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Etmopterus benchleyi Vásquez, Ebert, & Long, 2015. To 51.5 cm (20.3 in) TL. Isla Guadalupe, central Baja California (Personal communication: Scripps Institution of Oceanography Fish Collection, La Jolla, California); Nicaragua to Panama (Vásquez et al. 2015). Depth: 836–1,470 m (2,742 –4,822 ft) (min.: Vásquez et al. 2015; max.: Morera et al. 2019).

Etmopterus benchleyi V��squez, Ebert, & Long, 2015. To 51.5 cm (20.3 in) TL. Isla Guadalupe, central Baja California (Personal communication: Scripps Institution of Oceanography Fish Collection, La Jolla, California); Nicaragua to Panama (V��squez et al. 2015). Depth: 836���1,470 m (2,742 ���4,822 ft) (min.: V��squez et al. 2015; max.: Morera et al. 2019)., Published as part of Love, Milton S., Bizzarro, Joseph J., Cornthwaite, Maria, Frable, Benjamin W. & Maslenikov, Katherine P., 2021, Checklist of marine and estuarine fishes from the Alaska-Yukon Border, Beaufort Sea, to Cabo San Lucas, Mexico, pp. 1-285 in Zootaxa 5053 (1) on page 21, DOI: 10.11646/zootaxa.5053.1.1, http://zenodo.org/record/5578008, {"references":["Vasquez, V. E., Ebert, D. A. & Long, D. J. (2015) Etmopterus benchleyi n. sp., a new lanternshark (Squaliformes: Etmopteridae) from the central eastern Pacific Ocean. Journal of the Ocean Science Foundation, 17, 43 - 55.","Morera, R. B., Calderon, F. C. & Hernandez, J. M. V. (2019) Abundancia, biomasa y estructura de la ictiofauna demersal en el oceano Pacifico de Centroamerica, basadas en datos de prospeccion pesquera realizados a bordo del B / W Miguel Oliver. Revista Ciencias Las Marinas y Costeras, 12, 27 - 47. https: // doi. org / 10.15359 / revmar. 12 - 1.2"]}

Etmopterus spinax (Linnaeus, 1758) Etmopterus spinax: Gilat & Gelman, 1984a; Gilat & Gelman, 1984b (Heb.); Hornung et al. 1993 (Poll.); Galil & Goren, 1994; Golani, 1996; Pisanty & Golani, 1996; Golani, 1997 (Heb.); Goren & Galil, 2002; Galil, 2004; Golani, 2005; Golani, 2006, Goren & Galil, 2015; Paz et al. 2018 (gen.). SOMNIOSINAE, Published as part of Golani, Daniel, 2021, An updated Checklist of the Mediterranean fishes of Israel, with illustrations of recently recorded species and delineation of Lessepsian migrants, pp. 1-108 in Zootaxa 4956 (1) on page 8, DOI: 10.11646/zootaxa.4956.1.1, http://zenodo.org/record/4691195, {"references":["Gilat, E. & Gelman, A. (1984 a) On sharks and fishes observed using underwater photography during a deep-water cruise in the eastern Mediterranean. Fisheries Research, 2, 257 - 271. https: // doi. org / 10.1016 / 0165 - 7836 (84) 90029 - 8","Gilat, E. & Gelman, A. (1984 b) Shark populations in deep waters off the Israeli Mediterranean coast and Cyprus. Fisheries and Fishbreeding in Israel, 17, 3 - 15 [in Hebrew]","Hornung, H., Krom, M. D., Cohen, Y. & Bernhard, M. (1993) Trace metal content in deep-water sharks from the eastern Mediterranean Sea. Marine Biology, 115, 331 - 338. https: // doi. org / 10.1007 / BF 00346351","Galil, B. S. & Goren, M. (1994) The deep sea Levantine fauna. - New records and rare occurrences. Senckenbergiana maritima, 25 (1 / 3), 41 - 52.","Pisanty, S. & Golani, D. (1996) Vertical distribution of the demersal fish on the continental slope of Israel (Eastern Mediterranean). In: Armentrout, N. B. (Ed.), Condition of the World's Aquatic Habitats. Proceedings of the World Fisheries Congress, Theme 1. Oxford & IBH Publishing Company, New Delhi, pp. 386 - 395.","Golani, D. (1997) Handbook of the Fishes of Israel. Keter Publishing House Ltd. Jrusalem. 269 pp. [in Hebrew].","Goren, M. & Galil, B. S. (2002) Records of Cataetyx laticeps and Ophidion barbatum (Ophidiiformes) in the eastern Mediterranean, with comments on the deep sea ichthyofauna. Cybium, 26, 150 - 152.","Galil, B. S. (2004) The limit of the sea: the bathyal fauna of the Levantine Sea. Scientia Marina, 68 (suppl.), 63 - 72. https: // doi. org / 10.3989 / scimar. 2004.68 s 363","Goren, M. & Galil, B. S. (2015) A checklist of the deep sea fishes of the Levant Sea, Mediterranean Sea. Zootaxa, 3994 (4), 507 - 530. https: // doi. org / 10.11646 / zootaxa. 3994.4.2","Paz, G., Yudkovsky, Y., Shaish, L., Stern, N., Lubinevski, H., Mineis, H., Douek, J., Galil, B., Goren M. & Rinkevich, B. (2018) Initiating the DNA barcoding of Eastern Mediterranean deep-sea biota. Mediterranean Marine Science, 19 (3), 416 - 429. https: // doi. org / 10.12681 / mms. 14146"]}

Etmopterus bigelowi Shirai & Tachikawa, 1993 Blurred Lanternshark Etmopterus bigelowi Shirai & Tachikawa, 1993: 487, figs. 1, 2, 5. Holotype: HUMZ 100176. Type locality: off Angola, 11°37'S, 05°13'W, southeastern Atlantic. Local synonymy: Etmopterus bigelowi: Compagno, 1999: 114; Compagno et al., 2005: 95, fig., pl. 8; Ebert, 2013: 83, fig. 104; Ebert et al., 2013 a: 125, fig., pl. 10; Ebert & Mostarda, 2015: 30, fig.; Ebert, 2015: 75, fig. 81; Ebert & Mostarda, 2015: 27, fig.; Ebert & van Hees, 2015: 144; Compagno, 2016: 1186; Weigmann, 2016: 894. South Africa voucher material: FAKU 46064, FSFL-S 427, HUMZ 74378. Several uncatalogued SAM specimens from off Mossel Bay (WC). South Africa distribution: Scattered records from off the west coast (NC) to KZN. Remarks: The occurrence of this species off South Africa is patchy, but this may be due to misidentification with E. pusillus. Conservation status: LC (2020).

Etmopterus granulosus (Günther, 1880) Southern Lanternshark Spinax granulosus G̹nther, 1880: 19, fig. C, pl. 2. Holotype: BMNH 1879.5.14.460. Type locality: Southwestern coast of South America, Challenger station 305. Local synonymy: Etmopterus granulosus: Gilchrist, 1922b: 49; Barnard, 1925: 49, fig. 8, pl. 2; Smith, 1949a: 58, fig. 50; Bigelow & Schroeder, 1957: 55; Smith, 1965: 58, fig. 50; Compagno, 1984a: 77, fig.; Bass et al., 1986: 55 (Cape Point); Compagno et al., 1989: 28, pl.; Straube et al., 2011a: 138; Ebert, 2013: 84, fig. 117; Ebert et al., 2013 a: 132, fig., pl. 9; Ebert & Mostada, 2013: 31, fig.; Ebert, 2015: 78, fig. 91; Ebert & Mostada, 2015: 25, fig.; Ebert & van Hees, 2015: 144; Straube et al., 2015: 11; Weigmann, 2016: 895. Spinax granulosus: Norman, 1935: 37 (SW of Cape Town, 34°08'S, 17°33'E). Etmopterus cf. granulosus: Compagno et al., 1991: 63; Ebert et al., 1992: 605; Compagno, 1999: 114. Etmopterus baxteri: Compagno et al., 2005: 94, pl. 8; Compagno, 2016: 1185. South Africa voucher material: SAIAB 25710, SAIAB 26341, SAIAB 26342, SAIAB 26343, SAIAB 26344, SAIAB 26345, SAIAB 26346, SAIAB 26347, SAIAB 26348, SAIAB 26349, SAIAB 26350, SAIAB 26351, SAIAB 26352, SAIAB 26353, SAIAB 26354, SAIAB 26355, SAIAB 26356, SAIAB 26357, SAIAB 26358, SAIAB 26359, SAIAB 26360, SAIAB 26361, SAIAB 26362, SAIAB 26363, SAIAB 26364, SAIAB 26365, SAIAB 26366, SAIAB 26367, SAIAB 26368, SAIAB 26369, SAIAB 26370, SAIAB 26371, SAIAB 26372, SAIAB 26373, SAIAB 26374, SAIAB 26375, SAIAB 26430, SAIAB 26431, SAIAB 26432, SAIAB 26433, SAIAB 26434, SAIAB 26435, SAIAB 27170, SAIAB 27171, SAIAB 27172, SAIAB 27173, SAIAB 27174, SAIAB 27175, SAIAB 27176, SAIAB 27177, SAIAB 27178, SAIAB 27179, SAIAB 27180, SAIAB 27181, SAIAB 27182, SAIAB 27184, SAIAB 27185, SAIAB 27186, SAIAB 30297, SAIAB 81676, SAIAB 186424, SAIAB 186462. South Africa distribution: West coast from Cape Columbine (WC) to about Algoa Bay (EC). Remarks: The taxonomic history of this species off South Africa, like most members of this genus, has been rather convoluted, although Straube et al. (2015) clarified the name status of this species. Etmopterus granulosus is one of the most common lanternsharks off the South African west coast. It is wide-ranging throughout the Southern Hemisphere at higher latitudes. Conservation status: LC (2018)., Published as part of Ebert, David A., Wintner, Sabine P. & Kyne, Peter M., 2021, An annotated checklist of the chondrichthyans of South Africa, pp. 1-127 in Zootaxa 4947 (1) on page 29, DOI: 10.11646/zootaxa.4947.1.1, http://zenodo.org/record/4614567, {"references":["Gilchrist, J. D. F. (1922 b) Deep-sea fishes procured by the S. A. \" Pickle \" (Part I.). Special Report (3), Report of the Fisheries and Marine Biological Survey, Union of South Africa, 2, 41 - 79.","Barnard, K. H. (1925) A monograph of the marine fishes of South Africa. Part I (Amphioxus, Cyclostomata, Elasmobranchii, and Teleostei-Isospondyli to Heterosomata). Annals of the South African Museum, 21, 1 - 418.","Smith, J. L. B. (1949 a) The Sea Fishes of Southern Africa. South Africa Central News Agency Ltd., 550 pp.","Bigelow, H. B. & Schroeder, W. C. (1957) A study of the sharks of the suborder Squaloidea. Bulletin of the Museum of Comparative Zoology, 117 (1), 1 - 150.","Smith, J. L. B. (1965) The Sea Fishes of Southern Africa. 5 th Edition. Central News Agency Ltd., 580 pp.","Compagno, L. J. V. (1984 a) FAO Species Catalogue. Sharks of the World. An Annotated and Illustrated Catalogue of Shark Species Known to Date. FAO Fisheries Synopsis. Vol. 4. No. 125. Part 1. Hexanchiformes to Lamniformes. FAO, Rome, pp. 1 - 250.","Bass, A. J. & Compagno, L. J. V. (1986) Families Echinorhinidae, Proscyllidae, Odontaspidiidae, Mitsukurinidae. In: Smith, M. M. & Heemstra, P. C. (Eds.), Smith's Sea Fishes. Macmillan, Johannesburg, pp. 63 + 103 + 104 - 105.","Compagno, L. J. V., Ebert, D. A. & Smale, M. J. (1989) Guide to the Sharks and Rays of Southern Africa. Struik Publishers, Cape Town, 158 pp.","Ebert D. A. (2013) Deep-sea cartilaginous fishes of the Indian Ocean. Vol. 1. Sharks. FAO Species Catalogue for Fishery Purposes No. 8. Vol. 1. FAO, Rome, 256 pp.","Ebert, D. A., Fowler, S. & Compagno, L. J. V. (2013) Sharks of the World: A Fully Illustrated Guide to the Sharks of the World. Wild Nature Press, Plymouth, 528 pp.","Ebert, D. A. (2015) Deep-sea cartilaginous fishes of the Southeastern Atlantic Ocean. FAO Species Catalogue for Fishery Purposes No. 9. FAO, Rome, 251 pp.","Ebert, D. A. & van Hees, K. E. (2015) Beyond jaws: rediscovering the \" Lost Sharks \" of southern Africa. African Journal of Marine Science, 37, 141 - 156. https: // doi. org / 10.2989 / 1814232 X. 2015.1048730","Straube, N., Leslie, R. W., Clerkin, P. J., Ebert, D. A., Rochel, E., Corrigan, S. L., Chenhong, L. & Naylor, G. J. P. (2015) On the occurrence of the southern lanternshark, Etmopteus granulosus, off South Africa, with comments on the validity of E. compagnoi. Deepsea Research II: Topical Studies in Oceanography, 115, 11 - 17. https: // doi. org / 10.1016 / j. dsr 2.2014.04.004","Weigmann, S. (2016) Annotated checklist of the living sharks, batoids and chimaeras (Chondrichthyes) of the world, with a focus on biogeographical diversity. Journal of Fish Biology, 88, 837 - 1037. https: // doi. org / 10.1111 / jfb. 12874","Norman, J. R. (1935) Coast fishes, Part. I. The South Atlantic. ' Discovery' Reports, 12, 1 - 58.","Compagno, L. J. V., Ebert, D. A. & Cowley, P. D. (1991) Distribution of offshore demersal cartilaginous fishes (class Chondrichthyes) of the west coast of southern Africa, with notes on their systematics. South African Journal of Marine Science, 11, 43 - 139. https: // doi. org / 10.2989 / 025776191784287664","Ebert, D. A., Compagno, L. J. V. & Cowley, P. D. (1992) A preliminary investigation of the feeding ecology of squaloid sharks off the west coast of southern Africa. South African Journal of Marine Science, 12, 601 - 609. https: // doi. org / 10.2989 / 02577619209504727","Compagno, L. J. V. (1999) An overview of chondrichthyan systematics and biodiversity in southern Africa. Transactions of the Royal Society of South Africa, 54, 75 - 120. https: // doi. org / 10.1080 / 00359199909520406","Compagno, L., Dando, M. & Fowler, S. (2005) Field Guide to the Sharks of the World. Harper Collins Publishers Ltd, London, 368 pp.","Compagno, L. J. V. (2016) Sharks. In: Carpenter, K. E. & De Angelis, N. (Eds.), The Living Marine Resources of the Eastern Central Atlantic. Vol. 2. Bivalves, Gastropods, Hagfishes, Sharks, Batoid Fishes, and Chimaeras. FAO Species Identification Guide for Fishery Purposes. FAO, Rome, pp. 1122 - 1336."]}

Etmopterus sculptus Ebert, Compagno, & De Vries, 2011 Sculpted Lanternshark Etmopterus sculptus Ebert, Compagno, & De Vries, 2011: 279, figs. 1–2. Holotype: SAM 37569. Type locality: 33°22.9'S, 17°29.1'E, off Cape Town, South Africa. Local synonymy: Etmopterus lucifer: Gilchrist, 1922b: 49; Barnard, 1925: 50; Smith, 1949a: 59, fig. 52; Bigelow & Schroeder, 1957: 56; Smith, 1965: 59, fig. 52; Bass et al., 1976: 25, figs. 17, 18c; Compagno, 1984: 79, fig.; Compagno et al., 2005: 102, fig., pl. 8. Spinax lucifer: Norman, 1935: 37. Etmopterus brachyurus: Bass et al., 1986: 55, fig. 5.11; Compagno et al., 1989: 30, pl.; Compagno et al., 1991: 61. Etmopterus cf. brachyurus: Ebert et al., 1992: 604; Compagno, 1999: 114; Compagno, 2016: 1186, fig. Etmopterus sculptus: Ebert, 2013: 83, fig. 108; Ebert et al., 2013 a: 139, fig., pl. 10; Ebert & Mostada, 2013: 33, fig.; Ebert, 2015: 82, fig. 95; Ebert & Mostada, 2015: 29, fig.; Ebert & van Hees, 2015: 144; Weigmann, 2016: 897. South Africa voucher material: Holotype: SAM 37569. Paratypes: SAM 33011, 37570 (2 specimens), 37571 (2 specimens). Non-types: SAIAB 6191, SAIAB 6195, SAIAB 2430, SAIAB 2431, SAIAB 2432, SAIAB 2588, SAIAB 21930, SAIAB 21931, SAIAB 25308, SAIAB 25309, SAIAB 25735, SAIAB 26256, SAIAB 26258, SAIAB 26259, SAIAB 26265, SAM F41924 -3, SAM F41924 -4. South Africa distribution: Occurs along the entire South African coast from the Orange River (NC) to KZN border with Mozambique. Remarks: Etmopterus sculptus was long misidentified in the South African literature either as E. brachyurus or E. lucifer, but both of these species occur in the Western Pacific. Etmopterus sculptus is one of the most common lanternshark species found in South African waters (after E. granulosus). Conservation status: LC (2019)., Published as part of Ebert, David A., Wintner, Sabine P. & Kyne, Peter M., 2021, An annotated checklist of the chondrichthyans of South Africa, pp. 1-127 in Zootaxa 4947 (1) on pages 29-30, DOI: 10.11646/zootaxa.4947.1.1, http://zenodo.org/record/4614567, {"references":["Gilchrist, J. D. F. (1922 b) Deep-sea fishes procured by the S. A. \" Pickle \" (Part I.). Special Report (3), Report of the Fisheries and Marine Biological Survey, Union of South Africa, 2, 41 - 79.","Barnard, K. H. (1925) A monograph of the marine fishes of South Africa. Part I (Amphioxus, Cyclostomata, Elasmobranchii, and Teleostei-Isospondyli to Heterosomata). Annals of the South African Museum, 21, 1 - 418.","Smith, J. L. B. (1949 a) The Sea Fishes of Southern Africa. South Africa Central News Agency Ltd., 550 pp.","Bigelow, H. B. & Schroeder, W. C. (1957) A study of the sharks of the suborder Squaloidea. Bulletin of the Museum of Comparative Zoology, 117 (1), 1 - 150.","Smith, J. L. B. (1965) The Sea Fishes of Southern Africa. 5 th Edition. Central News Agency Ltd., 580 pp.","Bass, A. J., D'Aubrey, J. D. & Kistnasamy, N. (1976) Sharks of the east coast of southern Africa. VI. The families Oxynotidae, Squalidae, Dalatiidae and Echinorhinidae. Investigational Report. Oceanographic Research Institute, Durban, 45, 1 - 103.","Compagno, L. J. V. (1984 a) FAO Species Catalogue. Sharks of the World. An Annotated and Illustrated Catalogue of Shark Species Known to Date. FAO Fisheries Synopsis. Vol. 4. No. 125. Part 1. Hexanchiformes to Lamniformes. FAO, Rome, pp. 1 - 250.","Compagno, L., Dando, M. & Fowler, S. (2005) Field Guide to the Sharks of the World. Harper Collins Publishers Ltd, London, 368 pp.","Norman, J. R. (1935) Coast fishes, Part. I. The South Atlantic. ' Discovery' Reports, 12, 1 - 58.","Bass, A. J. & Compagno, L. J. V. (1986) Families Echinorhinidae, Proscyllidae, Odontaspidiidae, Mitsukurinidae. In: Smith, M. M. & Heemstra, P. C. (Eds.), Smith's Sea Fishes. Macmillan, Johannesburg, pp. 63 + 103 + 104 - 105.","Compagno, L. J. V., Ebert, D. A. & Smale, M. J. (1989) Guide to the Sharks and Rays of Southern Africa. Struik Publishers, Cape Town, 158 pp.","Compagno, L. J. V., Ebert, D. A. & Cowley, P. D. (1991) Distribution of offshore demersal cartilaginous fishes (class Chondrichthyes) of the west coast of southern Africa, with notes on their systematics. South African Journal of Marine Science, 11, 43 - 139. https: // doi. org / 10.2989 / 025776191784287664","Ebert, D. A., Compagno, L. J. V. & Cowley, P. D. (1992) A preliminary investigation of the feeding ecology of squaloid sharks off the west coast of southern Africa. South African Journal of Marine Science, 12, 601 - 609. https: // doi. org / 10.2989 / 02577619209504727","Compagno, L. J. V. (1999) An overview of chondrichthyan systematics and biodiversity in southern Africa. Transactions of the Royal Society of South Africa, 54, 75 - 120. https: // doi. org / 10.1080 / 00359199909520406","Compagno, L. J. V. (2016) Sharks. In: Carpenter, K. E. & De Angelis, N. (Eds.), The Living Marine Resources of the Eastern Central Atlantic. Vol. 2. Bivalves, Gastropods, Hagfishes, Sharks, Batoid Fishes, and Chimaeras. FAO Species Identification Guide for Fishery Purposes. FAO, Rome, pp. 1122 - 1336.","Ebert D. A. (2013) Deep-sea cartilaginous fishes of the Indian Ocean. Vol. 1. Sharks. FAO Species Catalogue for Fishery Purposes No. 8. Vol. 1. FAO, Rome, 256 pp.","Ebert, D. A., Fowler, S. & Compagno, L. J. V. (2013) Sharks of the World: A Fully Illustrated Guide to the Sharks of the World. Wild Nature Press, Plymouth, 528 pp.","Ebert, D. A. (2015) Deep-sea cartilaginous fishes of the Southeastern Atlantic Ocean. FAO Species Catalogue for Fishery Purposes No. 9. FAO, Rome, 251 pp.","Ebert, D. A. & van Hees, K. E. (2015) Beyond jaws: rediscovering the \" Lost Sharks \" of southern Africa. African Journal of Marine Science, 37, 141 - 156. https: // doi. org / 10.2989 / 1814232 X. 2015.1048730","Weigmann, S. (2016) Annotated checklist of the living sharks, batoids and chimaeras (Chondrichthyes) of the world, with a focus on biogeographical diversity. Journal of Fish Biology, 88, 837 - 1037. https: // doi. org / 10.1111 / jfb. 12874"]}

