (2014), Tirado-Sánchez et al. (2016). Order Heterodontiformes Van der Laan et al. (2020) was used for the classifica- Family Heterodontidae tion and Fricke et al. (2020) for the nomenclature. For the 3. Heterodontus francisci (Girard, 1855) – Dormilón systematic listing, we used the following sequence: subclass, cohort, order, family, species scientific name and common cornudo – Mainland. Pers. obs., Béarez (1996), Martínez-Ortiz and García-Domínguez (2013). DD 4. Heterodontus mexicanus Taylor & Castro-Aguirre, 1972 – Dormilón búfalo – Mexican hornshark Mainland/Galápagos. Pers. obs., Béarez (1996), Jiménez-Prado and Béarez (2004), Martínez-Ortiz and García-Domínguez (2013), Tirado-Sánchez et al. (2016). DD 5. Heterodontus quoyi (Fréminville, 1840) – Dormilón de Galápagos – Galápagos Mainland/Galápagos. Béarez (1996), McCosker and Rosenblatt (2010), Martínez-Ortiz and García- Figure 1. – Geographical location of the study areas (Manta, Puerto López, Santa Rosa and Galápagos Islands) from Ecuador, Central Domínguez (2013), Hearn et al. (2014), Tirado- Eastern Pacific Ocean. Sánchez et al. (2016). DD

240 Cybium 2020, 44(3) Ca l l e -Mo r á n & Bé a r e z Checklist of marine cartilaginous fishes from continental and insular Ecuador

Order Orectolobiformes Mainland. Pers. obs., Martínez-Ortiz and García- Family Rhincodontidae Domínguez (2013). EN 6. Rhincodon typus Smith, 1828 – Tiburón ballena – Family Megachasmidae Mainland/Galápagos. Pers. obs., Béarez (1996), 14. Megachasma pelagios Taylor, Compagno & Struh- McCosker and Rosenblatt (2010), Martínez-Ortiz and saker, 1983 – Tiburón bocudo – Megamouh shark García-Domínguez (2013), Hearn et al. (2014), Tira- Mainland. Martínez-Ortiz and García-Domínguez do-Sánchez et al. (2016). EN (2013), Martínez-Ortiz et al. (2017). LC

Family Family Cetorhinidae 7. Ginglymostoma unami Del Moral-Flores, Ramíz- 15. Cetorhinus maximus (Gunnerus, 1765) – Tiburón Antonio, Angulo & Pérez-Ponce de León, 2015 – Gata peregrino – nodriza – Pacific Mainland/Galápagos. Béarez (1996), Martínez-Ortiz Mainland. Pers. obs., Béarez (1996), Jiménez-Pra- and García-Domínguez (2013), Ebert (2016). EN do and Béarez (2004), Martínez-Ortiz and García- Domínguez (2013). NE Family Alopiidae 16. Alopias pelagicus Nakamura, 1935 – Tiburón zorro Order pelágico – Pelagic thresher Family Odontaspididae Mainland/Galápagos. Pers. obs., Béarez (1996), 8. ferox (Risso, 1810) – Solrayo – Small- Jiménez-Prado and Béarez (2004), McCosker and tooth tiger Rosenblatt (2010), Martínez-Ortiz and García- Mainland/Galápagos. Acuña-Marrero et al. (2013), Domínguez (2013), Hearn et al. (2014), Tirado- Martínez-Ortiz and García-Domínguez (2013), Hearn Sánchez et al. (2016). EN et al. (2014), Ebert (2016), Estupiñán-Montaño et al. 17. Alopias superciliosus Lowe, 1841 – Tiburón zorro (2016), Tirado-Sánchez et al. (2016). VU ojón – Bigeye thresher 9. Odontaspis noronhai (Maul, 1955) – Solrayo – Big- Mainland/Galápagos. Pers. obs., Béarez (1996), eye sand tiger Jiménez-Prado and Béarez (2004), McCosker and Mainland/Galápagos. Martínez-Ortiz and García- Rosenblatt (2010), Martínez-Ortiz and García- Domínguez (2013). LC Domínguez (2013), Hearn et al. (2014), Tirado- Sánchez et al. (2016), Ebert (2016). VU Family Pseudocarchariidae 18. Alopias vulpinus (Bonnaterre, 1788) – Tiburón zorro 10. Pseudocarcharias kamoharai (Matsubara, 1936) – común – Common thresher Tiburón cocodrilo – Mainland/Galápagos. Pers. obs., Béarez (1996), Mainland. Pers. obs., Jiménez-Prado and Béa- Jiménez-Prado and Béarez (2004), Martínez-Ortiz rez (2004), Martínez-Ortiz and García-Domínguez and García-Domínguez (2013), Robertson and Allen (2013), Ebert (2016). LC (2015). VU