Etmopterus sentosus Bass, D’Aubrey, & Kistnasamy, 1976 Thorny Lanternshark Etmopterus sentosus Bass, D’Aubrey, & Kistnasamy, 1976: 22, figs. 15, 18a. Holotype: SAIAB 6201 [ex ORI 2369]. Type locality: near Bazaruto Island, southern Mozambique. Local synonymy: Etmopterus sentosus: Bass et al., 1976: 22, figs. 15, 18a; Compagno, 1984a: 64, fig.; Bass et al., 1986: 57, fig. 5.15; Compagno et al., 1989: 30, pl.; Compagno, 1999: 115; Compagno et al., 2005: 107, fig., pl. 8.; Ebert, 2013: 82, fig. 101; Ebert et al., 2013 a: 139, fig., pl. 11; Ebert & van Hees, 2015: 144; Weigmann, 2016: 897. South Africa voucher material: SAIAB 6200 [former ORI 1391]. South Africa distribution: Occurs from off Durban to northern KZN. Remarks: This distinctive lanternshark species appears to be common where it occurs, especially off southern Mozambique. Conservation status: LC (2019)., Published as part of Ebert, David A., Wintner, Sabine P. & Kyne, Peter M., 2021, An annotated checklist of the chondrichthyans of South Africa, pp. 1-127 in Zootaxa 4947 (1) on page 30, DOI: 10.11646/zootaxa.4947.1.1, http://zenodo.org/record/4614567, {"references":["Bass, A. J., D'Aubrey, J. D. & Kistnasamy, N. (1976) Sharks of the east coast of southern Africa. VI. The families Oxynotidae, Squalidae, Dalatiidae and Echinorhinidae. Investigational Report. Oceanographic Research Institute, Durban, 45, 1 - 103.","Compagno, L. J. V. (1984 a) FAO Species Catalogue. Sharks of the World. An Annotated and Illustrated Catalogue of Shark Species Known to Date. FAO Fisheries Synopsis. Vol. 4. No. 125. Part 1. Hexanchiformes to Lamniformes. FAO, Rome, pp. 1 - 250.","Bass, A. J. & Compagno, L. J. V. (1986) Families Echinorhinidae, Proscyllidae, Odontaspidiidae, Mitsukurinidae. In: Smith, M. M. & Heemstra, P. C. (Eds.), Smith's Sea Fishes. Macmillan, Johannesburg, pp. 63 + 103 + 104 - 105.","Compagno, L. J. V., Ebert, D. A. & Smale, M. J. (1989) Guide to the Sharks and Rays of Southern Africa. Struik Publishers, Cape Town, 158 pp.","Compagno, L. J. V. (1999) An overview of chondrichthyan systematics and biodiversity in southern Africa. Transactions of the Royal Society of South Africa, 54, 75 - 120. https: // doi. org / 10.1080 / 00359199909520406","Compagno, L., Dando, M. & Fowler, S. (2005) Field Guide to the Sharks of the World. Harper Collins Publishers Ltd, London, 368 pp.","Ebert D. A. (2013) Deep-sea cartilaginous fishes of the Indian Ocean. Vol. 1. Sharks. FAO Species Catalogue for Fishery Purposes No. 8. Vol. 1. FAO, Rome, 256 pp.","Ebert, D. A., Fowler, S. & Compagno, L. J. V. (2013) Sharks of the World: A Fully Illustrated Guide to the Sharks of the World. Wild Nature Press, Plymouth, 528 pp.","Ebert, D. A. & van Hees, K. E. (2015) Beyond jaws: rediscovering the \" Lost Sharks \" of southern Africa. African Journal of Marine Science, 37, 141 - 156. https: // doi. org / 10.2989 / 1814232 X. 2015.1048730","Weigmann, S. (2016) Annotated checklist of the living sharks, batoids and chimaeras (Chondrichthyes) of the world, with a focus on biogeographical diversity. Journal of Fish Biology, 88, 837 - 1037. https: // doi. org / 10.1111 / jfb. 12874"]}

Etmopterus viator Straube, 2011 Traveler Lanternshark Etmopterus viator Straube, in Straube et al., 2011: 143, figs. 2a, b, 3, 5. Holotype: MNHN 2008-1899. Type locality: Kerguelen Plateau, 49°39'29''S, 72°45'00''E, Indian Ocean. Local synonymy: Etmopterus viator; Straube, 2011a: 143, figs. 2a, b, 3, 5 (distribution including South Africa); Ebert, 2013: 84, fig. 116; Ebert et al., 2013 a: 142, fig., pl. 10; Ebert & Mostada, 2013: 34, fig.; Ebert, 2015: 75, fig. 86; Ebert & van Hees, 2015: 144; Weigmann, 2016: 898. South Africa voucher material: None. South Africa distribution: Straube (2011) mentions that this species was recorded from three locations in South African waters, but provides no location information or cites any specimens in museum collection. Remarks: This appears to be a wide-ranging species around seamounts and islands in the Southern Ocean. Conservation status: LC (2018)., Published as part of Ebert, David A., Wintner, Sabine P. & Kyne, Peter M., 2021, An annotated checklist of the chondrichthyans of South Africa, pp. 1-127 in Zootaxa 4947 (1) on page 30, DOI: 10.11646/zootaxa.4947.1.1, http://zenodo.org/record/4614567, {"references":["Straube, N., Kriwet, J. & Schliewen, U. K. (2011 b) Cryptic diversity and species assignment of large lantern sharks of the Etmopterus spinax clade from the Southern Hemisphere (Squaliformes, Etmopteridae). Zoologica Scripta, 40 (1), 61 - 75. https: // doi. org / 10.1111 / j. 1463 - 6409.2010.00455. x","Ebert D. A. (2013) Deep-sea cartilaginous fishes of the Indian Ocean. Vol. 1. Sharks. FAO Species Catalogue for Fishery Purposes No. 8. Vol. 1. FAO, Rome, 256 pp.","Ebert, D. A., Fowler, S. & Compagno, L. J. V. (2013) Sharks of the World: A Fully Illustrated Guide to the Sharks of the World. Wild Nature Press, Plymouth, 528 pp.","Ebert, D. A. (2015) Deep-sea cartilaginous fishes of the Southeastern Atlantic Ocean. FAO Species Catalogue for Fishery Purposes No. 9. FAO, Rome, 251 pp.","Ebert, D. A. & van Hees, K. E. (2015) Beyond jaws: rediscovering the \" Lost Sharks \" of southern Africa. African Journal of Marine Science, 37, 141 - 156. https: // doi. org / 10.2989 / 1814232 X. 2015.1048730","Weigmann, S. (2016) Annotated checklist of the living sharks, batoids and chimaeras (Chondrichthyes) of the world, with a focus on biogeographical diversity. Journal of Fish Biology, 88, 837 - 1037. https: // doi. org / 10.1111 / jfb. 12874"]}

Etmopterus pusillus (Lowe, 1839) Smooth Lanternshark Acanthidium pusillum Lowe, 1839: 91. Syntypes: BMNH 1855.11.29.27 (2 specimens). Type locality: Madeira, eastern Atlantic. Local synonymy: Etmopterus pusillus: Bass et al., 1976: 23, fig. 16 (KZN); Compagno, 1984: 82, fig.; Bass et al., 1986: 56, fig. 5.14; Compagno et al., 1989: 30, pl.; Compagno et al., 1991: 64; Ebert et al., 1992: 605; Compagno, 1999: 115; Compagno et al., 2005: 107, fig., pl. 7; Ebert, 2013: 83, fig. 105; Ebert et al., 2013 a: 137, fig., pl. 9; Ebert & Mostada, 2013: 30, fig.; Ebert, 2015: 80, fig. 93; Ebert & Mostada, 2015: 27, fig.; Ebert & van Hees, 2015: 144; Compagno, 2016: 1188; Weigmann, 2016: 897. South Africa voucher material: SAIAB 6182 [former ORI 1104], SAIAB 6183 [former ORI 1477], SAIAB 6184 [former ORI 2866], SAIAB 6185 [former ORI 2867], SAIAB 6186 [former ORI 2868], SAIAB 6187 [former ORI 2869]. South Africa distribution: West coast from the Orange River (NC) to Cape Point (WC) and off KZN, but not yet confirmed from the EC. Remarks: Widespread, but patchy off South Africa, this species is often misidentified with E. bigelowi. Conservation status: LC (2020)., Published as part of Ebert, David A., Wintner, Sabine P. & Kyne, Peter M., 2021, An annotated checklist of the chondrichthyans of South Africa, pp. 1-127 in Zootaxa 4947 (1) on page 29, DOI: 10.11646/zootaxa.4947.1.1, http://zenodo.org/record/4614567, {"references":["Lowe, R. T. (1839) A supplement to a synopsis of the fishes of Madeira. Proceedings of the Zoological Society of London, 1839 (Pt. 7), 76 - 92.","Bass, A. J., D'Aubrey, J. D. & Kistnasamy, N. (1976) Sharks of the east coast of southern Africa. VI. The families Oxynotidae, Squalidae, Dalatiidae and Echinorhinidae. Investigational Report. Oceanographic Research Institute, Durban, 45, 1 - 103.","Compagno, L. J. V. (1984 a) FAO Species Catalogue. Sharks of the World. An Annotated and Illustrated Catalogue of Shark Species Known to Date. FAO Fisheries Synopsis. Vol. 4. No. 125. Part 1. Hexanchiformes to Lamniformes. FAO, Rome, pp. 1 - 250.","Bass, A. J. & Compagno, L. J. V. (1986) Families Echinorhinidae, Proscyllidae, Odontaspidiidae, Mitsukurinidae. In: Smith, M. M. & Heemstra, P. C. (Eds.), Smith's Sea Fishes. Macmillan, Johannesburg, pp. 63 + 103 + 104 - 105.","Compagno, L. J. V., Ebert, D. A. & Smale, M. J. (1989) Guide to the Sharks and Rays of Southern Africa. Struik Publishers, Cape Town, 158 pp.","Compagno, L. J. V., Ebert, D. A. & Cowley, P. D. (1991) Distribution of offshore demersal cartilaginous fishes (class Chondrichthyes) of the west coast of southern Africa, with notes on their systematics. South African Journal of Marine Science, 11, 43 - 139. https: // doi. org / 10.2989 / 025776191784287664","Ebert, D. A., Compagno, L. J. V. & Cowley, P. D. (1992) A preliminary investigation of the feeding ecology of squaloid sharks off the west coast of southern Africa. South African Journal of Marine Science, 12, 601 - 609. https: // doi. org / 10.2989 / 02577619209504727","Compagno, L. J. V. (1999) An overview of chondrichthyan systematics and biodiversity in southern Africa. Transactions of the Royal Society of South Africa, 54, 75 - 120. https: // doi. org / 10.1080 / 00359199909520406","Compagno, L., Dando, M. & Fowler, S. (2005) Field Guide to the Sharks of the World. Harper Collins Publishers Ltd, London, 368 pp.","Ebert D. A. (2013) Deep-sea cartilaginous fishes of the Indian Ocean. Vol. 1. Sharks. FAO Species Catalogue for Fishery Purposes No. 8. Vol. 1. FAO, Rome, 256 pp.","Ebert, D. A., Fowler, S. & Compagno, L. J. V. (2013) Sharks of the World: A Fully Illustrated Guide to the Sharks of the World. Wild Nature Press, Plymouth, 528 pp.","Ebert, D. A. (2015) Deep-sea cartilaginous fishes of the Southeastern Atlantic Ocean. FAO Species Catalogue for Fishery Purposes No. 9. FAO, Rome, 251 pp.","Ebert, D. A. & van Hees, K. E. (2015) Beyond jaws: rediscovering the \" Lost Sharks \" of southern Africa. African Journal of Marine Science, 37, 141 - 156. https: // doi. org / 10.2989 / 1814232 X. 2015.1048730","Compagno, L. J. V. (2016) Sharks. In: Carpenter, K. E. & De Angelis, N. (Eds.), The Living Marine Resources of the Eastern Central Atlantic. Vol. 2. Bivalves, Gastropods, Hagfishes, Sharks, Batoid Fishes, and Chimaeras. FAO Species Identification Guide for Fishery Purposes. FAO, Rome, pp. 1122 - 1336.","Weigmann, S. (2016) Annotated checklist of the living sharks, batoids and chimaeras (Chondrichthyes) of the world, with a focus on biogeographical diversity. Journal of Fish Biology, 88, 837 - 1037. https: // doi. org / 10.1111 / jfb. 12874"]}

Etmopterus alphus Ebert, Straube, Leslie, & Weigmann, 2016 Whitecheek Lanternshark Etmopterus alphus Ebert, Straube, Leslie, & Weigmann, 2016: 2, figs. 1–6. Holotype: SAM MB-F37564. Type locality: east of the Zambezi River, central Mozambique, 18°14'S, 37°31'E. Local synonymy: Etmopterus alphus: Ebert et al., 2016: 2, figs. 1–6. South Africa voucher material: SAIAB 190352. South Africa distribution: New record. Recently recorded off Durban (KZN). Remarks: This recently described species was known only from off Mozambique, but is reported here for the first time in South African waters. It also occurs off Madagascar and on the Madagascar Ridge. Conservation status: LC (2019).

Etmopterus bigelowi Shirai & Tachikawa, 1993 Blurred Lanternshark Etmopterus bigelowi Shirai & Tachikawa, 1993: 487, figs. 1, 2, 5. Holotype: HUMZ 100176. Type locality: off Angola, 11°37'S, 05°13'W, southeastern Atlantic. Local synonymy: Etmopterus bigelowi: Compagno, 1999: 114; Compagno et al., 2005: 95, fig., pl. 8; Ebert, 2013: 83, fig. 104; Ebert et al., 2013 a: 125, fig., pl. 10; Ebert & Mostarda, 2015: 30, fig.; Ebert, 2015: 75, fig. 81; Ebert & Mostarda, 2015: 27, fig.; Ebert & van Hees, 2015: 144; Compagno, 2016: 1186; Weigmann, 2016: 894. South Africa voucher material: FAKU 46064, FSFL-S 427, HUMZ 74378. Several uncatalogued SAM specimens from off Mossel Bay (WC). South Africa distribution: Scattered records from off the west coast (NC) to KZN. Remarks: The occurrence of this species off South Africa is patchy, but this may be due to misidentification with E. pusillus. Conservation status: LC (2020)., Published as part of Ebert, David A., Wintner, Sabine P. & Kyne, Peter M., 2021, An annotated checklist of the chondrichthyans of South Africa, pp. 1-127 in Zootaxa 4947 (1) on page 28, DOI: 10.11646/zootaxa.4947.1.1, http://zenodo.org/record/4614567, {"references":["Shirai, S. & Tachikawa, H. (1993) Taxonomic resolution of the Etmopterus pusillus species group (Elasmobranchii, Etmopteridae), with description of E. bigelowi, n. sp. Copeia, 1993 (2), 483 - 495. https: // doi. org / 10.2307 / 1447149","Compagno, L. J. V. (1999) An overview of chondrichthyan systematics and biodiversity in southern Africa. Transactions of the Royal Society of South Africa, 54, 75 - 120. https: // doi. org / 10.1080 / 00359199909520406","Compagno, L., Dando, M. & Fowler, S. (2005) Field Guide to the Sharks of the World. Harper Collins Publishers Ltd, London, 368 pp.","Ebert D. A. (2013) Deep-sea cartilaginous fishes of the Indian Ocean. Vol. 1. Sharks. FAO Species Catalogue for Fishery Purposes No. 8. Vol. 1. FAO, Rome, 256 pp.","Ebert, D. A., Fowler, S. & Compagno, L. J. V. (2013) Sharks of the World: A Fully Illustrated Guide to the Sharks of the World. Wild Nature Press, Plymouth, 528 pp.","Ebert, D. A. & Mostarda, E. (2015) Identification Guide to the Deep-sea Cartilaginous Fishes of the Southeastern Atlantic Ocean. FishFinder Programme, FAO, Rome, 70 pp.","Ebert, D. A. (2015) Deep-sea cartilaginous fishes of the Southeastern Atlantic Ocean. FAO Species Catalogue for Fishery Purposes No. 9. FAO, Rome, 251 pp.","Ebert, D. A. & van Hees, K. E. (2015) Beyond jaws: rediscovering the \" Lost Sharks \" of southern Africa. African Journal of Marine Science, 37, 141 - 156. https: // doi. org / 10.2989 / 1814232 X. 2015.1048730","Compagno, L. J. V. (2016) Sharks. In: Carpenter, K. E. & De Angelis, N. (Eds.), The Living Marine Resources of the Eastern Central Atlantic. Vol. 2. Bivalves, Gastropods, Hagfishes, Sharks, Batoid Fishes, and Chimaeras. FAO Species Identification Guide for Fishery Purposes. FAO, Rome, pp. 1122 - 1336.","Weigmann, S. (2016) Annotated checklist of the living sharks, batoids and chimaeras (Chondrichthyes) of the world, with a focus on biogeographical diversity. Journal of Fish Biology, 88, 837 - 1037. https: // doi. org / 10.1111 / jfb. 12874"]}

Etmopterus compagnoi Fricke & Koch, 1990 Brown Lanternshark Etmopterus compagnoi Fricke & Koch, 1990: 2, figs. 1–2. Holotype: SMNS 8999. Type locality: off Cape Town, Western Cape Province, South Africa, 34°41'S, 18°37'E, southeastern Atlantic. Local synonymy: Etmopterus spinax: Gilchrist, 1922b: 49; Barnard, 1925; Barnard, 1947: 20, fig. 5, pl. 3; Smith, 1949a: 59, fig. 51; Smith, 1965: 59, fig. 51; Compagno, 1984: 85, fig. Spinax spinax: Norman, 1935: 37. Etmopterus gracilispinis: Karrer, 1973: 199; Shcherbachev et al., 1978: 186; Compagno, 1984: 76, fig.; Compagno et al., 1989: 28, pl.; Compagno, 2016: 1187. Etmopterus sp.: Bass et al., 1986: 57, fig. 5.16. Etmopterus compagnoi: Fricke & Koch, 1990: 2, figs. 1–2; Compagno et al., 1991: 61; Ebert et al., 1992: 605; Compagno, 1999: 114; Straube et al., 2011a: 146; Ebert, 2013: 84, fig. 114; Ebert et al., 2013 a: 128, fig., pl. 10; Ebert & Mostarda, 2013: 34, fig.; Ebert, 2015: 76, fig. 87; Ebert & Mostarda, 2015: 28, fig.; Ebert & van Hees, 2015: 144; Straube et al., 2015: 11; Weigmann, 2016: 894. Etmopterus unicolor: Compagno et al., 2005: 108, fig., pl. 9. South Africa voucher material: SAIAB 25737, SAIAB 26271, SAIAB 27586, SAIAB 27587, SAIAB 27588, SAIAB 27589, SAIAB 27590, SAIAB 27591, SAIAB 27592, SAIAB 27593, SAIAB 27594, SAIAB 27595, SAIAB 27596, SAIAB 27597, SAIAB 27598, SAIAB 61706, SAIAB 87361, SAIAB 186423, SAIAB 186458, SAIAB 189169, SAIAB 193038. South Africa distribution: The Orange River (NC) to at least Port Alfred (EC) and possibly KZN. Remarks: Etmopterus compagnoi has a convoluted taxonomic history with most early literature accounts referring to it as either E. spinax or E. gracilispinis. However, Straube et al. (2015) found this species to be distinct from these other lanternsharks. Etmopterus compagnoi was thought to be a South African endemic (Compagno et al., 1989; Straube et al., 2015). However, it appears to have a wider geographic range occurring from southern Namibia to Cape Point (WC), where it is quite common, and east to off Port Alfred (EC) (Ebert, 2015). It also occurs off southern Mozambique and the northern Madagascar Plateau. However, specimens from outside the WC and EC should be closely examined to confirm their identification. Conservation status: LC (2019)., Published as part of Ebert, David A., Wintner, Sabine P. & Kyne, Peter M., 2021, An annotated checklist of the chondrichthyans of South Africa, pp. 1-127 in Zootaxa 4947 (1) on page 28, DOI: 10.11646/zootaxa.4947.1.1, http://zenodo.org/record/4614567, {"references":["Fricke, R. & Koch, I. (1990) A new species of the lantern shark genus Etmopterus from southern Africa (Elasmobranchii: Squalidae). Stuttgarter Beitrage zur Naturkunde, Series A (Biologie), 450, 1 - 9.","Gilchrist, J. D. F. (1922 b) Deep-sea fishes procured by the S. A. \" Pickle \" (Part I.). Special Report (3), Report of the Fisheries and Marine Biological Survey, Union of South Africa, 2, 41 - 79.","Barnard, K. H. (1925) A monograph of the marine fishes of South Africa. Part I (Amphioxus, Cyclostomata, Elasmobranchii, and Teleostei-Isospondyli to Heterosomata). Annals of the South African Museum, 21, 1 - 418.","Barnard, K. H. (1947) A Pictorial Guide to South African Fishes. Marine and Freshwater. Maskew Miller Ltd, Cape Town, 226 pp.","Smith, J. L. B. (1949 a) The Sea Fishes of Southern Africa. South Africa Central News Agency Ltd., 550 pp.","Smith, J. L. B. (1965) The Sea Fishes of Southern Africa. 5 th Edition. Central News Agency Ltd., 580 pp.","Compagno, L. J. V. (1984 a) FAO Species Catalogue. Sharks of the World. An Annotated and Illustrated Catalogue of Shark Species Known to Date. FAO Fisheries Synopsis. Vol. 4. No. 125. Part 1. Hexanchiformes to Lamniformes. FAO, Rome, pp. 1 - 250.","Norman, J. R. (1935) Coast fishes, Part. I. The South Atlantic. ' Discovery' Reports, 12, 1 - 58.","Karrer, C. (1973) Uber Fische aus dem S ʾ dostatlantik. Mitteilungen aus dem Zoologischen Museum in Berlin, 49 (1), 191 - 257. https: // doi. org / 10.1002 / mmnz. 4830490113","Shcherbachev, Y. N. (1978) Long-nosed chimaeras (Rhinochimaeridae, Chimaeriformes) from the waters of South Africa. Trudy Institute Okeanology P. P. Shirshova, 111, 7 - 9.","Compagno, L. J. V., Ebert, D. A. & Smale, M. J. (1989) Guide to the Sharks and Rays of Southern Africa. Struik Publishers, Cape Town, 158 pp.","Compagno, L. J. V. (2016) Sharks. In: Carpenter, K. E. & De Angelis, N. (Eds.), The Living Marine Resources of the Eastern Central Atlantic. Vol. 2. Bivalves, Gastropods, Hagfishes, Sharks, Batoid Fishes, and Chimaeras. FAO Species Identification Guide for Fishery Purposes. FAO, Rome, pp. 1122 - 1336.","Bass, A. J. & Compagno, L. J. V. (1986) Families Echinorhinidae, Proscyllidae, Odontaspidiidae, Mitsukurinidae. In: Smith, M. M. & Heemstra, P. C. (Eds.), Smith's Sea Fishes. Macmillan, Johannesburg, pp. 63 + 103 + 104 - 105.","Compagno, L. J. V., Ebert, D. A. & Cowley, P. D. (1991) Distribution of offshore demersal cartilaginous fishes (class Chondrichthyes) of the west coast of southern Africa, with notes on their systematics. South African Journal of Marine Science, 11, 43 - 139. https: // doi. org / 10.2989 / 025776191784287664","Ebert, D. A., Compagno, L. J. V. & Cowley, P. D. (1992) A preliminary investigation of the feeding ecology of squaloid sharks off the west coast of southern Africa. South African Journal of Marine Science, 12, 601 - 609. https: // doi. org / 10.2989 / 02577619209504727","Compagno, L. J. V. (1999) An overview of chondrichthyan systematics and biodiversity in southern Africa. Transactions of the Royal Society of South Africa, 54, 75 - 120. https: // doi. org / 10.1080 / 00359199909520406","Ebert D. A. (2013) Deep-sea cartilaginous fishes of the Indian Ocean. Vol. 1. Sharks. FAO Species Catalogue for Fishery Purposes No. 8. Vol. 1. FAO, Rome, 256 pp.","Ebert, D. A., Fowler, S. & Compagno, L. J. V. (2013) Sharks of the World: A Fully Illustrated Guide to the Sharks of the World. Wild Nature Press, Plymouth, 528 pp.","Ebert, D. A. & Mostarda, E. (2013) Identification Guide to the Deep-sea Cartilaginous Fishes of the Indian Ocean. FishFinder Programme, FAO, Rome, 76 pp.","Ebert, D. A. (2015) Deep-sea cartilaginous fishes of the Southeastern Atlantic Ocean. FAO Species Catalogue for Fishery Purposes No. 9. FAO, Rome, 251 pp.","Ebert, D. A. & Mostarda, E. (2015) Identification Guide to the Deep-sea Cartilaginous Fishes of the Southeastern Atlantic Ocean. FishFinder Programme, FAO, Rome, 70 pp.","Ebert, D. A. & van Hees, K. E. (2015) Beyond jaws: rediscovering the \" Lost Sharks \" of southern Africa. African Journal of Marine Science, 37, 141 - 156. https: // doi. org / 10.2989 / 1814232 X. 2015.1048730","Straube, N., Leslie, R. W., Clerkin, P. J., Ebert, D. A., Rochel, E., Corrigan, S. L., Chenhong, L. & Naylor, G. J. P. (2015) On the occurrence of the southern lanternshark, Etmopteus granulosus, off South Africa, with comments on the validity of E. compagnoi. Deepsea Research II: Topical Studies in Oceanography, 115, 11 - 17. https: // doi. org / 10.1016 / j. dsr 2.2014.04.004","Weigmann, S. (2016) Annotated checklist of the living sharks, batoids and chimaeras (Chondrichthyes) of the world, with a focus on biogeographical diversity. Journal of Fish Biology, 88, 837 - 1037. https: // doi. org / 10.1111 / jfb. 12874","Compagno, L., Dando, M. & Fowler, S. (2005) Field Guide to the Sharks of the World. Harper Collins Publishers Ltd, London, 368 pp."]}

Etmopterus alphus Ebert, Straube, Leslie, & Weigmann, 2016 Whitecheek Lanternshark Etmopterus alphus Ebert, Straube, Leslie, & Weigmann, 2016: 2, figs. 1–6. Holotype: SAM MB-F37564. Type locality: east of the Zambezi River, central Mozambique, 18°14'S, 37°31'E. Local synonymy: Etmopterus alphus: Ebert et al., 2016: 2, figs. 1–6. South Africa voucher material: SAIAB 190352. South Africa distribution: New record. Recently recorded off Durban (KZN). Remarks: This recently described species was known only from off Mozambique, but is reported here for the first time in South African waters. It also occurs off Madagascar and on the Madagascar Ridge. Conservation status: LC (2019)., Published as part of Ebert, David A., Wintner, Sabine P. & Kyne, Peter M., 2021, An annotated checklist of the chondrichthyans of South Africa, pp. 1-127 in Zootaxa 4947 (1) on page 28, DOI: 10.11646/zootaxa.4947.1.1, http://zenodo.org/record/4614567, {"references":["Weigmann, S. (2016) Annotated checklist of the living sharks, batoids and chimaeras (Chondrichthyes) of the world, with a focus on biogeographical diversity. Journal of Fish Biology, 88, 837 - 1037. https: // doi. org / 10.1111 / jfb. 12874","Ebert, D. A., Straube, N., Leslie, R. W. & Weigmann, S. (2016) Etmopterus alphus n. sp.: a new lanternshark (Squaliformes: Etmopteridae) from the south-western Indian Ocean. African Journal of Marine Science, 38 (3), 329 - 340. https: // doi. org / 10.2989 / 1814232 X. 2016.1198275"]}

Centroscyllium fabricii (Reinhardt, 1825) Black Dogfish Spinax fabricii Reinhardt, 1825: 3. Syntypes: ZMUC P 07106 [ex 185] (1 stuffed). Type locality: Julianehåb, Greenland, western North Atlantic. Local synonymy: Centroscyllium fabricii: Compagno, 1984a: 47, fig.; Compagno et al., 1989: 28, pl.; Compagno et al., 1991: 56; Ebert et al., 1992: 603; Compagno, 1999: 114; Ebert et al., 2013 a: 122, fig., pl. 8; NPOA, 2013: 37; Ebert, 2015: 71, fig. 77; Ebert & Mostarda, 2015: 24, fig.; Ebert & van Hees, 2015: 144; Compagno, 2016: 1185; Weigmann, 2016: 893. South Africa voucher material: SAIAB 25708, SAIAB 25736, SAIAB 26338, SAIAB 26339, SAIAB 26340, SAIAB 26424, SAIAB 26425, SAIAB 26426, SAIAB 26427, SAIAB 26428, SAIAB 26429, SAIAB 27183, SAIAB 30309. South Africa distribution: From the Orange River (NC) to southwest of Cape Agulhas (WC). Remarks: Compagno (1984a) reported the first South African record of this species from off the southwestern Cape. Previous southern African records were from off Namibia (Bass et al., 1976). This is one of the most common species of deep-sea lanternsharks off the South African west coast. Conservation status: LC (2020). Genus Etmopterus Rafinesque, 1810a Lanternsharks Etmopterus Rafinesque, 1810a: 14. Type species: Etmopterus aculeatus Rafinesque, 1810a, by monotypy., Published as part of Ebert, David A., Wintner, Sabine P. & Kyne, Peter M., 2021, An annotated checklist of the chondrichthyans of South Africa, pp. 1-127 in Zootaxa 4947 (1) on page 27, DOI: 10.11646/zootaxa.4947.1.1, http://zenodo.org/record/4614567, {"references":["Reinhardt, J. C. H. (1825) Ichthyologiske bidrag. In: H. C. Orsted. Oversigt over det Kongelige Danske Videnskabernes Selskabs Forhandlinger og dets Medlemmers Arbeider (KjObenhavn), 1824 - 25, pp. 1 - 35.","Compagno, L. J. V. (1984 a) FAO Species Catalogue. Sharks of the World. An Annotated and Illustrated Catalogue of Shark Species Known to Date. FAO Fisheries Synopsis. Vol. 4. No. 125. Part 1. Hexanchiformes to Lamniformes. FAO, Rome, pp. 1 - 250.","Compagno, L. J. V., Ebert, D. A. & Smale, M. J. (1989) Guide to the Sharks and Rays of Southern Africa. Struik Publishers, Cape Town, 158 pp.","Compagno, L. J. V., Ebert, D. A. & Cowley, P. D. (1991) Distribution of offshore demersal cartilaginous fishes (class Chondrichthyes) of the west coast of southern Africa, with notes on their systematics. South African Journal of Marine Science, 11, 43 - 139. https: // doi. org / 10.2989 / 025776191784287664","Ebert, D. A., Compagno, L. J. V. & Cowley, P. D. (1992) A preliminary investigation of the feeding ecology of squaloid sharks off the west coast of southern Africa. South African Journal of Marine Science, 12, 601 - 609. https: // doi. org / 10.2989 / 02577619209504727","Compagno, L. J. V. (1999) An overview of chondrichthyan systematics and biodiversity in southern Africa. Transactions of the Royal Society of South Africa, 54, 75 - 120. https: // doi. org / 10.1080 / 00359199909520406","Ebert, D. A., Fowler, S. & Compagno, L. J. V. (2013) Sharks of the World: A Fully Illustrated Guide to the Sharks of the World. Wild Nature Press, Plymouth, 528 pp.","NPOA. (2013) National Plan of Action for the Conservation and Management of Sharks (NPOA-Sharks). Department of Agriculture, Forestry and Fisheries (DAFF), Rogge Bay, Cape Town, 63 pp.","Ebert, D. A. (2015) Deep-sea cartilaginous fishes of the Southeastern Atlantic Ocean. FAO Species Catalogue for Fishery Purposes No. 9. FAO, Rome, 251 pp.","Ebert, D. A. & Mostarda, E. (2015) Identification Guide to the Deep-sea Cartilaginous Fishes of the Southeastern Atlantic Ocean. FishFinder Programme, FAO, Rome, 70 pp.","Ebert, D. A. & van Hees, K. E. (2015) Beyond jaws: rediscovering the \" Lost Sharks \" of southern Africa. African Journal of Marine Science, 37, 141 - 156. https: // doi. org / 10.2989 / 1814232 X. 2015.1048730","Compagno, L. J. V. (2016) Sharks. In: Carpenter, K. E. & De Angelis, N. (Eds.), The Living Marine Resources of the Eastern Central Atlantic. Vol. 2. Bivalves, Gastropods, Hagfishes, Sharks, Batoid Fishes, and Chimaeras. FAO Species Identification Guide for Fishery Purposes. FAO, Rome, pp. 1122 - 1336.","Weigmann, S. (2016) Annotated checklist of the living sharks, batoids and chimaeras (Chondrichthyes) of the world, with a focus on biogeographical diversity. Journal of Fish Biology, 88, 837 - 1037. https: // doi. org / 10.1111 / jfb. 12874","Bass, A. J., D'Aubrey, J. D. & Kistnasamy, N. (1976) Sharks of the east coast of southern Africa. VI. The families Oxynotidae, Squalidae, Dalatiidae and Echinorhinidae. Investigational Report. Oceanographic Research Institute, Durban, 45, 1 - 103.","Rafinesque, C. S. (1810 a) s. n. In: Caratteri di alcuni nuovi generi e nuove specie di animali e piante della Sicilia, con varie osservazioni sopra i medisimi. Part 1 (involves fishes). Sanfilippo, Palermo, pp. 3 - 69. https: // doi. org / 10.5962 / bhl. title. 104418"]}

Etmopterus spinax (Linnaeus 1758) ���Velvet belly lantern shark Taxonomy. First record from Lebanon as Etmopterus spinax by Aguillar et al. (2018: 81). Distribution. Western Baltic Sea, northern North Sea, Mediterranean Sea, eastern Atlantic: Iceland and Norway to Senegal. Conservation. IUCN: Global (LC: 1 December 2008); Med. (LC: 25 March 2016). Capture and threats: Unknown. Occurrence: Very rare. Low priority for conservation action., Published as part of Bariche, Michel & Fricke, Ronald, 2020, The marine ichthyofauna of Lebanon: an annotated checklist, history, biogeography, and conservation status, pp. 1-157 in Zootaxa 4775 (1) on page 19, DOI: 10.11646/zootaxa.4775.1.1, http://zenodo.org/record/3983887

Etmopterus marshae, new species Marsha���s Lanternshark (Figures 1���5; Tables 1, 2) Holotype. PNM 15353, 205 mm [213 mm prior to preservation] total length (TL), maturing male, M/V DA-BFAR, otter trawl, bottom type sandy, station BFAR-583, between Luzon Island and Mindoro Island, Philippines, 13��46'12.6��� to 13��48'53.4��� S, 120��50'54.6��� to 120��50'40.2��� E, 322���337 m, 30 May 2011 (Figures 1a, b). Paratypes. 10 specimens: CAS 234011, males (2 specimens) 234 mm TL, mature, 150 mm TL, immature, females (8 specimens) 97���192 mm TL, maturity undetermined, M/V DA-BFAR, otter trawl, bottom type sandy, station BFAR-583, between Luzon Island and Mindoro Island, Philippines, 13��46'12.6��� to 13��48'53.4��� S, 120��50'54.6��� to 120��50'40.2��� E, 322���337 m, 30 May 2011. Diagnosis. Etmopterus marshae is a relatively small, slender species of linear���denticled Etmopterus that can be separated from its closest congeners within the E. lucifer clade by a combination of characteristics including the length of its luminescent lateral flank marking branches being relatively equal in length; only three other species have anterior and posterior flank branches about equal in length. The new species can be separated from the other species with relatively equal branch lengths, E. burgessi Schaaf-Da Silva & Ebert, 2006, E. evansi Last, S��ret & Burgess, 2002, and E. pycnolepis Kotlyar, 1990, by a combination of characteristics including a whitish cheek blotch, a purplish lateral body coloration sharply becoming black below line between pectoral and pelvic fins, a distinct pattern of paired dashes along the upper body and between pectoral and pelvic fins, dark and light banding pattern on its caudal fin, and central and upper caudal marking lengths distinctly different from these other three closely related species. Description. Values expressed as a percentage of total length (TL) for the holotype (PNM 15353), followed by the ranges for 10 paratypes (97���234 mm TL) are presented in Table 1. The following description proportions include the holotype followed by the paratypes in parentheses. Additional descriptive information on ontogenetic differences between the eight larger specimens (150���234 mm TL) and three smaller specimens (97���116 mm TL) is provided. Body fusiform, trunk sub-cylindrical, width 1.5 (1.1���2.1) in trunk height; head subconical, long, 24.4 (20.7��� 23.3)% TL, slightly depressed, height 0.6 (0.6���0.7) times width. Snout moderately long, conical in lateral view, in dorsal view triangular���shaped becoming rounded at snout���tip, head width 9.3 (8.6���12.0)% TL. Eyes oval-shape, large, length 3.8 (3.8���5.3) in head and 2.2 (1.3���1.7) times height of eye; orbits with anterior and posterior notches; moderately spaced, interorbital space 1.3 (1.0���1.4) in width of head; eye length 1.2 (1.4���1.9) times in interorbital distance. Spiracles small, semi-circular, greatest diameter 2.0 (1.0���2.1)% TL, 3.3 (2.2���4.0) times into length of eye, distance to eye 2.4 (1.8���2.8)% TL, eye���spiracle length 0.7 (0.3���0.7) times into height of eye. Nostrils large, oblique, length about equal to internarial width, less than eye diameter; anterior nasal flap reasonably developed, triangular, anterior tip extending across nasal opening, length 0.8 (0.3���1.0) times spiracle length. Gill openings small, narrow, slightly oblique, in horizontal series, heights about equal, except first opening maybe noticeably larger than last four openings, height of first gill slit 1.3 (1.3���2.0) times height of fifth gill opening, intergill length 1.0 (1.0���1.3) times in length of eye. Mouth broad, length 2.0 (2.2���2.7) times in width, slightly arched, width 0.4 (0.7���0.9) times preoral length. Teeth dissimilar in upper and lower jaw (Figure 2); upper teeth multicuspid in three functional series, functional teeth in lower jaw unicuspid in single series, with at least two series of replacement teeth in upper and lower jaw; multicuspid upper teeth small, perpendicular, with strong central cusp flanked by one or two lateral cusplets on each side, decreasing size distally; first lateral cusplet about one-half height of central cusp, second lateral cusp (if present) much reduced; teeth in lower jaw fused into single row, blade-like, oblique, slightly erect cusp. Tooth count in first row of upper jaw 30 (30���36) and in first row of lower jaw 34 (30���38). First dorsal fin small, anterior margin slightly curved, rounded at apex, becoming relatively straight posteriorly, length of first dorsal fin 7.8 (7.4���10.6)% TL, origin slightly behind pectoral fin insertion; fin base insertion well forward of pelvic-fin origin; pre���first dorsal length 1.6 (1.6���2.0) times interdorsal distance; firstdorsal-fin spine slightly curved near tip towards fin apex, height about equal to fin height, 1.0 (0.6���1.0) times into height of first dorsal fin, located behind pectoral fin rear margin. Second dorsal fin conspicuously larger than first dorsal fin, length of first dorsal fin 0.6 (0.6���0.8) times second dorsal fin, height of first dorsal fin 0.6 (0.5���0.9) times second dorsal fin; apex rounded, posterior margin sharply notched, free rear tip elongated, length 12.7 (10.9��� 13.7)% TL, pre���second dorsal length 2.9 (2.7���3.5) times interdorsal distance; second-dorsal-fin spine large, height slightly higher than fin height, relatively straight, except slightly curving near tip towards fin apex; origin over insertion of pelvic fins, forward of pelvic fin free rear tips. Interspace between first and second dorsal fins 0.8 (0.6��� 0.9) times prepectoral length. Pectoral fins relatively large, length 9.3 (8.6���10.7)% TL, subangular at free rear tips, base 2.3 (1.4���2.6) times in anterior margin, posterior margin frayed. Caudal peduncle length from second dorsal fin insertion to caudal origin relatively long, 14.1 (11.5���14.7)% TL, height about equal to or slightly greater than width, rounded, and tapering posteriorly; less than upper caudal fin length. Caudal fin elongated, slightly less than head length, subterminal notch relatively inconspicuous; length of lower preventral caudal fin margin less than one-half upper caudal fin margin. Dermal denticles hook-like, posteriorly directed rearwards; organized in distinct longitudinal rows laterally, characteristic of E. lucifer clade members. Distance between lateral rows uniformly even, not decreasing from dorsal head surface rearwards towards caudal peduncle. Ventral snout surface densely and uniformly covered with dermal denticles, except for bare area surrounding mouth region (Figure 3); pores (ampullae of Lorenzini) obscured by dense dermal denticle covering snout surface. Dorsal fins, including ceratotrichia, uniformly covered with longitudinal rows of dermal denticles extending continuously onto each from body trunk. Flank-mark-area denticles uniformly dense and pointed ventrally. Dorsal surface of individual denticles without distinct keel. Luminescent markings distinct, intricate; ventral head surface markings blackish, starting just posterior to and slightly lateral to snout tip, extending to level of nostrils and orbits at about anterior notch, descending below eye level to mouth, then extending upwards over mouth corners and encircling mouth; continuous with belly marking posterior to mouth at about first gill openings, demarcated by a faint band of transverse dermal folds across throat extending from below lower edges of first two gill openings on either side. Head dorsal surface photophore pattern faint, originating at about spiracles as single midline along back and flanked on each side by a row, and extending in parallel posteriorly to caudal fin origin. Belly marking originates behind mouth on posterior portion of transverse dermal folds and extends ventrally along pectoral fin bases slightly extending upwards to about level with fin base insertion, about level with ventral edges of gill openings, and posteriorly to pelvic fin bases; ventral surface of pectoral fin lighter along lower base and upper base where ceratotrichia originate, bisected by thin darker band; margin on pectoral���pelvic space clearly defined, line extending from rear margin of pectoral fin base nearly to pelvic fin insertion except for light to whitish mark at pelvic fin origin; dark ventral belly surface continuous onto caudal peduncle to about lower caudal fin origin (Figure 4). Flank markings well defined, consisting of both an anterior and a posterior branch of roughly even length; anterior branch length 9.3 (7.3���10.7)% TL, slender, nearly straight except slightly curving near anterior tip, stopping just before origin of pelvic fin; posterior branch noticeably curved above pelvic fin and thick, width at level of base end of second dorsal fin 2.9 (2.0���4.3)% TL, branch length 8.8 (7.8���10.7)% TL, extending just to distal free rear tip of second dorsal fin; origin anterior to pelvic fin insertion. Caudal base marking distinct, narrow, length 5.9 (1.5���2.5) times longer than base of flank marking. Upper caudal fin marking very narrow, its length 1.2 (0.6���0.9) times upper caudal marking. Vertebral counts: total count 81 (74���85); total precaudal count 58 (51���61); monospondylous 39 (35���40); diplospondylous precaudal 19 (14���22); caudal 23 (21���24). Ontogenetic differences between large and small specimens. Etmopterus marshae sp. nov. exhibits several subtle changes with growth between the three smallest specimens (97���116 mm TL) and the eight largest individuals, including the holotype (150���234 mm TL), the largest of which is an adult male. The following character differences between large and small type specimens included: prenarial length 3.9���4.1% vs 2.0���2.6% TL, preoral length 10.8���11.3% vs 9.3���10.9% TL, preorbital length 6.9���7.2% vs 5.3���6.8% TL, orbit length 5.2���6.9% vs 4.2���5.6% TL, orbit height 4.9���5.2 vs 3.0���4.0% TL, interorbital length 8.8���10.3% vs 7.3���8.7% TL, spiracle length 2.1���2.9% vs 1.0���2.1% TL. The coloration was more pronounced in the smaller individuals, but overall the color patterns were consistent in both size groups. Coloration. Etmopterus marshae in life is a dark purplish black dorsally and laterally on body trunk, becoming dark black ventrally; transition between lateral and ventral surfaces sharply demarcated, becoming distinctly black from ventral surface of snout tip to about midpoint of anterior eye notch, extending semi-circular below eye orbit, and from below posterior eye notch extending posteriorly to pectoral fin origin; distance between pectoral-pelvic fin length sharply demarcated except for lighter areas behind pectoral fin insertion and pelvic fin origin; distance between pelvic fin insertion and lower caudal fin origin lighter except for flank marking base behind pelvic fin insertion and dark black blotch between posterior flank marking and caudal fin base marking (Figure 1a). Eyeball a solid greenish. Faint white spot on each cheek located just behind eye. Dorsal view of body surface uniformly dark, without lighter or white stripe; prominent complex pattern of dashes along body trunk with 1���3 rows of dark dashes extends from above dorsal head surface above lateral line to caudal origin, with a separate paired row of dashes extending between pectoral and pelvic fins. Gills and area just above them purplish to lighter, but area below sharply demarcated to black. Pectoral and pelvic fins dark at base and along anterior fin edge, becoming translucent to white on remainder of fins. Dorsal fins purplish black to black at base and along anterior edges, becoming translucent to white on remainder of fins. Black lateral flank markings sharply demarcated by surrounding lighter colored lateral flanks; prominent lighter colored flank area extending dorsally approximately length of darker lateral flank markings; lighter purplish colored flank area extending on either side of dark-flankmarking base located just posterior to pelvic fins; prominent lighter colored flank area originates anterior to darkflank-base marking and extends anteriorly to about pectoral fin insertion; lighter colored flank area posterior to dark flank base and below posterior branch of lateral lighter flank coloration noticeable, but not as prominent as faint whitish flank area between pectoral and pelvic fins. Lateral lighter colored flank area between pectoral and pelvic fins sharply demarcates lateral and ventral surfaces; ventral surface black. Prominent black blotch between posterior tip of flank marking and lower caudal origin. Caudal fin with distinct dark black bar originating at upper caudal fin origin and fading posteriorly to a lighter or white, and with a prominent black caudal fin tip; upper and lower ventral lobe mostly black. Ventral surface mostly dark black around mouth and belly. A dark stripe originates between pelvic fin insertions and extends to caudal origin. Area around mouth distinctly black, surface bare of dermal denticles, sharply demarcated from coarse dermal denticles surrounding it. After preservation coloration similar except purplish coloration fades to light or dark brown; dorsal white stripe, white cheek spot, and whitish flank area and black lateral flank markings prominent; dark bar on caudal fin and darker dashes extending laterally on body still clearly visible; area around gills becoming whitish (Figure 1b). Size. A 234 mm TL male (CAS 234011) was mature, while a 213 mm TL male (Holotype PNM 15353) was maturing. The largest female was 192 mm TL, but its maturity was not determined. Birth size uncertain, although the three smallest paratypes ranging 97���116 mm TL all had umbilical scars, with the umbilical scar slightly open on the smallest specimen, suggesting that this individual was close to birth size. Distribution. The new species is known from 11 specimens collected on a single haul between the Philippine islands of Luzon Island and Mindoro (13�� 46��� 12.6���E 120�� 50��� 54.6���S to 13�� 48��� 53.4���E 120�� 50��� 40.2���S), and at a depth range of 322���337 m (Figure 5). The bottom type where the specimens were collected was sandy. Etymology. The species is named after Marsha Englebrecht for her innovative contributions to the field of elasmobranch husbandry. The proposed common name is Marsha���s Lanternshark., Published as part of Ebert, David A. & Van Hees, Kelley E., 2018, Etmopterus marshae sp. nov, a new lanternshark (Squaliformes: Etmopteridae) from the Philippine Islands, with a revised key to the Etmopterus lucifer clade, pp. 197-210 in Zootaxa 4508 (2) on pages 198-202, DOI: 10.11646/zootaxa.4508.2.3, http://zenodo.org/record/2607000, {"references":["Schaaf-DaSilva, J. A. & Ebert, D. A. (2006) Etmopterus burgessi sp. nov., a new species of lanternshark (Squaliformes: Etmopteridae) from Taiwan. Zootaxa, 1373, 53 - 64.","Last, P. R., Burgess, G. H. & Seret, B. (2002) Description of six new species of lantern-sharks of the genus Etmopterus (Squaloidea: Etmopteridae) from the Australasian region. Cybium, 26 (3), 203 - 223.","Kotlyar, A. N. (1990) Dogfish sharks of the genus Etmopterus Rafinesque from the Nazca and Sala y Gomez submarine ridges. Trudy Instituta Okeanologii Imeni P. P. Shirshova, 125, 127 - 147."]}