Family Order 11. Carcharodon (Linnaeus, 1758) – Jaquetón Family blanco – 19. brunneus (Gilbert, 1892) – Pejegato Mainland/Galápagos. Compagno et al. (1995), Béa- marrón – Brown rez (1996), Martínez-Ortiz and García-Domínguez Mainland. Béarez (1996), Martínez-Ortiz and García- (2013), Hearn et al. (2014), Tirado-Sánchez et al. Domínguez (2013), Ebert (2016). DD (2016). VU 20. Apristurus kampae Taylor, 1972 – Pejegato trompudo 12. oxyrinchus Rafinesque, 1810 – Marrajo dientu- – so – Shortfin mako Mainland/Galápagos. McCosker and Rosenblatt Mainland/Galápagos. Pers. obs., Béarez (1996), (2010), Martínez-Ortiz and García-Domínguez Jiménez-Prado and Béarez (2004), McCosker and (2013), Hearn et al. (2014), Ebert (2016), Tirado- Rosenblatt (2010), Martínez-Ortiz and García- Sánchez et al. (2016). DD Domínguez (2013), Hearn et al. (2014), Tirado- 21. Apristurus nasutus de Buen, 1959 – Pejegato hocicón Sánchez et al. (2016). EN – Largenose catshark 13. Isurus paucus Guitart, 1966 – Marrajo carite – Long- Mainland. Béarez (1996), Martínez-Ortiz and García- fin mako Domínguez (2013), Ebert (2016). DD

Cybium 2020, 44(3) 241 Checklist of marine cartilaginous fishes from continental and insular Ecuador Ca l l e -Mo r á n & Bé a r e z

22. Apristurus stenseni (Springer, 1979) – Pejegato pana- 32. maculata Kner & Steindachner, 1867 – Tollo meño – manchado – Spotted Galápagos. Hearn et al. (2014). DD Galápagos. McCosker and Rosenblatt (2010), Hearn et al. (2014), Tirado-Sánchez et al. (2016). VU Family Scyliorhinidae 23. giddingsi McCosker, Long & Baldwin, Family Carcharhinidae 2012 – Tiburón gato de Galápagos – Galápagos cat- 33. albimarginatus (Rüppell, 1837) – Tibu- shark rón de puntas blancas – Galápagos endemic. McCosker et al. (2012), Hearn et Mainland/Galápagos. Béarez (1996), McCosker and al. (2014), Ebert (2016). NE Rosenblatt (2010), Hearn et al. (2014), Robertson and Allen (2015), Tirado-Sánchez et al. (2016). VU Family Triakidae 34. Carcharhinus altimus (Springer, 1950) – Tiburón 24. Galeorhinus (Linnaeus, 1758) – Cazón – Tope baboso – shark Mainland/Galápagos. Béarez (1996), Jiménez-Prado Mainland. Béarez (1996), Martínez-Ortiz and García- and Béarez (2004), McCosker and Rosenblatt (2010), Domínguez (2013). VU Martínez-Ortiz and García-Domínguez (2013), Hearn 25. Mustelus albipinnis Castro-Aguirre, Antuna-Mendi- et al. (2014), Tirado-Sánchez et al. (2016). DD ola, González-Acosta & De la Cruz-Agüero, 20051 35. Carcharhinus brachyurus (Günther, 1870) – Tiburón – Tiburón de aleta lisa con margen blanco – White- cobrizo – margin fin smooth-hound Mainland. Béarez (1996), Martínez-Ortiz and García- Galápagos. Acuña-Marrero et al. (2013), Hearn et al. Domínguez (2013). NT (2014), Tirado-Sánchez et al. (2016). DD 36. Carcharhinus cerdale Gilbert, 1898 – Tiburón poroso; 26. Mustelus dorsalis Gill, 1864 – Tiburón de punta lisa – Pacific smalltail shark Sharptooth smooth-hound Mainland. Pers. obs., Jiménez-Prado and Béarez Mainland. Béarez (1996), Jiménez-Prado and Béarez (2004), Castro (2011), Robertson and Allen (2015). (2004), Robertson and Allen (2015). DD NE 27. Mustelus henlei (Gill, 1863) – Musola parda – Brown 37. Carcharhinus falciformis (Müller & Henle, 1839) – smooth-hound (Fig. 2A) Tiburón jaquetón – (Fig. 2B) Mainland. Pers. obs., Béarez (1996), Jiménez-Pra- Mainland/Galápagos. Pers. obs., Béarez (1996), do and Béarez (2004), Martínez-Ortiz and García- Jiménez-Prado and Béarez (2004), McCosker and Domínguez (2013). LC Rosenblatt (2010), Martínez-Ortiz and García- 28. Mustelus lunulatus Jordan & Gilbert, 1882 – Musola Domínguez (2013), Hearn et al. (2014), Tirado- segadora – Sicklefin smooth-hound Sánchez et al. (2016). VU Mainland. Pers. obs., Béarez (1996), Jiménez-Pra- 38. Carcharhinus galapagensis (Snodgrass & Heller, do and Béarez (2004), Martínez-Ortiz and García- 1905) – Tiburón de Galápagos – Galápagos shark Domínguez (2013), Briones-Mendoza et al. (2018). Mainland/Galápagos. Béarez (1996), Jiménez-Prado LC and Béarez (2004), McCosker and Rosenblatt (2010), 29. Mustelus mento Cope, 1877 – Perro liso moteado – Martínez-Ortiz and García-Domínguez (2013), Hearn Speckled smooth-hound et al. (2014), Tirado-Sánchez et al. (2016). LC Mainland/Galápagos. Béarez (1996), Jiménez-Prado 39. Carcharhinus leucas (Müller & Henle, 1839) – Tibu- and Béarez (2004), McCosker and Rosenblatt (2010), rón sarda – Hearn et al. (2014), Tirado-Sánchez et al. (2016). NT Mainland/Galápagos. Pers. obs., Béarez (1996), 30. Mustelus whitneyi Chirichigno F., 1973 – Musola Jiménez-Prado and Béarez (2004), Martínez-Ortiz prieta – Humpback smooth-hound and García-Domínguez (2013), Tirado-Sánchez et al. Mainland. Béarez (1996), Martínez-Ortiz and García- (2016). NT Domínguez (2013). VU 40. Carcharhinus limbatus (Müller & Henle, 1839) – 31. Triakis acutipinna Kato, 1968 – Tollo del Ecuador – Tiburón macuira – Sharpfin houndshark Mainland/Galápagos. Pers. obs., Béarez (1996), Mainland endemic. Béarez (1996), Martínez-Ortiz Jiménez-Prado and Béarez (2004), McCosker and and García-Domínguez (2013). EN Rosenblatt (2010), Martínez-Ortiz and García- Domínguez (2013), Hearn et al. (2014), Tirado- 1 Represents a controversial record that needs confirmation. Sánchez et al. (2016). NT

242 Cybium 2020, 44(3) Ca l l e -Mo r á n & Bé a r e z Checklist of marine cartilaginous fishes from continental and insular Ecuador

41. Carcharhinus longimanus (Poey, 1861) – Tiburón Domínguez (2013), Hearn et al. (2014), Tirado- oceánico – Sánchez et al. (2016). NT Mainland/Galápagos. Pers. obs., Béarez (1996), Jiménez-Prado and Béarez (2004), McCosker and Family Sphyrnidae Rosenblatt (2010), Martínez-Ortiz and García- 50. corona Springer, 1940 – Cornuda coronada – Domínguez (2013), Hearn et al. (2014), Tirado- Scalloped Sánchez et al. (2016). CR Mainland. Béarez (1996), Martínez-Ortiz and García- 42. Carcharhinus obscurus (Lesueur, 1818) – Tiburón Domínguez (2013). NT arenero – 51. Sphyrna lewini (Griffith & Smith, 1834) – Cornuda Mainland. Béarez (1996), Martínez-Ortiz and García- común – Domínguez (2013). EN Mainland/Galápagos. Pers. obs., Béarez (1996), 43. Carcharhinus plumbeus (Nardo, 1827) – Tiburón Jiménez-Prado and Béarez (2004), McCosker and trozo – Rosenblatt (2010), Martínez-Ortiz and García- Mainland/Galápagos. Béarez (1996), McCosker Domínguez (2013), Hearn et al. (2014), Tirado- and Rosenblatt (2010), Martínez-Ortiz and García- Sánchez et al. (2016). CR Domínguez (2013), Hearn et al. (2014), Tirado- 52. Sphyrna media Springer, 1940 – Cornuda cuchara – Sánchez et al. (2016). VU shark 44. Galeocerdo cuvier (Péron & Lesueur, 1822) – Tinto- Mainland. Béarez (1996), Martínez-Ortiz and García- rera tigre – Domínguez (2013). DD Mainland/Galápagos. Pers. obs., Béarez (1996), 53. Sphyrna mokarran (Rüppell, 1837) – Cornuda gigante Jiménez-Prado and Béarez (2004), McCosker and – Rosenblatt (2010), Martínez-Ortiz and García- Mainland/Galápagos. Béarez (1996), McCosker Domínguez (2013), Hearn et al. (2014), Tirado- and Rosenblatt (2010), Martínez-Ortiz and García- Sánchez et al. (2016). NT Domínguez (2013), Hearn et al. (2014), Tirado- 45. Nasolamia velox (Gilbert, 1898) – Cazón trompa blan- Sánchez et al. (2016). CR ca – (Fig. 2C) 54. Sphyrna vespertina Springer, 1940 – Cornuda de coro- Mainland/Galápagos. Pers. obs., Béarez (1996), na – Bonnethead shark Jiménez-Prado and Béarez (2004), McCosker and Mainland/Galápagos. Pers. obs., Béarez (1996), Rosenblatt (2010), Martínez-Ortiz and García- McCosker and Rosenblatt (2010), Martínez-Ortiz Domínguez (2013), Hearn et al. (2014), Tirado- and García-Domínguez (2013), Hearn et al. (2014), Sánchez et al. (2016). DD Del Moral-Flores et al. (2016), Tirado-Sánchez et al. 46. fronto (Jordan & Gilbert, 1882) – Tiburón (2016). LC galano – 55. Sphyrna zygaena (Linnaeus, 1758) – Cornuda prieta – Mainland. Béarez (1996), Martínez-Ortiz and García- Domínguez (2013), Del Moral-Flores et al. (2016). Mainland/Galápagos. Pers. obs., Béarez (1996), NT Jiménez-Prado and Béarez (2004), McCosker and 47. Prionace glauca (Linnaeus, 1758) – Tiburón azul – Rosenblatt (2010), Martínez-Ortiz and García- Domínguez (2013), Hearn et al. (2014), Tirado- Mainland/Galápagos. Pers. obs., Béarez (1996), Sánchez et al. (2016). VU Jiménez-Prado and Béarez (2004), McCosker and Rosenblatt (2010), Martínez-Ortiz and García- Order Domínguez (2013), Hearn et al. (2014), Tirado- Family Sánchez et al. (2016). NT 56. brasiliensis (Quoy & Gaimard, 1824) – Tollo 48. longurio (Jordan & Gilbert, 1882) – cigarro – Cookie-cutter shark Cazón picudo del Pacífico – Mainland/Galápagos. McCosker and Rosenblatt Mainland. Pers. obs., Béarez (1996), Martínez-Ortiz (2010), Martínez-Ortiz and García-Domínguez and García-Domínguez (2013). DD (2013), Hearn et al. (2014), Ebert (2016), Tirado- 49. Triaenodon obesus (Rüppell, 1837) – Tiburón corale- Sánchez et al. (2016). LC ro ñato – Whitetip shark Mainland/Galápagos. Pers. obs., Béarez (1996), Family Jiménez-Prado and Béarez (2004), McCosker and 57. Aculeola nigra de Buen, 1959 – Tollo negro de cachos Rosenblatt (2010), Martínez-Ortiz and García- – Hooktooth dogfish