A new species of lanternshark, Etmopterus marshae (Squaliformes: Etmopteridae), is described from the Philippine Islands, western North Pacific Ocean. The new species occurs along insular slopes at a depth range of 322–337 m. The new species resembles other members of the “Etmopterus lucifer” clade in having linear rows of dermal denticles, and most closely resembles E. burgessi from Taiwan and E. evansi and E. pycnolepis from the South Pacific. It can be distinguished from all other members of the E. lucifer clade by a combination of characteristics, including length of anterior and posterior flank branches being of relatively equal length, straight vs. curved anterior flank marking, relative lengths of caudal markings, coloration, and relatively small size. A revised key to the revised key to the Etmopterus lucifer clade is provided.

Genus Etmopterus Rafinesque 1810 Etmopterus Rafinesque 1810: 14. Type species: Etmopterus aculeatus Rafinesque 1810 by monotypy., Published as part of Ebert, David A. & Van Hees, Kelley E., 2018, Etmopterus marshae sp. nov, a new lanternshark (Squaliformes: Etmopteridae) from the Philippine Islands, with a revised key to the Etmopterus lucifer clade, pp. 197-210 in Zootaxa 4508 (2) on page 198, DOI: 10.11646/zootaxa.4508.2.3, http://zenodo.org/record/2607000

Key to the Etmopterus lucifer clade of Lanternsharks 1a. Lateral flank marking anterior and posterior branches dissimilar in length........................................ 2. 1b. Lateral flank marking anterior and posterior branches nearly equal in length...................11. (E. burgessi subclade). 2b. Lateral flank marking anterior branch noticeably longer than posterior branch in length.............3 (E. lucifer subclade). 2a. Lateral flank marking anterior branch noticeably shorter than posterior branch in length............5 (E. molleri subclade). 3a. Body slender; ventral snout surface between nostrils bare, without dermal denticles; central caudal marking length 6 %TL or more................................................. Etmopterus lailae (Central North Pacific: Hawaiian Islands) 3b. Body stout; ventral snout surface between nostrils uniformly covered with dermal denticles, and depending on the species may or may not have small bare patches; central caudal marking length less than 5.1 %TL............................... 4. 4a. Flank marking anterior branch nearly twice the length of the posterior branch; area between nostrils without small bare patches. Etmopterus lucifer...........................................(Western North Pacific: Japan to Philippines) 4b. Flank marking anterior branch only slightly longer than posterior branch; area between nostrils with small bare patches..................... Etmopterus sculptus. (Southeast Atlantic and Southwest Indian Ocean: Namibia to southern Mozambique) 5a. Posterior branch of lateral flank marking extends past second dorsal fin free rear tip................................ 6. 5b. Posterior branch of lateral flank marking extends does not past second dorsal fin free rear tip......................... 8. 6a. Flanks with elaborate pattern of conspicuous rows of dark dots and dashes............................................................................ Etmopterus dislineatus. (Southwestern Pacific: off northern Queensland, Australia) 6b. Flanks without elaborate pattern of conspicuous rows of dark dots and dashes..................................... 7. 7a. Anterior (6.0���10.6%) and posterior (9.3���12.7%) flank branches relatively shorter................................................................................................. Etmopterus alphus. (Southwestern Indian Ocean) 7b. Anterior (8.0���11.5%) and posterior (11.0���15.4%) flank branches relatively longer............................................................................ Etmopterus molleri. (Western Pacific: eastern Australasia to New Zealand) 8a. Caudal fin lacks a band or spot.......................................................................... 9. 8a. Caudal fin with a band or spot......................................................................... 10. 9a. Upper teeth with 4 or 5 pairs of cusplets on each side; dermal denticles not arranged in rows....................................................................... Etmopterus decacuspidatus. (Western North Pacific: South China Sea) 9b. Upper teeth with less than 4 or 5 pairs of cusplets on each side; dermal denticles arranged in rows............................................................................................. Etmopterus bullisi. (Western Atlantic) 10a. Length of posterior flank marking 11.4���12.6% TL, flank marking base length 2.8���3.4% TL, and length of caudal base marking length from tip of anterior finger to tip of posterior finger 7.0���7.9% TL. Etmopterus brachyurus.............................................................................. (Western Pacific: Japan to Philippines, possibly Australia) 10b. Length of posterior flank marking 9.1���11.2% TL, flank marking base length 3.5���4.9% TL, and length of caudal base marking length from tip of anterior finger to tip of posterior finger 10.6���14.1% TL.............................................................................................. Etmopterus samadiae. (Western Pacific: Papua New Guinea) 11a. Caudal fin without dark band at fin tip.......................... Etmopterus burgessi. (Western North Pacific: Taiwan) 11b. Caudal fin with dark band at fin tip...................................................................... 12. 12a. Caudal peduncle without dark saddle; caudal fin without dark band across middle of upper lobe and at fin tip.......................................... Etmopterus pycnolepis. (Southeastern Pacific: Nazca and Sala y Gomez submarine ridges) 12b. Caudal peduncle with dark saddle; caudal fin with dark band across middle of upper lobe and at fin tip................ 13. 13a. Caudal fin ventral lobe and upper post-ventral margin mostly white to translucent............................................... Etmopterus evansi. (Eastern Indian Ocean: off northern Western Australia, Indonesia, and Papua New Guinea) 13b. Caudal fin ventral lobe and upper post-ventral margin mostly black................................................................................................ Etmopterus marshae n. sp. (Western North Pacific: Philippines), Published as part of Ebert, David A. & Van Hees, Kelley E., 2018, Etmopterus marshae sp. nov, a new lanternshark (Squaliformes: Etmopteridae) from the Philippine Islands, with a revised key to the Etmopterus lucifer clade, pp. 197-210 in Zootaxa 4508 (2) on page 207, DOI: 10.11646/zootaxa.4508.2.3, http://zenodo.org/record/2607000

Genus Etmopterus Rafinesque, 1810a Lanternsharks Etmopterus Rafinesque, 1810a: 14. Type species: Etmopterus aculeatus Rafinesque, 1810a, by monotypy., Published as part of White, William T. & Ko'Ou, Alfred, 2018, An annotated checklist of the chondrichthyans of Papua New Guinea, pp. 1-82 in Zootaxa 4411 (1) on page 14, DOI: 10.11646/zootaxa.4411.1, http://zenodo.org/record/1221878

Etmopterus samadiae White, Ebert, Mana & Corrigan, 2017c Papuan Lanternshark Etmopterus samadiae White, Ebert, Mana & Corrigan, 2017c: 340, figs 1–6, 7d, 8. Holotype: NTUM 10078. Type locality: Madang, Papua New Guinea. Local synonymy: Etmopterus sp.— Fricke et al., 2014: 14 (Madang Province). Etmopterus samadiae —White et al. (2017c): 340, figs 1–6, 7d, 8 (northern PNG); White et al., 2018: 56, figs (PNG). PNG voucher material: (17 spec.) ASIZ P.73777 (paratype), adult male 230 mm TL, ASIZ P.73778 (paratype), female 188 mm TL, ASIZ P.73765 (paratype), pregnant female 277 mm TL, off Lae, Huon Gulf, Morobe Province, 06°51.841' S, 147°04.672' E, 395–406 m depth, 22 Aug. 2010; ASIZ P0080739 (6 specimens), 235–275 mm TL, off Madang, 5°00.43' S, 145°49.74' E, 460–466 m depth, 2 Oct. 2010; ASIZ P0080734, female 224 mm TL, off Madang, 5°01.27' S, 145°49.74' E, 502–529 m depth, 2 Oct. 2010; NTUM 10078 (holotype), adult male 265 mm TL, east of Malmal Passage, Madang, 05°07' S, 145°50' E, 527–539 m depth, 30 Nov. 2012; NTUM 10313 (paratype), female 269 mm TL, northern Cape King William, Morobe Province, 06°00' S, 147°38' E, 785 m depth, 10 Dec. 2012; NTUM 10314 (paratype), female 258 mm TL, Astrolabe Bay, Madang, 05°22' S, 145°48' E, 420–490 m depth, 14 Dec 2012; NTUM 10315 (paratype), female 154 mm TL, Astrolabe Bay, Madang, 05°22' S, 145°48' E, 340–385 m depth, 14 Dec. 2012; NTUM 10316 (3 specimens; paratypes), female 177 mm TL, subadult male 201 mm TL, female 228 mm TL, west of Kairiru Island, East Sepik, 03°19' S, 143°27' E, 422–425 m depth, 19 Dec. 2012. Remarks: First recorded in PNG by Fricke et al. (2014) as Etmopterus sp.; recently described as a new species by White et al. (2017c), endemic to PNG. Most similar to Etmopterus brachyurus (Smith & Radcliffe in Smith, 1912) from the Northwest Pacific. Endemic to PNG., Published as part of White, William T. & Ko'Ou, Alfred, 2018, An annotated checklist of the chondrichthyans of Papua New Guinea, pp. 1-82 in Zootaxa 4411 (1) on page 15, DOI: 10.11646/zootaxa.4411.1, http://zenodo.org/record/1221878, {"references":["Fricke, R., Allen, G. R., Andrefouet, S., Chen, W. - J., Hamel, M. A., Laboute, P., Mana, R., Hui, T. H. & Uyeno, D. (2014) Checklist of the marine and estuarine fishes of Madang District, Papua New Guinea, western Pacific Ocean, with 820 new records. Zootaxa, 3832 (1), 1 - 247. https: // doi. org / 10.11646 / zootaxa. 3832.1.1","Finucci, B., White, W. T., Kemper, J. & Naylor, G. J. P. (2018) Redescription of Chimaera ogilbyi (Chimaeriformes; Chimaeridae) from the Indo-Australian region. Zootaxa, 4375, 191 - 210. https: // doi. org / 10.11646 / zootaxa. 4375.2.2"]}

Etmopterus fusus Last, Burgess & Séret, 2002 Pygmy Lanternshark Etmopterus fusus Last, Burgess & Séret 2002: 217, Figs 1b, 3b, 5b, 7b. Holotype: CSIRO H 3149-06. Type locality: off Dampier Archipelago, Western Australia. Local synonymy: Etmopterus fusus —White et al. (2017c): 339, Figs 11c, 13 (PNG); White et al., 2018: 54, figs (PNG). PNG voucher material: (1 spec.) NTUM 10318, female 256 mm TL, north of Taviltae, Madang, 500–510 m depth, 17 Dec. 2012. Remarks: First recorded from PNG by White et al. (2017c) based on a specimen collected from deepwater. Previously known from only 7 specimens from northwestern Australia in depths of 430–550 m; possibly also off Java in Indonesia at depths of 120– 200 m., Published as part of White, William T. & Ko'Ou, Alfred, 2018, An annotated checklist of the chondrichthyans of Papua New Guinea, pp. 1-82 in Zootaxa 4411 (1) on pages 14-15, DOI: 10.11646/zootaxa.4411.1, http://zenodo.org/record/1221878, {"references":["Last, P. R., Burgess, G. H. & Seret, B. (2002) Description of six new species of lantern-sharks of the genus Etmopterus (Squaloidea: Etmopteridae) from the Australasian region. Cybium, 26 (3), 203 - 223.","Finucci, B., White, W. T., Kemper, J. & Naylor, G. J. P. (2018) Redescription of Chimaera ogilbyi (Chimaeriformes; Chimaeridae) from the Indo-Australian region. Zootaxa, 4375, 191 - 210. https: // doi. org / 10.11646 / zootaxa. 4375.2.2"]}

Etmopterus evansi Last, Burgess & Séret, 2002 Blackmouth Lanternshark Etmopterus evansi Last, Burgess & Séret 2002: 214, Figs 2c, 4c, 6c, 8c. Holotype: CSIRO H 3141-16. Type locality: off Rowley Shoals, Western Australia. Local synonymy: Etmopterus evansi —White et al., 2017c: 339, Figs 11b, 12 (Madang and Manus Provinces); White et al., 2018: 52, figs (PNG). PNG voucher material: (4 spec.) ASIZ P. P0080732, adult male 343 mm TL, west of Manus Island, 679–685 m depth, 29 Sep. 2010; ASIZ P. P0080733, female 177 mm TL, Astrolabe Bay, Madang, 608–610 m depth, 5 Oct. 2010; NTUM 10312, juvenile male 172 mm TL, Astrolabe Bay, Madang, 520–575m depth, 14 Dec. 2012; NTUM 10317, male 299 mm TL, east of Cape Croisiles, Madang, 680–689 m depth, 16 Dec. 2012;. Remarks: First recorded from PNG by White et al. (2017c) based on four specimens collected from deepwater. Previously known from northwestern Australia and eastern Indonesia at depths of 430–555 m, thus represent the deepest records for this species (to 689 m depth)., Published as part of White, William T. & Ko'Ou, Alfred, 2018, An annotated checklist of the chondrichthyans of Papua New Guinea, pp. 1-82 in Zootaxa 4411 (1) on page 14, DOI: 10.11646/zootaxa.4411.1, http://zenodo.org/record/1221878, {"references":["Last, P. R., Burgess, G. H. & Seret, B. (2002) Description of six new species of lantern-sharks of the genus Etmopterus (Squaloidea: Etmopteridae) from the Australasian region. Cybium, 26 (3), 203 - 223.","Finucci, B., White, W. T., Kemper, J. & Naylor, G. J. P. (2018) Redescription of Chimaera ogilbyi (Chimaeriformes; Chimaeridae) from the Indo-Australian region. Zootaxa, 4375, 191 - 210. https: // doi. org / 10.11646 / zootaxa. 4375.2.2"]}

Etmopterus evansi Last, Burgess & Séret, 2002 Blackmouth Lanternshark Etmopterus evansi Last, Burgess & Séret 2002: 214, Figs 2c, 4c, 6c, 8c. Holotype: CSIRO H 3141-16. Type locality: off Rowley Shoals, Western Australia. Local synonymy: Etmopterus evansi —White et al., 2017c: 339, Figs 11b, 12 (Madang and Manus Provinces); White et al., 2018: 52, figs (PNG). PNG voucher material: (4 spec.) ASIZ P. P0080732, adult male 343 mm TL, west of Manus Island, 679–685 m depth, 29 Sep. 2010; ASIZ P. P0080733, female 177 mm TL, Astrolabe Bay, Madang, 608–610 m depth, 5 Oct. 2010; NTUM 10312, juvenile male 172 mm TL, Astrolabe Bay, Madang, 520–575m depth, 14 Dec. 2012; NTUM 10317, male 299 mm TL, east of Cape Croisiles, Madang, 680–689 m depth, 16 Dec. 2012;. Remarks: First recorded from PNG by White et al. (2017c) based on four specimens collected from deepwater. Previously known from northwestern Australia and eastern Indonesia at depths of 430–555 m, thus represent the deepest records for this species (to 689 m depth).