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Mainland. Jiménez-Prado and Béarez (2004), Martín- COHORT ez-Ortiz and García-Domínguez (2013). DD Order Torpediniformes 58. nigrum Garman, 1899 – Tollo negro Family peine – Combtooth dogfish 65. ommata (Jordan & Gilbert, 1890) – Raya Mainland/Galápagos. Béarez (1996), Jiménez-Prado eléctrica ocelada – Ocellated and Béarez (2004), McCosker and Rosenblatt (2010), Mainland. Béarez (1996), Martínez-Ortiz and García- Martínez-Ortiz and García-Domínguez (2013), Hearn Domínguez (2013). VU tschudii et al. (2014), Ebert (2016), Tirado-Sánchez et al. 66. Heckel, 1846 – Raya temblera – Apron ray (2016). DD Mainland. Béarez (1996), Martínez-Ortiz and García- 59. litvinovi Parin & Kotlyar, 1990 – Linterna Domínguez (2013). NT ojuelo – Smalleye lanternshark 67. entemedor Jordan & Starks, 1895 – Raya Mainland. Martínez-Ortiz and García-Domínguez eléctrica gigante – Giant electric ray (2013). DD Mainland/Galápagos. Pers. obs., Béarez (1996), Jiménez-Prado and Béarez (2004), Martínez-Ortiz Family and García-Domínguez (2013), Tirado-Sánchez et al. 60. owstonii Garman, 1906 –Sapata lija – (2016). DD 68. Narcine leoparda Carvalho, 2001 – Raya eléctrica Mainland. Martínez-Ortiz and García-Domínguez leopardo – Leopard numbfish (2013). VU Mainland. Pers. obs., Estupiñán-Montaño and Galván- 61. pacificus Bigelow & Schroeder, 19442 – Magaña (2015), Romero-Caicedo et al. (2015). NT Tollo negro dormilón – Mainland. Martínez-Ortiz and García-Domínguez Family Torpedinidae (2013). DD 69. Tetronarce tremens (de Buen, 1959) – - no – Chilean torpedo Family Mainland/Galápagos. Pers. obs., Béarez (1996), 62. squamosus (Bonnaterre, 1788) – Quel- Jiménez-Prado and Béarez (2004), McCosker and vacho negro – Leafscale Rosenblatt (2010), Martínez-Ortiz and García- Mainland/Galápagos. Acuña-Marrero et al. (2013), Domínguez (2013), Hearn et al. (2014), Tirado- Sánchez et al. (2016). DD Martínez-Ortiz and García-Domínguez (2013), Hearn et al. (2014), Ebert (2016), Tirado-Sánchez et al. Order (2016). VU Family Trygonorrhinidae 70. Zapteryx xyster Jordan & Evermann, 1896 – Guitarra Family Echinorhinidae bruja – Southern banded guitarfish (Fig. 2E) 63. cookei Pietschmann, 1928 – Tiburón Mainland. Pers. obs., Jiménez-Prado and Béa- negro espinoso – rez (2004), Martínez-Ortiz and García-Domínguez Mainland/Galápagos. Pers. obs., Béarez (1996), (2013). DD McCosker and Rosenblatt (2010), Martínez-Ortiz and García-Domínguez (2013), Hearn et al. (2014), Ebert Family Rhinobatidae (2016), Tirado-Sánchez et al. (2016). DD 71. Pseudobatos glaucostigmus (Jordan & Gilbert, 1883) – Guitarra punteada – Speckled guitarfish Order Squatiniformes Mainland. Béarez (1996), Martínez-Ortiz and García- Family Squatinidae Domínguez (2013). DD 64. Squatina armata (Philippi, 1887) – Pez ángel del Pací- 72. Pseudobatos leucorhynchus (Günther, 1867) – Guita- fico sur – Chilean (Fig. 2D) rra trompa blanca – Whitesnout guitarfish (Fig. 2F) Mainland. Pers. obs., Béarez (1996), Jiménez-Prado Mainland. Pers. obs., Béarez (1996), Jiménez-Pra- and Béarez (2004), Acero et al. (2016). DD do and Béarez (2004), Martínez-Ortiz and García- Domínguez (2013). NT 2 This record could rather represent Somniosus antarcticus 73. Pseudobatos planiceps (Garman, 1880) – Guitarra del Whitley, 1939. Pacífico – Pacific guitarfish

244 Cybium 2020, 44(3) Ca l l e -Mo r á n & Bé a r e z Checklist of marine cartilaginous fishes from continental and insular Ecuador

Figure 2. – Pictures of some comon species from the Ecuadorian coastal small-scale (© P. Béarez).A : Mustelus henlei (596 mm TL, Puerto López, 17 Nov. 2004). B: Carcharhinus falciformis (1095 mm TL, Puerto López, 13 May 2006). C: Nasolamia velox (1060 mm TL, Puerto López, 15 Dec. 2005). D: Squatina armata (940 mm TL, Salango, 22 Apr. 1994). E: Zapteryx xyster (410 mm TL, Puerto López, 19 Nov. 2011). F: Pseudobatos leucorhynchus (664 mm TL, Salango, 8 Nov. 2008). G: Pseudobatos prahli (765 mm TL, Puerto López, 8 Nov. 2008). H: Rostroraja equatorialis (560 mm TL, Puerto López, 19 Nov. 2011). I: halleri (437 mm TL, Salango, 13 Jul. 1994). J: Myliobatis peruvianus (1920 mm TL, Salango, 11 Nov. 2008).

Cybium 2020, 44(3) 245 Checklist of marine cartilaginous fishes from continental and insular Ecuador Ca l l e -Mo r á n & Bé a r e z

Mainland/Galápagos. Pers. obs., Béarez (1996), Mar- 83. peruana McEachran & Miyake, 1984 – tínez-Ortiz and García-Domínguez (2013), Tirado- Raya peruana – Peruvian Sánchez et al. (2016). DD Mainland. Ebert (2016). DD 74. Pseudobatos prahli Acero P. & Franke, 1995 – Guita- 84. Bathyraja spinosissima (Beebe & Tee-Van, 1941) – rra de Gorgona – Gorgona guitarfish (Fig. 2G) Raya blanca – Spiny skate Mainland. Pers. obs., Jiménez-Prado and Béarez Galápagos. McCosker and Rosenblatt (2010), Hearn (2004), Payán et al. (2010), Martínez-Ortiz and et al. (2014), Ebert (2016), Tirado-Sánchez et al. García-Domínguez (2013). DD (2016). LC 85. Notoraja martinezi Concha, Ebert & Long, 2016 – Family Pristidae Raya de púas – Barbedwire-tailed skate 75. Pristis pristis (Linnaeus, 1758) – Pez sierra común – Mainland. Ebert (2016). NE Common 86. Sympterygia brevicaudata (Cope, 1877) – Raya espi- Mainland. Pers. obs., McEachran and Notarbartolo nosa – Shorttail fanskate di Sciara (1995), Jiménez-Prado and Béarez (2004), Mainland. Béarez (1996), Martínez-Ortiz and García- Faria et al. (2013). CR Domínguez (2013). DD