Etmopterus samadiae, new species Papuan Lanternshark (Figures 1–10; Table 1) Etmopterus sp.— Fricke et al., 2014: 14 (Madang) Holotype. NTUM 10078 (tissue accession GN 17184), adult male 265 mm TL, east of Malmal Passage, Madang, Papua New Guinea, 05°07' S, 145°50' E, 527–539 m depth, 30 Nov 2012. Paratypes. 9 specimens: ASIZ P.73777, adult male 230 mm TL, ASIZ P.73778, female 188 mm TL, ASIZ P.73765, pregnant female 277 mm TL, off Lae, Huon Gulf, Morobe Province, Papua New Guinea, 06°51.841' S, 147°04.672' E, 395–406 m depth, 22 Aug 2010; NTUM 10313 (tissue accession GN 17195), female 269 mm TL, northern Cape King William, Morobe Province, Papua New Guinea, 06°00' S, 147°38' E, 785 m depth, 10 Dec 2012; NTUM 10314 (tissue accession GN 17197), female 258 mm TL, Astrolabe Bay, Madang, Papua New Guinea, 05°22' S, 145°48' E, 420–490 m depth, 14 Dec 2012; NTUM 10315 (tissue accession GN 17198), female 154 mm TL, Astrolabe Bay, Madang, Papua New Guinea, 05°22' S, 145°48' E, 340–385 m depth, 14 Dec 2012; NTUM 10316 (3 specimens; tissue accessions GN 17210–2), female 177 mm TL, subadult male 201 mm TL, female 228 mm TL, west of Kairiru Island, East Sepik, Papua New Guinea, 03°19' S, 143°27' E, 422–425 m depth, 19 Dec 2012. Diagnosis. Etmopterus samadiae is a relatively small, slender, species of linear–denticled Etmopterus that can be separated from its closest congeners within the E. lucifer clade by a combination of characteristics including the length of its anterior flank markings being slightly shorter than its posterior branch, long caudal base marking, and irregular and variable number of black, horizontal, dash-like marks on sides of body. The new species is morphologically and genetically (based on the NADH 2 marker) closest to E. brachyurus, but differs from this species in having a shorter posterior caudal marking (2.8–4.4 vs. 4.2–6.1% TL), a longer caudal base marking (10.6–14.1 vs. 7.0–7.8% TL), and flank marking with a slightly shorter posterior branch (9.1–11.2 vs. 11.4–12.6% TL). Description. Values expressed as a percentage of total length (TL) for the holotype, followed by the range of values for 9 paratypes (Table 1). Body fusiform, trunk sub-cylindrical (Fig. 1), width 1.1 (0.7–1.7) in trunk height; head sub-conical, long, 21.3 (20.6–23.7)% TL, slightly depressed, height 0.7 (0.5–0.8) times width. Snout moderately long, conical in lateral view, in dorsal view triangular–shaped becoming rounded at snout–tip (Fig. 3), head width 8.2 (9.8–11.8)% TL. Eyes oval-shape, large, orbit length 3.8 (3.0–3.7) in head and 2.6 (2.0–3.3) times orbit height; orbits with anterior and posterior notches; moderately spaced, inter–orbital space 1.2 (1.2–1.5) in width of head and orbit length 1.2 (1.1–1.4) times in inter–orbital distance. Spiracles small, semi-circular, greatest diameter 0.9 (1.4–2.5)% TL, 6.1 (2.7–4.0) times orbit length, distance to eye 2.9 (1.4–2.6)% TL, eye–spiracle length 0.7 (1.0–1.9) in orbit height. Nostrils large, oblique, length almost equal to internarial width, less than orbit length; anterior nasal flap well developed, triangular, anterior tip extending across nasal opening, length 1.0 (0.5–0.9) times spiracle length. Gill openings small, narrow, slightly oblique, in horizontal series, subequal in height, inter-gill length 4.9 (3.0–4.7)% TL. Mouth broad, length 3.4 (3.3–4.8) times in width, slightly arched, width 0.8 (0.7–0.9) times preoral length. Teeth dissimilar in upper and lower jaw (Fig. 4); upper teeth multicuspid in three functional series, functional teeth in lower jaw unicuspid in single series; multicuspid upper teeth small, upright, with strong central cusp flanked by 2 or 3 lateral cusplets on each side, decreasing in size distally; teeth in lower jaw fused into single row, blade-like, cusp oblique. Tooth count in first row of upper jaw 33 (27–28) and in first row of lower jaw 35 (28–31). First dorsal fin small, rounded at apex, length of first dorsal fin 8.6 (8.5–9.9)% TL, origin just anterior to pectoral-fin free rear tip; fin base insertion well anterior of pelvic-fin origin; pre–first dorsal fin length 1.3 (1.4–1.8) times inter–dorsal distance; first dorsal–fin spine straight, short, 1.6 (1.3–2.0) times height of first dorsal fin, located over pectoral fin rear margin. Second dorsal fin conspicuously larger, more erect than first dorsal fin, length of first dorsal fin 0.6 (0.6–0.8) times second dorsal fin, height of first dorsal fin 0.4 (0.5–0.7) times second dorsal fin; apex sub–angular, posterior margin concave, free rear tip elongated, length 13.2 (12.2–13.6)% TL, pre–second dorsal length 2.5 (2.6–3.1) times inter–dorsal distance; second dorsal–fin spine large, height slightly taller than fin, curved near tip towards fin apex; origin posterior to insertion of pelvic fins, over pelvic fin free rear tips. Interspace between first and second dorsal fins 1.0 (0.8–1.0) times pre–pectoral length. Pectoral fins relatively large, length 9.4 (10.1–11.3)% TL, subangular at free rear tips, base 2.0 (1.7–2.1) times in anterior margin, posterior margin nearly straight. Caudal peduncle relatively long, dorsal-caudal space 16.1 (13.6–15.3)% TL, height slightly greater than width, rounded, and tapering posteriorly. Caudal fin elongate, subequal to head length, terminal lobe distinct; length of lower preventral caudal fin margin less than one-half upper caudal fin margin. E. samadiae n.sp. E. fusus E. evansi E. brachyurus Holotype Paratypes NTUM n = 4 n = 4 Min. Max. 10318 Min. Max. Min. Max. Total length (mm) 265 154 277 256 172.0 343.0 224 350 ......continued on the next page E. samadiae n.sp. E. fusus E. evansi E. brachyurus Holotype Paratypes NTUM n = 4 n = 4 ......continued on the next page E. samadiae n.sp. E. fusus E. evansi E. brachyurus Holotype Paratypes NTUM n = 4 n = 4 Min. Max. 10318 Min. Max. Min. Max. Dermal denticles hook-like, posteriorly directed rearwards; organized in distinct rows laterally, characteristic of E. lucifer clade members (Fig. 5, 6). Distance between lateral rows mostly consistent along length, decreasing only very slightly towards caudal peduncle. Flank mark area denticles more dense and pointed ventrally. Luminescent markings distinct, intricate (Fig. 7 D); ventral head surface markings blackish, starting from almost at snout tip, extending to level of nostrils and orbits at just below level of anterior notch, then extending straight from just below posterior eye notch towards gill slits, weakly demarcated from belly marking by a weak band of transverse dermal folds across throat extending from below lower edges of first three gill openings on either side. Head dorsal surface photophore pattern as follows: a single midline along back originating at level of first gill slits extending posteriorly to caudal–fin origin; scattered photophores on paler fontanelle area of central head and also on paler area above orbits (as short dashes in some paratypes). Belly marking originates behind mouth on posterior portion of transverse dermal folds and extends ventrally along pectoral fin bases extending upwards to level with fin origin and upper edges of gill slits, and posteriorly to pelvic fin bases; ventral surface of pectoral fin very dark along lower base and upper base where ceratotrichia originate, bisected by distinct lighter to white band or blotch forming a two–prong fork pattern; margin on lateral trunk (pectoral–pelvic space) very clearly defined, line extending from rear margin of pectoral–fin base nearly to pelvic–fin insertion except for paler area at pelvic-fin origin; dark ventral belly surface continuous onto caudal peduncle to about lower caudal fin origin. Flank markings well defined (Fig. 7 d), consisting of both an anterior and posterior branch; anterior branch relatively short, length 10.3 (8.2–10.5)% TL, slender, nearly straight, extending well anterior to pelvic–fin origin; posterior branch straight, slightly thicker, width at level of base end of second dorsal fin 0.6 (0.5–0.8)% TL, slightly longer than anterior branch (shorter than anterior branch in paratype ASIZ P.73777), length 11.2 (9.1– 10.8)% TL, extending to just anterior to second dorsal-fin free rear tip but well posterior to second dorsal-fin insertion; base of flank marking wide, origin slightly posterior to pelvic-fin insertion. Ventral caudal base marking distinct, short, length about equal to base of flank marking, not extending as a saddle on to caudal peduncle, anterior finger very short, posterior finger long, total length (including base) 11.4 (10.6–14.1)% TL. No central caudal marking. Posterior caudal fin marking very narrow, its length 3.2 (2.8–4.4) % TL. Vertebral counts: total counts 80 (80–86), total precaudal counts 55 (55–59); monospondylous 34 (36–37); diplospondylous precaudal 21 (19–22); caudal 25 (25–27). Coloration. In life, greyish to silvery black dorsally and laterally on body, becoming dark black ventrally; transition between lateral and ventral surfaces well demarcated by a paler lateral stripe below the flank markings in most specimens (less distinct in holotype and some paratypes). Dorsal midline with a broad, pale stripe originating just posterior to midpoint of inter-spiracle space, extending to first dorsal fin origin, continuing between dorsal fins, and from second dorsal fin insertion to upper caudal fin origin (Fig. 2); most prominent anterior to first dorsal fin and between dorsal fins, less prominent along upper caudal peduncle; a single row of dark photophores extending along middle of white dorsal stripe. Body with variable number and arrangement of short, horizontal, dash-like black markings (Fig. 8); most numerous in paratype NTUM 10314 (Fig. 8). Pectoral and pelvic fins dark at base and along anterior fin edge, becoming translucent to white on remainder of fins. Dorsal fins dark at base and along anterior edges, becoming translucent to white on remainder of fins. Black lateral flank markings demarcated by surrounding lighter colored lateral flanks (not sharply demarcated in most preserved specimens). Paler colored flank area on lateral surface between pectoral and pelvic fins sharply demarcates lateral and ventral surfaces; ventral surface black. Caudal fin with a distinct, large dark blotch centrally, occupying the area between the posterior finger of the caudal base marking and the upper caudal marking; posterior margin dark. Ventral surface mostly dark black around mouth, belly, and with a dark stripe between pelvic fin insertions and caudal origin; snout not distinctly paler than mouth. After preservation coloration similar but with paler markings often less obvious and coloration becoming a light or dark brown; dorsal median pale stripe and whitish flank area and black lateral flank markings less prominent but still distinct in most cases; dark blotch on central caudal fin obvious and darker pores extending laterally on body still clearly visible under microscope. Molecular analysis. The analysis of the NADH 2 data confirms that Etmopterus samadiae belongs in the E. lucifer clade and represents a monophyletic lineage that is distinct from, but most closely related to Etmopterus brachyurus (Fig. 9). It should be noted that this inference is based on a single mitochondrial marker. Inclusion of multiple nuclear markers could affect the presented inference. Size. Specimens examined ranged in size from 154 to 277 mm TL. A 277 mm TL female (ASIZ P.73765) was found to be pregnant (embryo caudal fin visible in cloaca) and two males 230 (ASIZ P.73777) and 265 mm TL (NTUM 10078) were determined to be mature. Distribution. The new species is known from off the northern Papua New Guinea mainland, from west of Kairiru Island in East Sepik Province to off Lae in the Huon Gulf (Morobe Province) and at a depth range of 340 to 785 m (Fig. 10). Etymology. The species is named after Dr Sarah Samadi (MNHN) who was one of the key principal investigators of the 2010 and 2012 MNHN expeditions in Papua New Guinea from which all the type specimens were obtained. The proposed English common name is Papuan Lanternshark.

Etmopterus samadiae, new species Papuan Lanternshark (Figures 1���10; Table 1) Etmopterus sp.��� Fricke et al., 2014: 14 (Madang) Holotype. NTUM 10078 (tissue accession GN 17184), adult male 265 mm TL, east of Malmal Passage, Madang, Papua New Guinea, 05��07' S, 145��50' E, 527���539 m depth, 30 Nov 2012. Paratypes. 9 specimens: ASIZ P.73777, adult male 230 mm TL, ASIZ P.73778, female 188 mm TL, ASIZ P.73765, pregnant female 277 mm TL, off Lae, Huon Gulf, Morobe Province, Papua New Guinea, 06��51.841' S, 147��04.672' E, 395���406 m depth, 22 Aug 2010; NTUM 10313 (tissue accession GN 17195), female 269 mm TL, northern Cape King William, Morobe Province, Papua New Guinea, 06��00' S, 147��38' E, 785 m depth, 10 Dec 2012; NTUM 10314 (tissue accession GN 17197), female 258 mm TL, Astrolabe Bay, Madang, Papua New Guinea, 05��22' S, 145��48' E, 420���490 m depth, 14 Dec 2012; NTUM 10315 (tissue accession GN 17198), female 154 mm TL, Astrolabe Bay, Madang, Papua New Guinea, 05��22' S, 145��48' E, 340���385 m depth, 14 Dec 2012; NTUM 10316 (3 specimens; tissue accessions GN 17210���2), female 177 mm TL, subadult male 201 mm TL, female 228 mm TL, west of Kairiru Island, East Sepik, Papua New Guinea, 03��19' S, 143��27' E, 422���425 m depth, 19 Dec 2012. Diagnosis. Etmopterus samadiae is a relatively small, slender, species of linear���denticled Etmopterus that can be separated from its closest congeners within the E. lucifer clade by a combination of characteristics including the length of its anterior flank markings being slightly shorter than its posterior branch, long caudal base marking, and irregular and variable number of black, horizontal, dash-like marks on sides of body. The new species is morphologically and genetically (based on the NADH 2 marker) closest to E. brachyurus, but differs from this species in having a shorter posterior caudal marking (2.8���4.4 vs. 4.2���6.1% TL), a longer caudal base marking (10.6���14.1 vs. 7.0���7.8% TL), and flank marking with a slightly shorter posterior branch (9.1���11.2 vs. 11.4���12.6% TL). Description. Values expressed as a percentage of total length (TL) for the holotype, followed by the range of values for 9 paratypes (Table 1). Body fusiform, trunk sub-cylindrical (Fig. 1), width 1.1 (0.7���1.7) in trunk height; head sub-conical, long, 21.3 (20.6���23.7)% TL, slightly depressed, height 0.7 (0.5���0.8) times width. Snout moderately long, conical in lateral view, in dorsal view triangular���shaped becoming rounded at snout���tip (Fig. 3), head width 8.2 (9.8���11.8)% TL. Eyes oval-shape, large, orbit length 3.8 (3.0���3.7) in head and 2.6 (2.0���3.3) times orbit height; orbits with anterior and posterior notches; moderately spaced, inter���orbital space 1.2 (1.2���1.5) in width of head and orbit length 1.2 (1.1���1.4) times in inter���orbital distance. Spiracles small, semi-circular, greatest diameter 0.9 (1.4���2.5)% TL, 6.1 (2.7���4.0) times orbit length, distance to eye 2.9 (1.4���2.6)% TL, eye���spiracle length 0.7 (1.0���1.9) in orbit height. Nostrils large, oblique, length almost equal to internarial width, less than orbit length; anterior nasal flap well developed, triangular, anterior tip extending across nasal opening, length 1.0 (0.5���0.9) times spiracle length. Gill openings small, narrow, slightly oblique, in horizontal series, subequal in height, inter-gill length 4.9 (3.0���4.7)% TL. Mouth broad, length 3.4 (3.3���4.8) times in width, slightly arched, width 0.8 (0.7���0.9) times preoral length. Teeth dissimilar in upper and lower jaw (Fig. 4); upper teeth multicuspid in three functional series, functional teeth in lower jaw unicuspid in single series; multicuspid upper teeth small, upright, with strong central cusp flanked by 2 or 3 lateral cusplets on each side, decreasing in size distally; teeth in lower jaw fused into single row, blade-like, cusp oblique. Tooth count in first row of upper jaw 33 (27���28) and in first row of lower jaw 35 (28���31). First dorsal fin small, rounded at apex, length of first dorsal fin 8.6 (8.5���9.9)% TL, origin just anterior to pectoral-fin free rear tip; fin base insertion well anterior of pelvic-fin origin; pre���first dorsal fin length 1.3 (1.4���1.8) times inter���dorsal distance; first dorsal���fin spine straight, short, 1.6 (1.3���2.0) times height of first dorsal fin, located over pectoral fin rear margin. Second dorsal fin conspicuously larger, more erect than first dorsal fin, length of first dorsal fin 0.6 (0.6���0.8) times second dorsal fin, height of first dorsal fin 0.4 (0.5���0.7) times second dorsal fin; apex sub���angular, posterior margin concave, free rear tip elongated, length 13.2 (12.2���13.6)% TL, pre���second dorsal length 2.5 (2.6���3.1) times inter���dorsal distance; second dorsal���fin spine large, height slightly taller than fin, curved near tip towards fin apex; origin posterior to insertion of pelvic fins, over pelvic fin free rear tips. Interspace between first and second dorsal fins 1.0 (0.8���1.0) times pre���pectoral length. Pectoral fins relatively large, length 9.4 (10.1���11.3)% TL, subangular at free rear tips, base 2.0 (1.7���2.1) times in anterior margin, posterior margin nearly straight. Caudal peduncle relatively long, dorsal-caudal space 16.1 (13.6���15.3)% TL, height slightly greater than width, rounded, and tapering posteriorly. Caudal fin elongate, subequal to head length, terminal lobe distinct; length of lower preventral caudal fin margin less than one-half upper caudal fin margin. E. samadiae n.sp. E. fusus E. evansi E. brachyurus Holotype Paratypes NTUM n = 4 n = 4 Min. Max. 10318 Min. Max. Min. Max. Total length (mm) 265 154 277 256 172.0 343.0 224 350 ......continued on the next page E. samadiae n.sp. E. fusus E. evansi E. brachyurus Holotype Paratypes NTUM n = 4 n = 4 ......continued on the next page E. samadiae n.sp. E. fusus E. evansi E. brachyurus Holotype Paratypes NTUM n = 4 n = 4 Min. Max. 10318 Min. Max. Min. Max. Dermal denticles hook-like, posteriorly directed rearwards; organized in distinct rows laterally, characteristic of E. lucifer clade members (Fig. 5, 6). Distance between lateral rows mostly consistent along length, decreasing only very slightly towards caudal peduncle. Flank mark area denticles more dense and pointed ventrally. Luminescent markings distinct, intricate (Fig. 7 D); ventral head surface markings blackish, starting from almost at snout tip, extending to level of nostrils and orbits at just below level of anterior notch, then extending straight from just below posterior eye notch towards gill slits, weakly demarcated from belly marking by a weak band of transverse dermal folds across throat extending from below lower edges of first three gill openings on either side. Head dorsal surface photophore pattern as follows: a single midline along back originating at level of first gill slits extending posteriorly to caudal���fin origin; scattered photophores on paler fontanelle area of central head and also on paler area above orbits (as short dashes in some paratypes). Belly marking originates behind mouth on posterior portion of transverse dermal folds and extends ventrally along pectoral fin bases extending upwards to level with fin origin and upper edges of gill slits, and posteriorly to pelvic fin bases; ventral surface of pectoral fin very dark along lower base and upper base where ceratotrichia originate, bisected by distinct lighter to white band or blotch forming a two���prong fork pattern; margin on lateral trunk (pectoral���pelvic space) very clearly defined, line extending from rear margin of pectoral���fin base nearly to pelvic���fin insertion except for paler area at pelvic-fin origin; dark ventral belly surface continuous onto caudal peduncle to about lower caudal fin origin. Flank markings well defined (Fig. 7 d), consisting of both an anterior and posterior branch; anterior branch relatively short, length 10.3 (8.2���10.5)% TL, slender, nearly straight, extending well anterior to pelvic���fin origin; posterior branch straight, slightly thicker, width at level of base end of second dorsal fin 0.6 (0.5���0.8)% TL, slightly longer than anterior branch (shorter than anterior branch in paratype ASIZ P.73777), length 11.2 (9.1��� 10.8)% TL, extending to just anterior to second dorsal-fin free rear tip but well posterior to second dorsal-fin insertion; base of flank marking wide, origin slightly posterior to pelvic-fin insertion. Ventral caudal base marking distinct, short, length about equal to base of flank marking, not extending as a saddle on to caudal peduncle, anterior finger very short, posterior finger long, total length (including base) 11.4 (10.6���14.1)% TL. No central caudal marking. Posterior caudal fin marking very narrow, its length 3.2 (2.8���4.4) % TL. Vertebral counts: total counts 80 (80���86), total precaudal counts 55 (55���59); monospondylous 34 (36���37); diplospondylous precaudal 21 (19���22); caudal 25 (25���27). Coloration. In life, greyish to silvery black dorsally and laterally on body, becoming dark black ventrally; transition between lateral and ventral surfaces well demarcated by a paler lateral stripe below the flank markings in most specimens (less distinct in holotype and some paratypes). Dorsal midline with a broad, pale stripe originating just posterior to midpoint of inter-spiracle space, extending to first dorsal fin origin, continuing between dorsal fins, and from second dorsal fin insertion to upper caudal fin origin (Fig. 2); most prominent anterior to first dorsal fin and between dorsal fins, less prominent along upper caudal peduncle; a single row of dark photophores extending along middle of white dorsal stripe. Body with variable number and arrangement of short, horizontal, dash-like black markings (Fig. 8); most numerous in paratype NTUM 10314 (Fig. 8). Pectoral and pelvic fins dark at base and along anterior fin edge, becoming translucent to white on remainder of fins. Dorsal fins dark at base and along anterior edges, becoming translucent to white on remainder of fins. Black lateral flank markings demarcated by surrounding lighter colored lateral flanks (not sharply demarcated in most preserved specimens). Paler colored flank area on lateral surface between pectoral and pelvic fins sharply demarcates lateral and ventral surfaces; ventral surface black. Caudal fin with a distinct, large dark blotch centrally, occupying the area between the posterior finger of the caudal base marking and the upper caudal marking; posterior margin dark. Ventral surface mostly dark black around mouth, belly, and with a dark stripe between pelvic fin insertions and caudal origin; snout not distinctly paler than mouth. After preservation coloration similar but with paler markings often less obvious and coloration becoming a light or dark brown; dorsal median pale stripe and whitish flank area and black lateral flank markings less prominent but still distinct in most cases; dark blotch on central caudal fin obvious and darker pores extending laterally on body still clearly visible under microscope. Molecular analysis. The analysis of the NADH 2 data confirms that Etmopterus samadiae belongs in the E. lucifer clade and represents a monophyletic lineage that is distinct from, but most closely related to Etmopterus brachyurus (Fig. 9). It should be noted that this inference is based on a single mitochondrial marker. Inclusion of multiple nuclear markers could affect the presented inference. Size. Specimens examined ranged in size from 154 to 277 mm TL. A 277 mm TL female (ASIZ P.73765) was found to be pregnant (embryo caudal fin visible in cloaca) and two males 230 (ASIZ P.73777) and 265 mm TL (NTUM 10078) were determined to be mature. Distribution. The new species is known from off the northern Papua New Guinea mainland, from west of Kairiru Island in East Sepik Province to off Lae in the Huon Gulf (Morobe Province) and at a depth range of 340 to 785 m (Fig. 10). Etymology. The species is named after Dr Sarah Samadi (MNHN) who was one of the key principal investigators of the 2010 and 2012 MNHN expeditions in Papua New Guinea from which all the type specimens were obtained. The proposed English common name is Papuan Lanternshark., Published as part of White, William T., Ebert, David A., Mana, Ralph R. & Corrigan, Shannon, 2017, Etmopterus samadiae n. sp., a new lanternshark (Squaliformes: Etmopteridae) from Papua New Guinea, pp. 339-354 in Zootaxa 4244 (3) on pages 340-349, DOI: 10.11646/zootaxa.4244.3.3, http://zenodo.org/record/427411, {"references":["Fricke, R., Allen, G. R., Andrefouet, S., Chen, W. - J., Hamel, M. A., Laboute, P., Mana, R., Hui, T. H. & Uyeno, D. (2014) Checklist of the marine and estuarine fishes of Madang District, Papua New Guinea, western Pacific Ocean, with 820 new records. Zootaxa, 3834 (1), 1 - 247."]}