Order Family Gurgesiellidae Family Rajidae 87. Gurgesiella furvescens de Buen, 1959 – Raya falsa 76. ecuadoriensis (Beebe & Tee-Van, 1941) – sureña – Dusky finless skate Raya de Ecuador – Ecuador skate Mainland/Galápagos. Grove and Lavenberg (1997), Mainland endemic. Jiménez-Prado and Béarez (2004), McCosker and Rosenblatt (2010), Hearn et al. (2014), Last et al. (2016). DD Ebert (2016), Tirado-Sánchez et al. (2016). LC 77. Rajella eisenhardti Long & McCosker, 1999 – Raya de Galápagos – Galápagos grey skate Order Galápagos endemic. McCosker and Rosenblatt (2010), Family Dasyatidae dipterurus Hearn et al. (2014), Ebert (2016), Tirado-Sánchez et 88. (Jordan & Gilbert, 1880) – Raya látigo diamante – Diamond al. (2016). DD Mainland/Galápagos. Béarez (1996), McCosker 78. Rajella nigerrima (de Buen, 1960) – Raya negra – and Rosenblatt (2010), Martínez-Ortiz and García- Blackish skate Domínguez (2013), Hearn et al. (2014), Tirado- Mainland. Martínez-Ortiz and García-Domínguez Sánchez et al. (2016). DD (2013), Ebert (2016). LC 89. Hypanus longus (Garman, 1880) – Raya látigo coluda 79. Rostroraja equatorialis (Jordan & Bollman, 1890) – – Raya ecuatorial – Equatorial ray (Fig. 2H) Mainland. Pers. obs., Béarez (1996), Jiménez-Prado Mainland. Pers. obs., Béarez (1996), Jiménez-Pra- and Béarez (2004), McCosker and Rosenblatt (2010), do and Béarez (2004), Martínez-Ortiz and García- Martínez-Ortiz and García-Domínguez (2013), Hearn Domínguez (2013). DD et al. (2014), Tirado-Sánchez et al. (2016). DD Rostroraja velezi 80. (Chirichigno F., 1973) – Raya de 90. Telatrygon acutirostra (Nishida & Nakaya, 1988)3 – Vélez – Velez ray Raya águda – Sharpnose stingray Mainland/Galápagos. Pers. obs., Béarez (1996), Mainland. Aguilar (2008). NT Jiménez-Prado and Béarez (2004), McCosker and 91. Pteroplatytrygon violacea (Bonaparte, 1832) – Raya Rosenblatt (2010), Martínez-Ortiz and García- látigo violeta – Pelagic stingray Domínguez (2013), Hearn et al. (2014), Tirado- Mainland/Galápagos. Jiménez-Prado and Béarez Sánchez et al. (2016). DD (2004), McCosker and Rosenblatt (2010), Martínez- Ortiz and García-Domínguez (2013), Hearn et al. Family (2014), Tirado-Sánchez et al. (2016). LC 81. Bathyraja abyssicola (Gilbert, 1896) – Raya abisal – 92. meyeni (Müller & Henle, 1841) – Raya Deepsea skate redonda – Round ribbontail ray Galápagos. Cerutti-Pereyra et al. (2018). DD Galápagos. McCosker and Rosenblatt (2010), Hearn 82. Bathyraja aguja (Kendall & Radcliffe, 1912) – Raya et al. (2014), Tirado-Sánchez et al. (2016). VU aguja – Aguja skate Mainland. Martínez-Ortiz and García-Domínguez 3 This record by Aguilar (2008) likely represents an undescribed (2013). DD species.

246 Cybium 2020, 44(3) Ca l l e -Mo r á n & Bé a r e z Checklist of marine cartilaginous fishes from continental and insular Ecuador

Family Domínguez (2013), Hearn et al. (2014), Tirado- 93. Styracura pacifica (Beebe & Tee-Van, 1941) – Chupa- Sánchez et al. (2016). NE re del Pacífico – Pacific chupare Galápagos. McCosker and Rosenblatt (2010), Hearn Family Myliobatidae et al. (2014), Tirado-Sánchez et al. (2016). NE 103. Aetomylaeus asperrimus (Gilbert, 1898) – Raya águi- la áspera – Rough Family Mainland/Galápagos. McCosker and Rosenblatt 94. Urobatis halleri (Cooper, 1863) – Raya redonda de (2010), Martínez-Ortiz and García-Domínguez Haller – Haller’s round ray (Fig. 2I) (2013), Hearn et al. (2014), Tirado-Sánchez et al. Mainland. Pers. obs., Béarez (1996), Jiménez-Pra- (2016). DD do and Béarez (2004), Martínez-Ortiz and García- 104. Myliobatis californica Gill, 1865 – Raya murciélago – Domínguez (2013). LC Bat eagle ray 95. Urobatis tumbesensis (Chirichigno F. & McEachran, Galápagos. Grove and Lavenberg (1997), Tirado- 1979) – Raya psicodélica – Tumbes Sánchez et al. (2016). LC Mainland. Pers. obs., Jiménez-Prado and Béa- 105. Myliobatis longirostris Applegate & Fitch, 1964 – rez (2004), Martínez-Ortiz and García-Domínguez Raya águila picuda – Snouted eagle ray (2013). DD Mainland. Pers. obs., Jiménez-Prado and Béa- 96. aspidura (Jordan & Gilbert, 1882) – Raya rez (2004), Martínez-Ortiz and García-Domínguez redonda de rabo espinoso – Spiny-tail round ray (2013). NT Mainland. Béarez (1996), Jiménez-Prado and Béarez 106. Myliobatis peruvianus Garman, 1913 – Raya águila (2004). DD peruana – Peruvian eagle ray (Fig. 2J) 97. Urotrygon chilensis (Günther, 1879) – Raya redonda Mainland/Galápagos. Pers. obs., Béarez (1996), chilena – Chilean round ray McCosker and Rosenblatt (2010), Martínez-Ortiz and Mainland. Pers. obs., Béarez (1996), Jiménez-Pra- García-Domínguez (2013), Hearn et al. (2014), Tira- do and Béarez (2004), Martínez-Ortiz and García- do-Sánchez et al. (2016). DD Domínguez (2013). DD 98. Urotrygon munda Gill, 1863. – Raya redonda munda Family Rhinopteridae – Munda round ray 107. Rhinoptera steindachneri Evermann & Jerkins, 1891 Mainland. Jiménez-Prado and Béarez (2004), Robert- – Gavilán negro – Pacific cownose ray son and Allen (2015). DD Mainland/Galápagos. Pers. obs., Béarez (1996), 99. Urotrygon rogersi (Jordan & Starks, 1895) – Raya Jiménez-Prado and Béarez (2004), McCosker and redonda de Rogers – Roger’s round ray Rosenblatt (2010), Martínez-Ortiz and García- Mainland. Pers. obs., Jiménez-Prado and Béa- Domínguez (2013), Tirado-Sánchez et al. (2016). NT rez (2004), Martínez-Ortiz and García-Domínguez (2013). DD Family 108. birostris (Walbaum, 1792) – Manta gigante – Family Gymnuridae Giant 100. Gymnura crebripunctata (Peters, 1869) – Raya mari- Mainland/Galápagos. Béarez (1996), Jiménez-Prado posa – Longsnout and Béarez (2004), McCosker and Rosenblatt (2010), Mainland. Pers. obs., Béarez (1996), Jiménez-Prado Martínez-Ortiz and García-Domínguez (2013), Hearn and Béarez (2004). DD et al. (2014), Tirado-Sánchez et al. (2016). VU 101. Gymnura marmorata (Cooper, 1864) – Raya maripo- 109. Mobula mobular (Bonnaterre, 1788) – Manta mobula sa californiana – butterfly ray – Devil fish Mainland. Pers. obs., Béarez (1996), Jiménez-Pra- Mainland/Galápagos. Tirado-Sánchez et al. (2016), do and Béarez (2004), Martínez-Ortiz and García- Lezama-Ochoa et al. (2019). EN Domínguez (2013). LC 110. Mobula munkiana Notarbartolo-di-Sciara, 1987 – Manta de Munk – Munk’s devil ray Family Aetobatidae Mainland/Galápagos. Pers. obs., Jiménez-Prado and 102. laticeps (Gill, 1865) – Chucho pintado – Béarez (2004), McCosker and Rosenblatt (2010), Pacific Martínez-Ortiz and García-Domínguez (2013), Hearn Mainland/Galápagos. Pers. obs., Béarez (1996), et al. (2014), Tirado-Sánchez et al. (2016). VU Jiménez-Prado and Béarez (2004), McCosker and 111. Mobula tarapacana (Philippi, 1892) – Manta cornuda Rosenblatt (2010), Martínez-Ortiz and García- – Chilean devil ray