Etmopterus lailae, new species Laila���s Lanternshark (Figures 1���3 a, 4���5, Table 1) Holotype. BPBM 40183, 368 mm TL, immature male, R/V Townsend Cromwell cruise 8805, leg 2, station 216, Koko Seamount, 35�� 16.48��N 171�� 17.13��E to 35�� 16.55��N 171�� 17.20��E, 314���358 m, 13 August 1988. Paratypes. BPBM 40174, 303 mm TL, immature male, R/ V Townsend Cromwell cruise 8805, leg 2, station 167, South Kanmu Seamount, 32�� 03��N 173�� 04��E to 32�� 02��N 173�� 06��E, 336���338 m, 6 August 1988; BPBM 40182, 265 mm TL, immature male, R/V Townsend Cromwell cruise 8805, leg 2, station 218, Koko Seamount, 35�� 17.05��N 171�� 22.01��E to 35�� 17.05��N 171�� 21.54��E, 368���384 m, 14 August 1988. Diagnosis. Etmopterus lailae is a moderately large, slender species of linear���denticled Etmopterus that can be separated from most of its congeners within the E. lucifer clade by the length of its anterior flank marking branch being much longer relative to its posterior branch; all other members of this genus, except for two species, have a posterior branch that is equal to or longer than the anterior branch. The only two species with an anterior branch relatively longer than the posterior branch, E. lucifer Jordan & Snyder, 1902 and E. sculptus Ebert, Compagno, & De Vries, 2011, can be separated from the new species by a lower spiral valve count (8���9 vs 14���16 for E. lailae), a slightly higher precaudal vertebral count (55���64 vs 53���57), and a higher number of teeth on the lower jaw (30���43 vs 26���28). Etmopterus lailae lacks dermal denticles between the nostrils on the ventral snout surface and on the dorsal fins, while the other two species have denticles present on the snout and dorsal fins. Description. Proportional measurements expressed as a percentage of total length (TL) are given for the holotype and the range for two paratypes (Table 1). Body fusiform, trunk sub-cylindrical, width 1.2 (0.7���1.0) in trunk height; head sub-conical, moderate-sized, length 14.9 (13.9���16.2)% TL, slightly depressed, height 1.6 (1.3���1.6) times width. Snout moderately long, triangular���shaped becoming rounded at snout���tip in dorsal view, slightly depressed in lateral view, head width 11.1 (9.2���10.9)% TL. Eyes oval-shape, large, length 2.9 (2.6���3.3) in head, 1.9 (1.6���1.8) times width of eye; orbits with anterior and posterior notches; inter���orbital space 1.6 (1.3���1.5) in width of head; eye length 1.3 (1.4���1.5) times in inter���orbital distance. Spiracles small, greatest diameter 1.9 (1.3���1.5)% TL, 2.7 (3.3���4.0) times into length of eye, distance to eye 2.4 (2.6)% TL, eye���spiracle length 0.9 (0.9) times into height of eye. Nostrils large, oblique, length 1.4 (1.5���1.6) times into internarial width, 2.7 (2.6���2.7) times eye diameter; anterior nasal flap well developed, triangular, anterior tip extending across nasal opening, length 0.6 (0.5) times into spiracle length. Gill openings small, slightly oblique, in horizontal series, height decreasing progressively posteriorly, first two openings noticeably larger than last three openings, fifth opening just in front of pectoral fin origin; height of first gill slit 1.8 (1.3���1.8) in height of fifth gill opening; inter-gill length about equal, 1.1 (1.2���1.3) times, to length of eye. Mouth broad, length 4.0 (3.0���3.3) times width, slightly arched, width 0.7 (0.6) in preoral length. ......continued on the next page Teeth dissimilar in upper and lower jaw (Fig. 2); upper jaw teeth with strong central cusp flanked on each side by two smaller lateral cusplets, less than one-half the height of median cusp, and decreasing in size distally; lower jaw teeth unicuspid, blade-like, oblique, fused into a single row. Tooth count in first row of upper jaw 24 (22���24) and in first row of lower jaw 28 (26���26). First dorsal fin small, length of first dorsal fin 9.0 (7.9���8.6)% TL, anterior margin slightly curved, rounded at apex, origin slightly behind pectoral fin free rear tip; fin base insertion well forward of pelvic-fin origin; pre���first dorsal length 1.6 (1.7���1.8) in inter���dorsal distance; first dorsal���fin spine nearly straight, short, (0.4���0.9) times into height of first dorsal fin, located posterior to pectoral fin posterior margin. Second dorsal fin noticeably larger than first dorsal fin, length of first dorsal fin 0.7 (0.6���0.7) into second dorsal fin, height of first dorsal fin (0.5���0.6) into second dorsal fin; apex broadly rounded, posterior margin concave, free rear tip elongated, length 13.0 (12.5��� 13.2)% TL, pre���second dorsal length 2.8 (2.9���3.1) in inter���dorsal distance; second dorsal���fin spine large, height about equal to or slightly higher than fin height, slightly curved near tip towards fin apex; origin behind over or slightly behind pelvic fin free rear tips. Interspace between first and second dorsal fins 2.8 (2.9���3.1) times into pre��� pectoral length. Pectoral fin length 9.0 (8.3)% TL, broadly rounded at free rear tips, base into anterior margin length ratio 1.7 (1.7���2.0), posterior margin nearly straight edged. Caudal peduncle rounded, relatively short, 12.0 (10.6���10.9)% TL, and tapering posteriorly; height slightly greater than width, 1.3 (1.0���1.4) times width; distance less than upper caudal fin length. Caudal fin elongated, slightly greater than head length, sub���terminal notch conspicuous; preventral caudal fin margin 2.4 (3.2���3.0) into dorsal caudal fin margin. Dermal denticles on dorsal body surface erect, thorn-like, curved rearwards, in distinct longitudinal rows extending from dorsal head surface to caudal fin; distance between rows appear to decrease behind pelvic fin insertions to caudal fin. Ventral snout surface with prominent pores (ampullae of Lorenzini) surrounded by dermal denticles, except for bare patch between nostrils and extending just behind posterior nostril edges; area above upper lip of jaw without dermal denticles (Fig. 3 a). Dorsal fins mostly naked, without dermal denticles extending on fin base or ceratotrichia. Luminescent markings distinct, covering ventral head surface, extending to level of nostrils and at about orbital anterior notch, descending below eye level to mouth, then extending upward over mouth corners, but not encircling mouth; discontinuous with belly marking posterior to mouth at about first gill openings, demarcated by a faint band of transverse dermal folds across throat extending from below lower edges of first and second gill openings on either side. Belly marking originates behind mouth on posterior portion of transverse dermal folds and extends ventrally along pectoral fin bases slightly extending upwards to about level with fin base insertion, about level with ventral edges of gill openings, and posteriorly to pelvic fin bases; ventral surface of pectoral fin with two very dark lobeshaped markings, one at fin origin and another at insertion, each extending from fin base to origin of ceratotrichia bisected by distinct lighter band forming a two���prong fork pattern; margin on pectoral���pelvic space clearly demarcated, line extending from rear margin of pectoral���fin insertion nearly to pelvic���fin origin except for lighter space at pelvic���fin origin; dark ventral belly surface terminates under free rear inner margin of second dorsal fin; ventral caudal peduncle surface with darker marking just behind cloaca and extending about halfway to lower caudal origin (Fig. 4). Flank markings distinctive, with anterior and posterior branches present; anterior branch slender, curving slightly over pelvic fins with a downward concavity, thin proximally near marking base, thickening medially before narrowing to acute tip distally; length relatively long, 11.1 (9.6���10.4)% TL as measured from marking base to tip, extending beyond origin of pelvic���fin; posterior branch nearly straight, relatively thick, length less than anterior branch, 6.3 (6.3���7.9)% TL, extending just beyond second dorsal fin base insertion, terminating below and before midpoint of inner fin margin; base of flank marking relatively narrow, origin posterior to pelvic fin insertion. Caudal central marking distinct, thick, ovoid-shaped, length 6.0 (6.3���6.8)% TL, longer than base width of flank marking. Caudal fin upper marking very narrow, its length 1.2 (1.0���1.1) times central caudal marking. Vertebral counts: total vertebral counts 86 (79���82); monospondylous 40 (40); diplospondylous 17 (13���15); total precaudal 57 (53���55); caudal 29 (26���27). Spiral valve count is 16 (14���16). Coloration. After preservation dorsal and lateral surface a light to medium brown, except for prominent dark brown lateral and caudal markings, ventral surfaces also a dark brown; transition between lateral and ventral surfaces sharply demarcated. Body with 1 to 3 rows of prominent dark photophores extending from head posteriorly along flanks to about upper caudal origin; an irregular row of dark photophores also extending between pectoral and pelvic fins. Ventral surface a dark brown to blackish around mouth, belly, between pelvic fin origins and lower caudal origin; area around mouth distinctly dark brownish, sharply contrasting the lighter brown snout; area across throat slightly lighter than area anterior and posterior to this region. Gills and area just below darker than lighter brown area above. Pectoral and pelvic fin bases light brown above, darker below, and with blackish posterior and inner margins; remainder of fins becoming translucent. Dorsal fins light brown at base, becoming lighter to translucent. Lateral and caudal flank markings prominent, sharply demarcated, but without lighter colored lateral flanks. Caudal fin after preservation lacks vertical dark bar or any noticeable markings except for upper caudal marking. Size. Maximum length is at least 370 mm TL (holotype: BPBM 40183) for an immature male, both paratypes are immature males; females were not available for examination. Distribution. The new species presently is known only from the Koko and South Kanmu seamounts, Northwestern Hawaiian Islands, and at a depth range of 314���384 m (Fig. 5). Etymology. The new species is named after Laila Mostello-Wetherbee, shark enthusiast and daughter of coauthor Brad Wetherbee. The proposed common name is Laila���s Lanternshark. Comparisons. Etmopterus lailae can be assigned to the ��� E. lucifer clade��� as defined by Straube et al. (2010) with its predominant lateral flank markings displaying conspicuous anterior and posterior branches. The members of this group are also referred to as linear denticle etmopterids due to the characteristic arrangement of distinct linear rows of denticles on the dorsal head surface that also extends to the flanks, caudal peduncle and caudal base. The ��� E. lucifer clade��� can further be subdivided into three distinct subgroups based on the relative lengths of the lateral flank marking branches (Ebert et al., 2016); anterior flank marking branch longer than posterior, anterior branch shorter than posterior, and anterior and posterior branches relatively equal in length. Eleven Etmopterus species are currently recognized to fall within the ��� E. lucifer clade���, of which six species (E. alphus Ebert, Straube, Leslie & Weigmann, 2016, E. brachyurus Smith & Radcliffe, 1912, E. bullisi Bigelow & Schroeder, 1957, E. decacuspidatus Chan, 1966, E. dislineatus Last, Burgess, & S��ret, 2002, and E. molleri Whitley, 1939) have an anterior flank branch that is shorter than the posterior, and three species (E. burgessi Schaaf-DaSilva & Ebert, 2006, E. evansi Last, Burgess, & S��ret, 2002, and E. pycnolepis, Kotlyar, 1990) have anterior and posterior branch lengths that are about equal in length (Ebert et al., 2011, 2016). The remaining two species, E. lucifer and E. sculptus Ebert, Compagno, & De Vries, 2011, each has an anterior branch length that is longer than the posterior, and appear to be closest to E. lailae based on the relative branch lengths of flank markings (Ebert et al. 2011). Etmopterus lailae can be separated from its two closest congeners by a combination of external morphological and internal meristic characteristics. Externally, E. lailae can be separated from E. sculptus by a shorter posterior flank branch marking (6.3���7.9 vs 7.5���8.6)% TL; a longer central caudal marking (6.0���6.8 vs 3.4���5.0)% TL; central caudal marking length 3.3���3.6 vs 1.4���1.9 times flank base marking width; and a shorter upper caudal marking (1.1���1.2 vs 2.0���2.5) relative to the central caudal marking. The ventral snout surface of E. lailae between the nostrils is bare, but posteriorly has dermal denticles surrounding tiny pores (ampullae of Lorenzini), while E. sculptus is rather uniformly covered with dermal denticles, with small naked patches along the upper lips, and around and between the nostrils. Internally, E. lailae compared to E. sculptus has a higher spiral valve turn count (14���16 vs 8���9), a lower precaudal vertebral count (53���57 vs 60���64), and a lower tooth count on the lower jaw (26��� 28 vs 36���43). The biogeography is also informative since E. lailae is only known from the north central Pacific Ocean, while E. sculptus occurs from off Namibia to southern Mozambique in the southern hemisphere (Ebert et al., 2011). The closest congener geographically and morphologically to E. lailae is the Japanese E. lucifer, but these two species can be separated by a combination of meristic and external morphological characteristics. Etmopterus lailae can be separated meristically from Japanese E. lucifer by having a higher spiral valve turn count (14���16 vs 8���9), a lower tooth count on the lower jaw (26���28 vs 30���39), and an overlapping although slightly lower precaudal vertebral count (53���57 vs 55���63) and a lower total vertebral count (79���86 vs 85���90), although it should be noted that precaudal and total vertebral counts vary widely depending on the region and within some regions (Last & Stevens, 2009; Last & Stewart, 2015). For example, Yamakawa et al. (1986) based on 65 Japanese E. lucifer specimens reported a range of 55���62 precaudal vertebral, while we found the range for five Japanese E. lucifer to be 59���63. Although Yamakawa et al. (1986) did not report on the number of total vertebrae from their study, we found this number to range from 85���90 based on five Japanese E. lucifer specimens, while E. lailae had a range of 79���86. Etmopterus lailae can be separated from E. lucifer by the following body ratios: mouth width shorter (6.6���7.6 vs 7.4���10.1)% TL; pre-pectoral length slightly shorter (21.8���22.6 vs 22.6���25.3)% TL; pectoral pelvic space shorter (21.5���24.4 vs 24.2���33.2)% TL; anterior pectoral fin length shorter (7.9���8.6 vs 8.8���11.0)% TL; first dorsal fin length longer (7.9���9.0 vs 5.4���8.0)% TL; second dorsal fin height relatively low, its height 26.3���34.3 vs 34.5��� 41.9% TL of its overall length. The flank markings are also informative in separating these two species: E. lailae when compared to E. lucifer has a shorter anterior flank marking branch (9.8���12.6% vs 12.7���14.0)%; a lateral flank marking base width that is slightly narrower (1.8���2.0% vs 2.1���2.5%); a slightly narrower flank base width 9.2���9.3 vs 9.6���13.9; a longer central caudal marking (6.0���6.8% vs 2.8���5.1%); central caudal marking length 3.3��� 3.6 vs 2.5���2.7 times flank base marking width. The arrangement of the dermal denticles, including their presence or absence, is informative when separating E. lailae from E. lucifer. The ventral snout surface of E. lailae has a naked patch between, and extending just posterior to, the space between the inner nostrils (Fig. 2 a). Also, beginning just posterior to the inner nostril opening is a bare patch, absent dermal denticles, extending rearwards to the upper lip except for small patch of denticles bisecting it about one-third the distance from the inner nostril opening to the upper lip; this area has several large pore openings. The area just anterior to the upper lip of the jaw, and traversing its length, also has a bare area with a few large scattered pores. Distinct pores cover the remainder of the snout ventral surface. In contrast, the snout ventral surface of E. lucifer is densely and relatively evenly covered by dermal denticles, but has no bare patches, except along the upper lip (Fig. 2 b); some small pores are apparent, but scattered. The dorsal fins of E. lailae are mostly naked, without dermal denticles extending onto the dorsal fins, while E. lucifer has linear rows of denticles present on the bases and extending onto the dorsal fins, including the ceratotrichia. The lateral dermal denticles extending the length of the body on E. lailae appear more sculpted than observed in E. lucifer. In addition to the above parameters these two species differ significantly in the size at maturity. All three known specimens of E. lailae are immature males, with the largest measuring 368 mm TL after preservation (370 mm TL before preservation), while E. lucifer from Japanese waters is a much smaller species with males maturing (before preservation) at a minimum length of 278 mm TL, with a maximum of 310 mm TL, based on the individuals examined in this study. The separation of ��� E. lucifer clade��� etmopterids, including the new species E. lailae, is complicated due to the inadequate original description of E. lucifer from Japan and the convoluted taxonomic history of this species from Japanese waters and elsewhere. The original description and type series of E. lucifer was based on a combination of two different etmopterid species, one form with an anterior branch longer than the posterior, and the other with the posterior branch being longer than the anterior. Examination of several of the syntypes by DAE and J.A. Schaaf-Da Silva (California Department Fish & Wildlife) confirms that two different species are among the type series. Furthermore, a lectotype has never been designated for the species, thus leaving open the question of what species exactly constitutes true E. lucifer, and leaves unresolved the status of the other species comprising the type series. Resolution to, and a description of, true E. lucifer with a lectotype designation are currently under investigation by DAE and N. Straube (Bavarian State Collection of Zoology)., Published as part of Ebert, David A., Papastamatiou, Yannis P., Kajiura, Stephen M. & Wetherbee, Bradley M., 2017, Etmopterus lailae sp. nov., a new lanternshark (Squaliformes: Etmopteridae) from the Northwestern Hawaiian Islands, pp. 371-382 in Zootaxa 4237 (2) on pages 372-379, DOI: 10.11646/zootaxa.4237.2.10, http://zenodo.org/record/343864, {"references":["Straube N., Iglesias, S. P., Sellos, D. Y., Kriwet, J. & Schliewen, U. K. (2010) Molecular phylogeny and node time estimation of bioluminescent lanternsharks (Elasmobranchii: Etmopteridae). Molecular Phylogenetics and Evolution, 56, 905 - 917. https: // doi. org / 10.1016 / j. ympev. 2010.04.042","Ebert, D. A., Straube, N., Leslie, R. W. & Weigmann, S. (2016) Etmopterus alphus n. sp.: a new lanternshark (Squaliformes: Etmopteridae) from the south-western Indian Ocean. African Journal of Marine Science, 38, 329 - 340. https: // doi. org / 10.2989 / 1814232 X. 2016.1198275","Schaaf-DaSilva, J. A. & Ebert, D. A. (2006) Etmopterus burgessi sp. nov., a new species of lanternshark (Squaliformes: Etmopteridae) from Taiwan. Zootaxa, 1373, 53 - 64.","Kotlyar, AN. (1990) Dogfish sharks of the genus Etmopterus Rafinesque from the Nazca and Sala y Gomez Submarine Ridges. Trudy Instituta Okeanologii Imeni P. P. Shirshova, 125, 127 - 147.","Ebert, D. A., Compagno, L. J. V. & DeVries, M. J. (2011) A new lantershark (Squaliformes: Etmopteridae: Etmopterus) from southern Africa. Copeia, 2011, 379 - 384.","Last, P. R. & Stevens, J. D. (2009) Sharks and rays of Australia. CSIRO Publishing, Melbourne, 644 pp.","Last, P. R. & Stewart, A. L. (2015) Family Etmopteridae. In: Roberts, C. D., Stewart, A. L., Struthers, C. D. (Eds.), Fishes of New Zealand. Lol. 2. Te Papa Press, Wellington, pp. 139 - 147."]}

Genus Etmopterus Rafinesque, 1810 Etmopterus Rafinesque, 1810: 14. Type species: Etmopterus aculeatus Rafinesque, 1810 by monotypy., Published as part of Ebert, David A., Papastamatiou, Yannis P., Kajiura, Stephen M. & Wetherbee, Bradley M., 2017, Etmopterus lailae sp. nov., a new lanternshark (Squaliformes: Etmopteridae) from the Northwestern Hawaiian Islands, pp. 371-382 in Zootaxa 4237 (2) on page 372, DOI: 10.11646/zootaxa.4237.2.10, http://zenodo.org/record/343864