Cybium 2020, 44(3) 247 Checklist of marine cartilaginous fishes from continental and insular Ecuador Ca l l e -Mo r á n & Bé a r e z

Mainland/Galápagos. McCosker and Rosenblatt (2010), Hearn et al. (2014), Tirado-Sánchez et al. (2016). EN 112. Mobula thurstoni (Lloyd, 1908) – Diablo chupasangre – Smoothtail mobula Mainland/Galápagos. Pers. obs., Béarez (1996), Jiménez-Prado and Béarez (2004), Martínez-Ortiz and García-Domínguez (2013), Lezama-Ochoa et al. (2019). NT

Subclass Order Chimaeriformes Family Callorhinchidae 113. Callorhinchus callorynchus (Linnaeus, 1758) – Peje- gallo – Plownose Figure 3. – Number of species per order of sharks, rays and chimae- Mainland. Pers. obs., Swing and Béarez (2006), Mar- ras in Ecuador (n = 119). tínez-Ortiz and García-Domínguez (2013). LC

Family The order with the highest number of species for sharks 114. Hydrolagus alphus Quaranta, Didier, Long & Ebert, was Carcharhiniformes with 37 species, and the lowest was 2006 – Quimera de Galápagos – Whitespot ghost- Squatiniformes with only 1 species. For rays the highest shark number of species was for Myliobatiformes with 25 species, Galápagos endemic. McCosker and Rosenblatt (2010), and the lowest was Torpediniformes with 5 species. The Ebert (2016), Tirado-Sánchez et al. (2016). DD order of Chimaeriformes included 7 species (Fig. 3). 115. Hydrolagus macrophthalmus de Buen, 1959 – Quime- Nearly half (58) of Ecuadorian chondrichthyans are only ra de ojo grande – Bigeye chimaera found along the mainland coast, while 12 are only found in Mainland. Martínez-Ortiz and García-Domínguez Galápagos, and 49 are shared between the two regions. It is (2013), Ebert (2016). DD interesting to note that several families of rays (e.g. urotry- 116. Hydrolagus mccoskeri Barnett, Didier, Long & Ebert, gonids, gymnurids) with limited swimming ability are absent 2006 – Quimera fantasma de Galápagos – Galápagos from the insular region. The endemism of Chondrichthyans ghostshark is rather low (5%) in Ecuadorian waters, accounting for just 2 Galápagos endemic. McCosker and Rosenblatt (2010), sharks, 2 rays, and 2 chimaeras. Among them, two are main- Ebert (2016), Tirado-Sánchez et al. (2016). DD land endemics (Triakis acutipinna, Dipturus ecuadoriensis) 117. Hydrolagus melanophasma James, Ebert, Long & and four are Galápagos endemics (Bythaelurus giddingsi, Didier, 2009 – Quimera negra del Pacífico oriental – Rajella eisenhardti, Hydrolagus alphus, H. mccoskeri). Eastern Pacific black ghostshark Among the 119 species of Ecuadorian chondrichthyans, Mainland. Aguirre-Villaseñor et al. (2013), Martínez- 4 are Critically Endangered, 9 are Endangered, 17 are Vul- Ortiz and García-Domínguez (2013), Ebert (2016). nerable, 17 are Near Threatened, 18 are Least Concern, 48 LC are Data Deficient, and 6 are Not Evaluated. It is important to note the high proportion of species without data to assess Family their status. This shows the low level of knowledge about 118. raleighana Goode & Bean, 1895 – Quimera chondrichthyans in general, and in this region of the world de Raleigh – Narrownose chimaera in particular. Mainland. Martínez-Ortiz and García-Domínguez (2013), Robertson and Allen (2015). LC 119. africana Compagno, Stehmann & DISCUSSION Ebert, 19904 – Pez fantasma africano – Paddlenose spookfish The number of chondrichthyans reported for Ecuador in Mainland. Ebert (2016). DD this study (119) is higher than that previously recorded for 4 The specimen mentioned as Rhinochimaera sp. by Martínez- continental Ecuador (Martínez-Ortiz and García-Domíngu- Ortiz and García-Domínguez (2013: 24) probably represents this ez, 2013) and the Galápagos Islands (Hearn et al., 2014), species. which totalled 112. It is also higher than what was recorded

248 Cybium 2020, 44(3) Ca l l e -Mo r á n & Bé a r e z Checklist of marine cartilaginous fishes from continental and insular Ecuador in Peruvian waters according to Cornejo et al. (2015) who REFERENCES cited 115 species (66 sharks, 43 rays, and 6 chimaeras) and for Chilean waters, for which 96 species (53 sharks, 39 rays, ACERO A., TAVERA J.J., ANGUILA R. & HERNÁNDEZ L., 2016. – A new southern Caribbean species of Angel shark and 4 chimaeras) were recorded by Lamilla and Bustamante (Chondrichthyes, Squaliformes, Squatinidae), including phyl- (2005). This is also the case for Costa Rican Pacific waters, ogeny and tempo of diversification of American species. Copeia, 104(2): 577-585. DOI: 10.1643/CI-15-292 where only 87 species (42 sharks, 39 rays, and 6 chimaeras) ACUÑA-MARRERO D., ZIMMERHACKEL J.S., MAYORGA J. were recorded by Espinoza et al. (2018), and Colombian & HEARN A., 2013. – First record of three shark species, Odontaspis ferox, Mustelus albipinnis and Centrophorus squa- Pacific waters which bears the lowest richness with only 67 mosus, from the Galápagos Islands. Mar. Biodivers. Rec., 6: species of elasmobranchs (36 sharks and 31 rays), recorded e87: 1-6. DOI: 10.1017/S1755267213000596 by Mejía-Falla and Navia (2019). However, the number of AGUILAR A.F., 2008. – acutirostra (Nishida & Nakaya, 1988) nuevo Registro de Raya del Género Dasyatis en Aguas chondrichthyans reported here is lower than that of Mexican ecuatorianas. 4 p. Guayaquil: Instituto Nacional de Pesca. Pacific waters for which 123 species (63 sharks, 55 rays, and Aguirre-Villaseñor H., Salas-Singh C., Madrid- 5 chimaeras) were recorded by Ehemann et al. (2018). Vera J., Martínez-Ortiz J., Didier D.A. & Ebert D.A., 2013. - New eastern Pacific Ocean records ofHydrolagus Of the 1,464 chondrichthyan species compiled world- melanophasma, with annotations of a juvenile female. J. Fish wide by Fricke et al. (2020), 8.1% exist in Ecuador: 10.7% Biol., 82(2): 714-724. DOI: 10.1111/jfb.12012 BÉAREZ P., 1996. – Lista de los peces marinos del Ecuador conti- sharks, 5.9% rays, and 12.5% chimaeras. Rays have the nental. Rev. Biol. Trop., 44(2): 731-741. poorest representation. Of this total number of known chon- Briggs J.C. & Bowen B.W., 2012. – A realignment of marine drichthyans, Chile has 6.6%, 7.9%, 4.6%, biogeographic provinces with particular reference to fish distri- butions. J. Biogeogr., 39: 12-30. DOI: 10.1111/j.1365- 5.9%, and 8.4%. 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Cerutti-Pereyra F., Yánez A.B., Ebert D.A., Arnés- The number of elasmobranchs reported in this study Urgellés C. & Salinas-de-León P., 2018. – New may be higher due to the fact that some unidentified spe- record and range extension of the deepsea skate, Bathyraja abyssicola (Chondrichthyes: Arhynchobatidae), in the Galapa- cies, which might prove to be undescribed species, were gos Islands. J. Ocean Sci. Found., 30: 85-89. DOI: 10.5281/ not taken into account. This is the case for Galeus sp. and zenodo.1400829 Hypanus sp. recorded by McCosker and Rosenblatt (2010), Compagno L.J.V., Krupp F. & Schneider W., 1995. – Tibu- rones. In: Guía FAO para la Identificación de Especies para los and successively by Hearn et al. (2014) and Tirado-Sánchez Fines de la Pesca. Pacífico centro-oriental (Fischer W., Krupp et al. (2016). Also, a chimaera mentioned by McCosker and F., Schneider W., Sommer C., Carpenter K.E. & Niem V.H., eds), Vertebrados, parte 1, vol. 2., pp. 647-743. Rome: FAO. 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