Etmopterus lailae, new species Laila’s Lanternshark (Figures 1–3 a, 4–5, Table 1) Holotype. BPBM 40183, 368 mm TL, immature male, R/V Townsend Cromwell cruise 8805, leg 2, station 216, Koko Seamount, 35° 16.48´N 171° 17.13´E to 35° 16.55´N 171° 17.20´E, 314–358 m, 13 August 1988. Paratypes. BPBM 40174, 303 mm TL, immature male, R/ V Townsend Cromwell cruise 8805, leg 2, station 167, South Kanmu Seamount, 32° 03´N 173° 04´E to 32° 02´N 173° 06´E, 336–338 m, 6 August 1988; BPBM 40182, 265 mm TL, immature male, R/V Townsend Cromwell cruise 8805, leg 2, station 218, Koko Seamount, 35° 17.05´N 171° 22.01´E to 35° 17.05´N 171° 21.54´E, 368–384 m, 14 August 1988. Diagnosis. Etmopterus lailae is a moderately large, slender species of linear–denticled Etmopterus that can be separated from most of its congeners within the E. lucifer clade by the length of its anterior flank marking branch being much longer relative to its posterior branch; all other members of this genus, except for two species, have a posterior branch that is equal to or longer than the anterior branch. The only two species with an anterior branch relatively longer than the posterior branch, E. lucifer Jordan & Snyder, 1902 and E. sculptus Ebert, Compagno, & De Vries, 2011, can be separated from the new species by a lower spiral valve count (8–9 vs 14–16 for E. lailae), a slightly higher precaudal vertebral count (55–64 vs 53–57), and a higher number of teeth on the lower jaw (30–43 vs 26–28). Etmopterus lailae lacks dermal denticles between the nostrils on the ventral snout surface and on the dorsal fins, while the other two species have denticles present on the snout and dorsal fins. Description. Proportional measurements expressed as a percentage of total length (TL) are given for the holotype and the range for two paratypes (Table 1). Body fusiform, trunk sub-cylindrical, width 1.2 (0.7–1.0) in trunk height; head sub-conical, moderate-sized, length 14.9 (13.9–16.2)% TL, slightly depressed, height 1.6 (1.3–1.6) times width. Snout moderately long, triangular–shaped becoming rounded at snout–tip in dorsal view, slightly depressed in lateral view, head width 11.1 (9.2–10.9)% TL. Eyes oval-shape, large, length 2.9 (2.6–3.3) in head, 1.9 (1.6–1.8) times width of eye; orbits with anterior and posterior notches; inter–orbital space 1.6 (1.3–1.5) in width of head; eye length 1.3 (1.4–1.5) times in inter–orbital distance. Spiracles small, greatest diameter 1.9 (1.3–1.5)% TL, 2.7 (3.3–4.0) times into length of eye, distance to eye 2.4 (2.6)% TL, eye–spiracle length 0.9 (0.9) times into height of eye. Nostrils large, oblique, length 1.4 (1.5–1.6) times into internarial width, 2.7 (2.6–2.7) times eye diameter; anterior nasal flap well developed, triangular, anterior tip extending across nasal opening, length 0.6 (0.5) times into spiracle length. Gill openings small, slightly oblique, in horizontal series, height decreasing progressively posteriorly, first two openings noticeably larger than last three openings, fifth opening just in front of pectoral fin origin; height of first gill slit 1.8 (1.3–1.8) in height of fifth gill opening; inter-gill length about equal, 1.1 (1.2–1.3) times, to length of eye. Mouth broad, length 4.0 (3.0–3.3) times width, slightly arched, width 0.7 (0.6) in preoral length. ......continued on the next page Teeth dissimilar in upper and lower jaw (Fig. 2); upper jaw teeth with strong central cusp flanked on each side by two smaller lateral cusplets, less than one-half the height of median cusp, and decreasing in size distally; lower jaw teeth unicuspid, blade-like, oblique, fused into a single row. Tooth count in first row of upper jaw 24 (22–24) and in first row of lower jaw 28 (26–26). First dorsal fin small, length of first dorsal fin 9.0 (7.9–8.6)% TL, anterior margin slightly curved, rounded at apex, origin slightly behind pectoral fin free rear tip; fin base insertion well forward of pelvic-fin origin; pre–first dorsal length 1.6 (1.7–1.8) in inter–dorsal distance; first dorsal–fin spine nearly straight, short, (0.4–0.9) times into height of first dorsal fin, located posterior to pectoral fin posterior margin. Second dorsal fin noticeably larger than first dorsal fin, length of first dorsal fin 0.7 (0.6–0.7) into second dorsal fin, height of first dorsal fin (0.5–0.6) into second dorsal fin; apex broadly rounded, posterior margin concave, free rear tip elongated, length 13.0 (12.5– 13.2)% TL, pre–second dorsal length 2.8 (2.9–3.1) in inter–dorsal distance; second dorsal–fin spine large, height about equal to or slightly higher than fin height, slightly curved near tip towards fin apex; origin behind over or slightly behind pelvic fin free rear tips. Interspace between first and second dorsal fins 2.8 (2.9–3.1) times into pre– pectoral length. Pectoral fin length 9.0 (8.3)% TL, broadly rounded at free rear tips, base into anterior margin length ratio 1.7 (1.7–2.0), posterior margin nearly straight edged. Caudal peduncle rounded, relatively short, 12.0 (10.6–10.9)% TL, and tapering posteriorly; height slightly greater than width, 1.3 (1.0–1.4) times width; distance less than upper caudal fin length. Caudal fin elongated, slightly greater than head length, sub–terminal notch conspicuous; preventral caudal fin margin 2.4 (3.2–3.0) into dorsal caudal fin margin. Dermal denticles on dorsal body surface erect, thorn-like, curved rearwards, in distinct longitudinal rows extending from dorsal head surface to caudal fin; distance between rows appear to decrease behind pelvic fin insertions to caudal fin. Ventral snout surface with prominent pores (ampullae of Lorenzini) surrounded by dermal denticles, except for bare patch between nostrils and extending just behind posterior nostril edges; area above upper lip of jaw without dermal denticles (Fig. 3 a). Dorsal fins mostly naked, without dermal denticles extending on fin base or ceratotrichia. Luminescent markings distinct, covering ventral head surface, extending to level of nostrils and at about orbital anterior notch, descending below eye level to mouth, then extending upward over mouth corners, but not encircling mouth; discontinuous with belly marking posterior to mouth at about first gill openings, demarcated by a faint band of transverse dermal folds across throat extending from below lower edges of first and second gill openings on either side. Belly marking originates behind mouth on posterior portion of transverse dermal folds and extends ventrally along pectoral fin bases slightly extending upwards to about level with fin base insertion, about level with ventral edges of gill openings, and posteriorly to pelvic fin bases; ventral surface of pectoral fin with two very dark lobeshaped markings, one at fin origin and another at insertion, each extending from fin base to origin of ceratotrichia bisected by distinct lighter band forming a two–prong fork pattern; margin on pectoral–pelvic space clearly demarcated, line extending from rear margin of pectoral–fin insertion nearly to pelvic–fin origin except for lighter space at pelvic–fin origin; dark ventral belly surface terminates under free rear inner margin of second dorsal fin; ventral caudal peduncle surface with darker marking just behind cloaca and extending about halfway to lower caudal origin (Fig. 4). Flank markings distinctive, with anterior and posterior branches present; anterior branch slender, curving slightly over pelvic fins with a downward concavity, thin proximally near marking base, thickening medially before narrowing to acute tip distally; length relatively long, 11.1 (9.6–10.4)% TL as measured from marking base to tip, extending beyond origin of pelvic–fin; posterior branch nearly straight, relatively thick, length less than anterior branch, 6.3 (6.3–7.9)% TL, extending just beyond second dorsal fin base insertion, terminating below and before midpoint of inner fin margin; base of flank marking relatively narrow, origin posterior to pelvic fin insertion. Caudal central marking distinct, thick, ovoid-shaped, length 6.0 (6.3–6.8)% TL, longer than base width of flank marking. Caudal fin upper marking very narrow, its length 1.2 (1.0–1.1) times central caudal marking. Vertebral counts: total vertebral counts 86 (79–82); monospondylous 40 (40); diplospondylous 17 (13–15); total precaudal 57 (53–55); caudal 29 (26–27). Spiral valve count is 16 (14–16). Coloration. After preservation dorsal and lateral surface a light to medium brown, except for prominent dark brown lateral and caudal markings, ventral surfaces also a dark brown; transition between lateral and ventral surfaces sharply demarcated. Body with 1 to 3 rows of prominent dark photophores extending from head posteriorly along flanks to about upper caudal origin; an irregular row of dark photophores also extending between pectoral and pelvic fins. Ventral surface a dark brown to blackish around mouth, belly, between pelvic fin origins and lower caudal origin; area around mouth distinctly dark brownish, sharply contrasting the lighter brown snout; area across throat slightly lighter than area anterior and posterior to this region. Gills and area just below darker than lighter brown area above. Pectoral and pelvic fin bases light brown above, darker below, and with blackish posterior and inner margins; remainder of fins becoming translucent. Dorsal fins light brown at base, becoming lighter to translucent. Lateral and caudal flank markings prominent, sharply demarcated, but without lighter colored lateral flanks. Caudal fin after preservation lacks vertical dark bar or any noticeable markings except for upper caudal marking. Size. Maximum length is at least 370 mm TL (holotype: BPBM 40183) for an immature male, both paratypes are immature males; females were not available for examination. Distribution. The new species presently is known only from the Koko and South Kanmu seamounts, Northwestern Hawaiian Islands, and at a depth range of 314–384 m (Fig. 5). Etymology. The new species is named after Laila Mostello-Wetherbee, shark enthusiast and daughter of coauthor Brad Wetherbee. The proposed common name is Laila’s Lanternshark. Comparisons. Etmopterus lailae can be assigned to the “ E. lucifer clade” as defined by Straube et al. (2010) with its predominant lateral flank markings displaying conspicuous anterior and posterior branches. The members of this group are also referred to as linear denticle etmopterids due to the characteristic arrangement of distinct linear rows of denticles on the dorsal head surface that also extends to the flanks, caudal peduncle and caudal base. The “ E. lucifer clade” can further be subdivided into three distinct subgroups based on the relative lengths of the lateral flank marking branches (Ebert et al., 2016); anterior flank marking branch longer than posterior, anterior branch shorter than posterior, and anterior and posterior branches relatively equal in length. Eleven Etmopterus species are currently recognized to fall within the “ E. lucifer clade”, of which six species (E. alphus Ebert, Straube, Leslie & Weigmann, 2016, E. brachyurus Smith & Radcliffe, 1912, E. bullisi Bigelow & Schroeder, 1957, E. decacuspidatus Chan, 1966, E. dislineatus Last, Burgess, & Séret, 2002, and E. molleri Whitley, 1939) have an anterior flank branch that is shorter than the posterior, and three species (E. burgessi Schaaf-DaSilva & Ebert, 2006, E. evansi Last, Burgess, & Séret, 2002, and E. pycnolepis, Kotlyar, 1990) have anterior and posterior branch lengths that are about equal in length (Ebert et al., 2011, 2016). The remaining two species, E. lucifer and E. sculptus Ebert, Compagno, & De Vries, 2011, each has an anterior branch length that is longer than the posterior, and appear to be closest to E. lailae based on the relative branch lengths of flank markings (Ebert et al. 2011). Etmopterus lailae can be separated from its two closest congeners by a combination of external morphological and internal meristic characteristics. Externally, E. lailae can be separated from E. sculptus by a shorter posterior flank branch marking (6.3–7.9 vs 7.5–8.6)% TL; a longer central caudal marking (6.0–6.8 vs 3.4–5.0)% TL; central caudal marking length 3.3–3.6 vs 1.4–1.9 times flank base marking width; and a shorter upper caudal marking (1.1–1.2 vs 2.0–2.5) relative to the central caudal marking. The ventral snout surface of E. lailae between the nostrils is bare, but posteriorly has dermal denticles surrounding tiny pores (ampullae of Lorenzini), while E. sculptus is rather uniformly covered with dermal denticles, with small naked patches along the upper lips, and around and between the nostrils. Internally, E. lailae compared to E. sculptus has a higher spiral valve turn count (14–16 vs 8–9), a lower precaudal vertebral count (53–57 vs 60–64), and a lower tooth count on the lower jaw (26– 28 vs 36–43). The biogeography is also informative since E. lailae is only known from the north central Pacific Ocean, while E. sculptus occurs from off Namibia to southern Mozambique in the southern hemisphere (Ebert et al., 2011). The closest congener geographically and morphologically to E. lailae is the Japanese E. lucifer, but these two species can be separated by a combination of meristic and external morphological characteristics. Etmopterus lailae can be separated meristically from Japanese E. lucifer by having a higher spiral valve turn count (14–16 vs 8–9), a lower tooth count on the lower jaw (26–28 vs 30–39), and an overlapping although slightly lower precaudal vertebral count (53–57 vs 55–63) and a lower total vertebral count (79–86 vs 85–90), although it should be noted that precaudal and total vertebral counts vary widely depending on the region and within some regions (Last & Stevens, 2009; Last & Stewart, 2015). For example, Yamakawa et al. (1986) based on 65 Japanese E. lucifer specimens reported a range of 55–62 precaudal vertebral, while we found the range for five Japanese E. lucifer to be 59–63. Although Yamakawa et al. (1986) did not report on the number of total vertebrae from their study, we found this number to range from 85–90 based on five Japanese E. lucifer specimens, while E. lailae had a range of 79–86. Etmopterus lailae can be separated from E. lucifer by the following body ratios: mouth width shorter (6.6–7.6 vs 7.4–10.1)% TL; pre-pectoral length slightly shorter (21.8–22.6 vs 22.6–25.3)% TL; pectoral pelvic space shorter (21.5–24.4 vs 24.2–33.2)% TL; anterior pectoral fin length shorter (7.9–8.6 vs 8.8–11.0)% TL; first dorsal fin length longer (7.9–9.0 vs 5.4–8.0)% TL; second dorsal fin height relatively low, its height 26.3–34.3 vs 34.5– 41.9% TL of its overall length. The flank markings are also informative in separating these two species: E. lailae when compared to E. lucifer has a shorter anterior flank marking branch (9.8–12.6% vs 12.7–14.0)%; a lateral flank marking base width that is slightly narrower (1.8–2.0% vs 2.1–2.5%); a slightly narrower flank base width 9.2–9.3 vs 9.6–13.9; a longer central caudal marking (6.0–6.8% vs 2.8–5.1%); central caudal marking length 3.3– 3.6 vs 2.5–2.7 times flank base marking width. The arrangement of the dermal denticles, including their presence or absence, is informative when separating E. lailae from E. lucifer. The ventral snout surface of E. lailae has a naked patch between, and extending just posterior to, the space between the inner nostrils (Fig. 2 a). Also, beginning just posterior to the inner nostril opening is a bare patch, absent dermal denticles, extending rearwards to the upper lip except for small patch of denticles bisecting it about one-third the distance from the inner nostril opening to the upper lip; this area has several large pore openings. The area just anterior to the upper lip of the jaw, and traversing its length, also has a bare area with a few large scattered pores. Distinct pores cover the remainder of the snout ventral surface. In contrast, the snout ventral surface of E. lucifer is densely and relatively evenly covered by dermal denticles, but has no bare patches, except along the upper lip (Fig. 2 b); some small pores are apparent, but scattered. The dorsal fins of E. lailae are mostly naked, without dermal denticles extending onto the dorsal fins, while E. lucifer has linear rows of denticles present on the bases and extending onto the dorsal fins, including the ceratotrichia. The lateral dermal denticles extending the length of the body on E. lailae appear more sculpted than observed in E. lucifer. In addition to the above parameters these two species differ significantly in the size at maturity. All three known specimens of E. lailae are immature males, with the largest measuring 368 mm TL after preservation (370 mm TL before preservation), while E. lucifer from Japanese waters is a much smaller species with males maturing (before preservation) at a minimum length of 278 mm TL, with a maximum of 310 mm TL, based on the individuals examined in this study. The separation of “ E. lucifer clade” etmopterids, including the new species E. lailae, is complicated due to the inadequate original description of E. lucifer from Japan and the convoluted taxonomic history of this species from Japanese waters and elsewhere. The original description and type series of E. lucifer was based on a combination of two different etmopterid species, one form with an anterior branch longer than the posterior, and the other with the posterior branch being longer than the anterior. Examination of several of the syntypes by DAE and J.A. Schaaf-Da Silva (California Department Fish & Wildlife) confirms that two different species are among the type series. Furthermore, a lectotype has never been designated for the species, thus leaving open the question of what species exactly constitutes true E. lucifer, and leaves unresolved the status of the other species comprising the type series. Resolution to, and a description of, true E. lucifer with a lectotype designation are currently under investigation by DAE and N. Straube (Bavarian State Collection of Zoology).

A new species of lanternshark, Etmopterus samadiae (Squaliformes: Etmopteridae), is described from off northern Papua New Guinea, in the western Central Pacific Ocean. The new species resembles other members of the “Etmopterus lucifer” clade in having linear rows of dermal denticles and most closely resembles E. brachyurus from the western North Pacific. The new species occurs along insular slopes between 340 and 785 m depth. The new species can be distinguished from other members of the E. lucifer clade by a combination of characteristics, including length of anterior flank branch markings being slightly shorter than its posterior branch, a longer caudal base marking, and irregular and variable number of black, horizontal, dash-like marks on sides of body. Molecular analysis based on the NADH2 marker further supports the distinction of E. samadiae from other members of the E. lucifer clade.

Three species of neonate and juvenile sharks (Heptranchias perlo, Squatina aculeata, Etmopterus spinax) were captured as by-catch from a commercial trawl fishing from depths between 360 and 430 m in the North-eastern Mediterranean Sea. Two neonates of H. perlo, five neonates of E. spinax and one juvenile of S. aculeata were for the first time identified in the above region.

Etmopterus spinax (Linnaeus, 1758) Squalus spinax Linnaeus, 1758: 233. Type locality: Genoa, Italy. Etmopterus spinax, Gilat & Gelman, 1984: 257; Pisanty & Golani, 1995: 389; Galil & Goren, 1994: 48; Jones et al., 2003: 78, Tab. 2. Distribution. Eastern Atlantic and Mediterranean; depth range 200���2490m., Published as part of Goren, Menachem & Galil, Bella S., 2015, A checklist of the deep sea fishes of the Levant Sea, Mediterranean Sea, pp. 507-530 in Zootaxa 3994 (4) on page 511, DOI: 10.11646/zootaxa.3994.4.2, http://zenodo.org/record/244394, {"references":["Gilat, E. & Gelman, A. (1984) On the sharks and fishes observed using underwater photography during a deep-water cruise in the Eastern Mediterranean. Fisheries Research, 2, 257 - 271. http: // dx. doi. org / 10.1016 / 0165 - 7836 (84) 90029 - 8","Pisanty, S. & Golani, D. (1995) Vertical distribution of demersal fish on the continental slope of Israel (Eastern Mediterranean) In: Armentraut, N. B. (Ed.), Condition of the World's Aquatic Habitat. Proceedings of the World Fisheries Congress. Theme 1. Oxford & IBH Publishing Company, New Delhi, pp. 386 - 395.","Galil, B. S. & Goren, M. (1994) The deep sea Levantine Fauna. New records and rare occurrences. Senckenbergiana Maritima, 25, 41 - 52.","Jones, E., Tselepides, A., Bagley, P., Collins, M. & Priede, I. (2003) Bathymetric distribution of some benthic and benthopelagic species attracted to baited cameras and traps in the deep eastern Mediterranean. Marine Ecology Progress Series, 251, 75 - 86. http: // dx. doi. org / 10.3354 / meps 251075"]}

A new species of lanternshark, Etmopterus lailae (Squaliformes: Etmopteridae), is described from the Northwestern Hawaiian Islands, in the central North Pacific Ocean. The new species resembles other members of the “Etmopterus lucifer” clade in having linear rows of dermal denticles, and most closely resembles E. lucifer from Japan. The new species occurs along insular slopes around seamounts at depths between 314–384 m. It can be distinguished from other members of the E. lucifer clade by a combination of characteristics, including a longer anterior flank marking branch, arrangement of dermal denticles on the ventral snout surface and body, flank and caudal markings, and meristic counts including number of spiral valve turns, and precaudal vertebrate. A key to species of the Etmopterus lucifer-clade is included.

The mesopelagic zone is a visual scene continuum in which organisms have developed various strategies to optimize photon capture. Here, we used light microscopy, stereology-assisted retinal topographic mapping, spectrophotometry and microspectrophotometry to investigate the visual ecology of deep-sea bioluminescent sharks [four etmopterid species (Etmopterus lucifer, E. splendidus, E. spinax and Trigonognathus kabeyai) and one dalatiid species (Squaliolus aliae)]. We highlighted a novel structure, a translucent area present in the upper eye orbit of Etmopteridae, which might be part of a reference system for counterillumination adjustment or acts as a spectral filter for camouflage breaking, as well as several ocular specialisations such as aphakic gaps and semicircular tapeta previously unknown in elasmobranchs. All species showed pure rod hexagonal mosaics with a high topographic diversity. Retinal specialisations, formed by shallow cell density gradients, may aid in prey detection and reflect lifestyle differences; pelagic species display areae centrales while benthopelagic and benthic species display wide and narrow horizontal streaks, respectively. One species (E. lucifer) displays two areae within its horizontal streak that likely allows detection of conspecifics' elongated bioluminescent flank markings. Ganglion cell topography reveals less variation with all species showing a temporal area for acute frontal binocular vision. This area is dorsally extended in T. kabeyai, allowing this species to adjust the strike of its peculiar jaws in the ventro-frontal visual field. Etmopterus lucifer showed an additional nasal area matching a high rod density area. Peak spectral sensitivities of the rod visual pigments (λmax) fall within the range 484–491 nm, allowing these sharks to detect a high proportion of photons present in their habitat. Comparisons with previously published data reveal ocular differences between bioluminescent and non-bioluminescent deep-sea sharks. In particular, bioluminescent sharks possess higher rod densities, which might provide them with improved temporal resolution particularly useful for bioluminescent communication during social interactions.

* □ Etmopterus spinax (Linnaeus, 1758) – Velvet belly; Lixinha-da-fundura ①②③, Quelmazinha ②, Lixinha ③, Published as part of Carneiro, Miguel, Martins, Rogélia, Landi, Monica & Costa, Filipe O., 2014, Updated checklist of marine fishes (Chordata: Craniata) from Portugal and the proposed extension of the Portuguese continental shelf, pp. 1-73 in European Journal of Taxonomy 73 on page 14, DOI: 10.5852/ejt.2014.73, http://zenodo.org/record/3866515

□ Etmopterus princeps Collet, 1904 – Great lanternshark; ①, Lixinha-da-fundura-grada ②, ③, Published as part of Carneiro, Miguel, Martins, Rogélia, Landi, Monica & Costa, Filipe O., 2014, Updated checklist of marine fishes (Chordata: Craniata) from Portugal and the proposed extension of the Portuguese continental shelf, pp. 1-73 in European Journal of Taxonomy 73 on page 13, DOI: 10.5852/ejt.2014.73, http://zenodo.org/record/3866515

Etmopterus brachyurus Smith & Radcliffe, 1912 Type locality: off Jolo Island, Philippines, 6 ��02'00" N, 120 �� 44 ' 40 " E. The two Taiwanese samples of E. brachyurus formed a single cluster together with a sample from Japan (GN 7423), and were sister to the E. cf. molleri clade from Taiwan (Figure 2, Supplementary Material 1)., Published as part of Straube, Nicolas, White, William T., Ho, Hsuan-Ching, Rochel, Elisabeth, Corrigan, Shannon, Li, Chenhong & Naylor, Gavin J. P., 2013, A DNA sequence-based identification checklist for Taiwanese chondrichthyans, pp. 256-278 in Zootaxa 3752 (1) on page 260, DOI: 10.11646/zootaxa.3752.1.16, http://zenodo.org/record/285357

Etmopterus splendidus Yano, 1988 Type locality: off Shimo-Koshiki Island, 31 ��46.0' N, 129 �� 43.8 ' E, Japan. The six Taiwanese samples of E. splendidus formed a single cluster together with an additional sample (GN 995) also from Taiwan (Figure 2, Supplementary Material 1). This cluster is sister to a clade comprising one sample of E. sentosus (GN 7402) from the Indian Ocean off Mozambique., Published as part of Straube, Nicolas, White, William T., Ho, Hsuan-Ching, Rochel, Elisabeth, Corrigan, Shannon, Li, Chenhong & Naylor, Gavin J. P., 2013, A DNA sequence-based identification checklist for Taiwanese chondrichthyans, pp. 256-278 in Zootaxa 3752 (1) on page 261, DOI: 10.11646/zootaxa.3752.1.16, http://zenodo.org/record/285357

Etmopterus joungi Knuckey, Ebert & Burgess, 2011 Type locality: Da-xi, Taiwan. The three Taiwanese samples of E. joungi formed a single cluster together with E. pusillus samples from Australia and Portugal. Specimens of E. pusillus included in Naylor et al. (2012 a, 2012 b) indicate haplotypes that are geographically widespread across the world���s oceans. A revision of this species is required to ascertain whether E. joungi is a western North Pacific sub-population of E. pusillus, and thus a junior synonym of E. pusillus, or whether it is a distinct species (Figure 2, Supplementary Material 1)., Published as part of Straube, Nicolas, White, William T., Ho, Hsuan-Ching, Rochel, Elisabeth, Corrigan, Shannon, Li, Chenhong & Naylor, Gavin J. P., 2013, A DNA sequence-based identification checklist for Taiwanese chondrichthyans, pp. 256-278 in Zootaxa 3752 (1) on page 261, DOI: 10.11646/zootaxa.3752.1.16, http://zenodo.org/record/285357

Etmopterus burgessi Schaaf da Silva & Ebert, 2006 Type locality: Da-xi fish market, northeastern Taiwan. The six Taiwanese samples of E. burgessi included in this analysis formed a single cluster, distinct from its congeners and sister to a clade comprising E. lucifer samples from New Zealand (Figure 2)., Published as part of Straube, Nicolas, White, William T., Ho, Hsuan-Ching, Rochel, Elisabeth, Corrigan, Shannon, Li, Chenhong & Naylor, Gavin J. P., 2013, A DNA sequence-based identification checklist for Taiwanese chondrichthyans, pp. 256-278 in Zootaxa 3752 (1) on page 261, DOI: 10.11646/zootaxa.3752.1.16, http://zenodo.org/record/285357

Etmopterus sheikoi (Dolganov, 1986) Type locality: Kyushu Palau Ridge, western North Pacific. The two Taiwanese E. sheikoi samples included in this analysis formed a single cluster that included a sample (GN 7440) from Taiwan. As discussed in Straube et al. (2010), the species is inferred to be the sister group to the E. lucifer clade. Potential misidentification can be ruled out as the morphology is highly distinctive and specimens analysed herein are vouchered. The phylogenetic affinities of E. sheikoi are especially interesting. Straube et al. (2010) showed that flank markings in Etmopterus mirror the relationships inferred from molecular data for all species, except E. sheikoi. From a morphological perspective, the species would be expected to cluster within the E. pusillus clade rather than the E. lucifer clade as suggested by the molecular data (Figure 2). This is an unusual case in which morphology and genetics are inconsistent and need further data collection and analysis from both approaches. This species was previously allocated to the genus Miroscyllium, but molecular data suggest this species belongs to the genus Etmopterus., Published as part of Straube, Nicolas, White, William T., Ho, Hsuan-Ching, Rochel, Elisabeth, Corrigan, Shannon, Li, Chenhong & Naylor, Gavin J. P., 2013, A DNA sequence-based identification checklist for Taiwanese chondrichthyans, pp. 256-278 in Zootaxa 3752 (1) on page 261, DOI: 10.11646/zootaxa.3752.1.16, http://zenodo.org/record/285357

Etmopterus joungi Knuckey, Ebert & Burgess, 2011 Type locality: Da-xi, Taiwan. The three Taiwanese samples of E. joungi formed a single cluster together with E. pusillus samples from Australia and Portugal. Specimens of E. pusillus included in Naylor et al. (2012 a, 2012 b) indicate haplotypes that are geographically widespread across the world’s oceans. A revision of this species is required to ascertain whether E. joungi is a western North Pacific sub-population of E. pusillus, and thus a junior synonym of E. pusillus, or whether it is a distinct species (Figure 2, Supplementary Material 1).

Etmopterus cf. molleri The 12 Taiwanese samples of E. cf. molleri included in this analysis formed a single cluster together with a Taiwan sample of E. molleri (GN 997), which was sister to an E. brachyurus clade (Figure 2, Supplementary Material 1). Naylor et al. (2012 a) commented that E. molleri samples from Taiwan need to be compared with the recently described E. burgessi to ensure they are not conspecific. This analysis includes the first sequenced samples of E. burgessi and shows they are quite distinct from this species. Samples of E. molleri from the type location of Australia are required to ascertain whether the Taiwan species represents a distinct species or whether they are conspecific., Published as part of Straube, Nicolas, White, William T., Ho, Hsuan-Ching, Rochel, Elisabeth, Corrigan, Shannon, Li, Chenhong & Naylor, Gavin J. P., 2013, A DNA sequence-based identification checklist for Taiwanese chondrichthyans, pp. 256-278 in Zootaxa 3752 (1) on page 261, DOI: 10.11646/zootaxa.3752.1.16, http://zenodo.org/record/285357

Etmopterus brachyurus Smith & Radcliffe, 1912 Type locality: off Jolo Island, Philippines, 6 °02'00" N, 120 ° 44 ' 40 " E. The two Taiwanese samples of E. brachyurus formed a single cluster together with a sample from Japan (GN 7423), and were sister to the E. cf. molleri clade from Taiwan (Figure 2, Supplementary Material 1).

The structure and composition of megabenthic epifauna from the mud volcano field known as El Laberinto and the nearby soft bottoms were analysed. Species abundance and biomass data were obtained from the Arrastre Region Sur-Atlantica bottom trawl surveys carried out routinely from 1993 to 2009. A high biodiversity of megabenthic epifauna was found with a total of 272 species. Univariate community descriptors displayed similar values in zones inside and outside El Laberinto and in different seasons with the exception of abundance, which was larger in the autumn, especially in the zone nearest the mud volcano field. The dominant species in terms of abundance in all zones were similar and included Pasiphaea sivado, Plesionika martia, Parapenaeus longirostris, Nezumia aequalis, Chimaera monstrosa, Galeus melastomus and Etmopterus spinax. This information could be useful for increasing knowledge on biodiversity and the occurrence of commercial species and for establishing conservation strategies for protection...

Etmopterus spinax (Linnaeus, 1758) Aegean Sea: 1200-27 (1 spc.), 14.07.1981, Gôkova Bay, trawl, 600 m, Nezih Bilecik.

Etmopterus spinax (Linnaeus, 1758) Aegean Sea: 1200-27 (1 spc.), 14.07.1981, G��kova Bay, trawl, 600 m, Nezih Bilecik., Published as part of Nurettin Meri��, L��tfiye Eryilmaz & M��fit ��zulug, 2007, A catalogue of the fishes held in the Istanbul University, Science Faculty, Hydrobiology Museum., pp. 29-54 in Zootaxa 1472 on page 32

Etmopterus lucifer, 2 specimens -Holotype. CAS-SU 6863, male, 278 mm TL, Misaki, Japan; CAS 23662, male, 308 mm TL, off Sandai, Japan., Published as part of Jayna A. Schaaf-da Silva & David A. Ebert, 2006, Etmopterus burgessi sp. nov., a new species of lanternshark (Squaliformes: Etmopteridae) from Taiwan., pp. 53-64 in Zootaxa 1373 on page 63

Etmopterus burgessi sp. nov. Broad-snout Lanternshark (Fig. 1, Table 1) Holotype. CAS 223476, 355 mm TL, adult male, Ta-Chi, Taiwan, 24�� 53' N, 122�� 01' E, collected by David A. Ebert, 11 May 1988. Paratypes. 3 specimens, all collected at Ta-Chi, Taiwan, 24�� 53' N, 122�� 01' E. CAS 223477 (Fig. 2), adult female, 406 mm TL, 22 May 2005; CAS 223478, juvenile female, 241 mm TL, 23 May 2005; CAS 223479, juvenile female, 239 mm TL, 21 May 2005. Diagnosis. A relatively large species of Etmopterus with the following combination of characters: snout broad, 8.0 (7.1-8.3)% TL; mouth broad and strongly arched; upper teeth in 2-3 series, each tooth with a median lanceolate cusp, flanked by three smaller lateral cusplets on each side; denticles short and robust, arranged in regular longitudinal rows over most of body; denticles conspicuous on ventral snout surface, and surface of second dorsal fin; second dorsal fin height short, 40.7 (34.6-36.2)% fin length; posterior branch of flank marking short, base of flank marking wide; caudal peduncle short, 11.4 (11.1-12.7)% TL; color in life charcoal grey above, darker grey to black below. Description. Body fusiform, relatively stout; head long 22.1 (21.1-22.2)% TL, not depressed, height 0.93 (0.65-0.77) times width. Pre-orbital snout long, rounded in lateral view, width broad, 8.0 (7.1-8.3)% TL, horizontal length 2.10 (2.10-2.20) in length of eye. Eye large, oval-shaped, length 5.22 (4.57-5.03) in head, 1.65 (1.73-2.09) times height. Spiracle large, semicircular, greatest diameter 3.06 (3.62-4.11) in eye length. Nasals very large, oblique; nostril width about equal to internarial distance; nasal flaps poorly developed, anterior flap triangular. Gill openings very large, slightly oblique, subequal in size, becoming progressively smaller posteriorly, height of first gill slit 2.1 (1.6-1.9)% TL. Mouth broad, strongly arched, width 1.18 (1.22-1.45) in preoral length. Teeth dissimilar in upper and lower jaws; upper teeth multicuspid in 2-3 series, each with a long median cusp flanked by lateral cusplets more than one half the length of median cusp; teeth in lower jaw in single series, unicuspid, interlocking, oblique, bladelike. Teeth in first row of upper jaw 26 (24-26), first row of lower jaw 40 (32-36). Vertebral count: 65. Dermal denticles small, low, unicuspid with erect thorn-like conical crowns, recurved posteriorly; arranged in regular longitudinal rows on trunk, back, caudal fin, and second dorsal fin. Dermal denticles irregular on abdomen, flank markings above pelvic fins, and caudal markings. Pelvic flank marking poorly defined, consisting of both an anterior and posterior branch; anterior branch long, slender, 8.1 (7.1-9.8)% TL, extending just forward of pelvicfin origin; posterior branch truncated distally, slightly shorter than anterior branch, 7.3 (8.4-9.2)% TL, terminating well past the second dorsal-fin insertion and in line with edge of free rear tip; base of flank marking wide. Caudal marking base broad, forming a saddle anteriorly, covering the entire ventral surface of posterior caudal peduncle; at about the lower caudal-lobe origin, marking extends dorsally to a level almost half of caudal peduncle height, tapering to a point forming the central caudal marking. Central caudal marking narrowly ovoid, slightly longer than eye length, 5.7 (4.7-6.4)% TL. Viewed laterally, central caudal marking is well apart from the flank marking, separated by at least one central caudal marking length. Upper caudal marking narrow, about equal to length of central caudal marking, 5.9 (6.4-7.4)% TL. Distal margins of fins largely covered with skin, not fringed with naked ceratotrichia. Each dorsal fin with a grooved spine just anterior to the fin origin. First dorsal fin spine small, less than half the length of second dorsal spine. First dorsal fin low, slightly rounded, length 7.4 (6.5-7.0)% TL; fin-spine origin well behind pectoral-fin rear tip; insertion of base well forward of pelvic-fin origin; much smaller than second dorsal fin. Length of first dorsal fin 1.65 (1.38-1.48) in second dorsal fin; height of first dorsal fin 1.62 (1.75-2.13) in second dorsal fin. Second dorsal fin relatively small, upright, concave posterior margin, free rear tip long, spine length 0.74 (0.41-0.51) in height of second dorsal fin; fin-spine origin just behind insertion of pelvic fin; interdorsal space 0.96 (0.92-1.06) in length from snout tip to pectoral-fin origin. Dorsal -caudal space relatively short 11.4 (11.1-12.7)% TL. Coloration. In life very darkly pigmented; charcoal gray to blackish. Dorsum, ventrum, and lateral surfaces have no obvious boundaries. Belly and flank markings may be weakly displayed; magnification shows an increased density of melanophores. Upper caudal lobe very dark; lower lobe and tips of fins sometimes lighter grey but not translucent. Size. Maximum length at least 355 mm for adult male (holotype) and 406 mm for adult female (paratype CAS 223477). Distribution. All four type specimens were collected on the continental slope off northeastern Taiwan at depths greater than 300 m. Etymology. The species is named in recognition of George Burgess for his contributions to the systematics of this genus. Discussion Etmopterus burgessi can be separated from other western North Pacific Etmopterus species based on a combination of the following characters: snout width, first gill slit length, caudal peduncle length, second dorsal fin height, flank and caudal markings, tooth morphology, and dermal denticle placement. The snout width of E. burgessi increases in proportion to head length (Fig. 3) compared to the other four regional examples. Etmopterus brachyurus and E. splendidus have snout widths that decrease slightly with increased head length. The sample size for E. lucifer and E. molleri are not robust, but both cluster closely with E. brachyurus. The gill slit length of E. burgessi is relatively long (Fig. 4) in proportion to head length. In the new species, as head length increases, so does the length of the first gill slit in proportion to the head length. For E. brachyurus, the relationship is negatively allometric, with head length increasing as the length of the first gill slit decreases proportionately. The length of the caudal peduncle is shorter in E. burgessi than in the other species examined. The measurement of this length is 11.4 (11.1-12.7)% TL in E. burgessi, but 13.7-15.5% TL in E. brachyurus and 12.8% TL in E. lucifer. Etmopterus molleri has a dorsal -caudal fin space of 14.1-14.9% TL, and E. splendidus has a measurement of 12.5-14.8% TL. The height of the second dorsal fin in E. burgessi is relatively low; its height ranges from to 34.6-40.7% of its overall length (Fig 5), and overlaps with that of E. lucifer which ranges from 34.5-44.7%. This measurement in E. brachyurus is slightly higher, with a range of 42.1-48.4% fin length. The height of the second dorsal fin in E. molleri is much higher than for its congeners, ranging from 50-60%. Yamakawa et al. (1986) reports similar range values for the three aforementioned species. Etmopterus splendidus has a second dorsal fin height to length of 31.4-47.6%. Although percentages of the height in overall length overlap between species, the second dorsal fin is generally lower in E. burgessi than in either E. molleri or E. brachyurus. Etmopterus burgessi has a flank marking that is distinct from three of the four species examined; E. splendidus, E. molleri, and E. brachyurus (Fig. 5). Comparatively, the anterior extension of E. burgessi has a broad base, 21.9 (21.7-29.3)% of anterior extension and short anterior branch, 78.1 (70.1-78.3)% of anterior extension. Etmopterus splendidus also has a broad flank marking base (34.3-46.3% of anterior extension) and a relatively short anterior branch (53.7-65.7% of anterior extension), but lacks a posterior branch entirely. The flank marking base is relatively narrow in both E. molleri (12.3-16.7% anterior extension) and E. brachyurus (12.1-16.8% anterior extension). The new species has a relatively short and blunt posterior branch, 41.4 (42.3-45.3)% of total flank marking length, that is similar to E. lucifer (29.1-37.3%). In comparison, the flank marking length is much shorter, its base broader, and the branches shorter in E. burgessi than either E. brachyurus or E. molleri. The upper tooth morphology of E. burgessi consists of a single medial cusp, flanked on each side by three lateral cusplets, a condition similar to E. splendidus (Yano, 1988). However, the upper tooth morphology of E. brachyurus (Smith and Radcliffe, 1912; Compagno et al., 2005; direct observation), E. lucifer (Compagno et al., 2005; direct observation), and E. molleri (Whitley, 1939; Compagno et al., 2005) consists of a medial cusp flanked by two cusplets on each side. The dermal denticles of E. burgessi are present over most of the body surface, and are useful for identification. Indeed, dermal denticles are present on the second dorsal fin surface of the new species. All of the other species examined have denticles on this surface except E. molleri, for which this area is relatively smooth (Yamakawa et al., 1986; direct observation). The ventral surface of the snout of E. burgessi has a thick grouping of dermal denticles, but in contrast, E. brachyurus displays a pattern where denticles are lacking around the nasals and along the medial section of the snout. The ventral snout of E. lucifer and E. molleri is sparsely covered with denticles. The known distribution of Etmopterus species in the western North Pacific is problematic given the apparent high degree of endemism within this group. At the present, E. burgessi is known only from off the east coast of Taiwan. The other four ��� E. lucifer group ��� etmopterids, E. brachyurus, E. lucifer, E. molleri, and E. splendidus have variously been reported from Taiwanese waters. However, despite extensive field sampling and examination of museum specimens, we can only confirm the presence of E. brachyurus and E. splendidus, both of which were quite commonly observed at Taiwanese fish markets. Etmopterus brachyurus, originally described from the Philippines (Smith & Radcliffe, 1912), has been reported throughout the western Pacific from Japan to Australia (Compagno et al., 2005). Etmopterus splendidus was originally described from southern Japan (Yano, 1988) and has since been found in Taiwan and possibly Java, Indonesia (Compagno et al., 2005). Although we did not collect E. lucifer or E. molleri both have variously been reported by several authors (Teng, 1959; Chen, 1963; Joung & Chen, 1992; Compagno et al. 2005) as occurring in Taiwanese waters. Shen and Ting (1972) documented two forms of E. lucifer from the waters around Taiwan. However, based on the illustrations of Shen and Ting (1972: Fig 2-A & 2-B) the two E. lucifer forms are actually examples of E. molleri and E. splendidus, respectively. Furthermore, according to Joung and Chen (1992), the description of E. lucifer by Chen (1963) is actually that of E. molleri; an observation with which we concur. Compagno et al. (2005) commented that the presence of E. lucifer, originally described from Japan, needed confirmation from some western Pacific localities. Etmopterus molleri, originally described from Australian waters, has been reported to occur throughout the western Pacific, including New Zealand, Taiwan, and Japan (Compagno et al., 2005). Although this species has been illustrated (Shen & Ting 1972) and reported (Joung & Chen, 1992) as occurring in Taiwanese waters, we could not confirm its occurrence., Published as part of Jayna A. Schaaf-da Silva & David A. Ebert, 2006, Etmopterus burgessi sp. nov., a new species of lanternshark (Squaliformes: Etmopteridae) from Taiwan., pp. 53-64 in Zootaxa 1373 on pages 54-61

Etmopterus molleri, 3 specimens -CAS-SU 23779, female, 347 mm TL, off Sagami Nada; CAS 11225, female, 293 mm TL; CAS 11225, female, 265mm TL, off Misaki, Japan., Published as part of Jayna A. Schaaf-da Silva & David A. Ebert, 2006, Etmopterus burgessi sp. nov., a new species of lanternshark (Squaliformes: Etmopteridae) from Taiwan., pp. 53-64 in Zootaxa 1373 on page 63