Two new species of short-snouted dogfish sharks of the genus Squalus Linnaeus, 1758, from southern Africa (Chondrichthyes: Squaliformes: Squalidae)

  • Sarah T. de F. L. Viana
  • Mark W. Lisher
  • Marcelo R. de Carvalho
Original Paper


Species composition of the genus Squalus from the eastern Atlantic and western Indian oceans is still poorly known, with three to four species often recognized: S. acanthias, S. megalops, S. blainvillei, and S. mitsukurii. A fifth species, S. acutipinnis, was recently redescribed as valid in the region, although morphological variation was noted among its type specimens that indicated the presence of two different forms. The second form is described herein as Squalus margaretsmithae sp. nov. and distinguished from S. acutipinnis based on external morphological characters. Squalus mahia sp. nov. is also described from southern Africa based mostly on material collected from Madagascar. Squalus mahia sp. nov. stands out from other Squalus species by having a conspicuously slender body, elongate and thin dorsal fin spines, narrow caudal fin, and other external proportions. These two species are often misidentified with the Australian S. megalops because they share a short snout, white postventral caudal margins, lanceolate and unicuspid dermal denticles, and a low number of vertebrae. Comparative analyses are provided here between Squalus margaretsmithae sp. nov., Squalus mahia sp. nov., and morphologically similar species that also share these features, including S. brevirostris, S. crassispinus, S. acutipinnis, S. altipinnis, and S. hemipinnis. A taxonomic evaluation of the nominal species S. megalops and its occurrence in the region is also discussed.


Squalus New species Taxonomy Eastern Atlantic Ocean Western Indian Ocean 



We thank O. Gon and R. Bills (SAIAB), D. Clark and M. Bougaardt (SAM), M. McGrouther and S. Reader (AMS), L. Rocha and J. Fong (CAS), P. Last, W. White, A. Graham and J. Pogonoski (CSIRO), T. Kawai and students (HUMZ), H. Motomura (KAUM), P. Pruvost (MNHN), J. Maclaine (NHM), A. Palandacic (NMW), M. Nakae (NSMT), and A. Datovo and M. Gianeti (MZUSP), J. Johnson, (QM), R. de Ruiter (RMNH), C. Lamour (ZMB), and K. Sakamoto (ZUMT) for their help with specimens and museum facilities. Special thanks to N. Mazungula and B. Konqobe (SAIAB), R. Thiel, I. Eidus and S. Weigmann (ZMH) for providing data, pictures and radiographs of some types. We also thank A. Veríssimo (CIBIO/VIMS), K. Jensen (KU) and J. N. Caira (UCONN) for sending photo vouchers for comparisons. This study was funded by the Fundação de Amparo a Pesquisa do Estado de São Paulo (FAPESP - Processo 2011/18861-7; 2014/26503-1), Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq 158773/2011-0) and Geddes Collection Visiting Fellowship (2013), Australian Museum (Sydney) to the first author. The second author thanks the crew of RV Dr. Fridtjof Nansen and Agulhas and Somali Current Large Marine Ecosystems Project (ASCLME). The third author is supported by a grant from CNPq (305271/2015-6).


  1. Baranes A (2003) Sharks from the Amirantes Islands, Seychelles, with a description of two new species of squaloids from the deep sea. Israel J Zool 49(1):33–65CrossRefGoogle Scholar
  2. Bass AJ, D’Aubrey JD, Kistnasamy N (1976) Sharks of the east coast of southearn Africa. VI The families Oxynotidae, Squalidae, Dalatiidae and Echinorhinidae. Invest Rep, Oceanogr Res Inst 45(1):1–103Google Scholar
  3. Bass AJ, Compagno LJV, Heemstra PC (1986) Squaliformes. In: Smith MM, Heemstra PC (eds) Smiths’ sea fishes. Macmillan, Johannesburg, pp 49–62Google Scholar
  4. Bianchi G (1986) Fichas FAO de identificaçao de espécies para propósitos comerciais. Guia de campo para espécies comerciais marinhas e de águas salobras de Angola. FAO, RomaGoogle Scholar
  5. Bianchi G, Carpenter KE, Roux JP, Molloy FJ, Boyer D, Boyer HJ (1999) FAO species identification field guide for fishery purposes. the living marine resources of Namibia. FAO, RomeGoogle Scholar
  6. Bigelow HB, Schroeder WC (1948) Sharks. In: J. Tee-Van et al. (eds) Fishes of the Western North Atlantic. Part 1. Memoir Sears Foundation for Marine Research, Yale University, New HavenGoogle Scholar
  7. Bigelow HB, Schroeder WC (1957) A study of the sharks of the suborder Squaloidea. Bull Mus Comp Zool, Harvard College 117(1):1–150Google Scholar
  8. Caira JN, Jensen K, Barbeau E (eds) (2016) Global Cestode Database. World Wide Web electronic publication. Accessed January 16 2017
  9. Cappetta H (1987) Chondrichthyes II. Mesozoic and Cenozoic Elasmobranchii. In: Schultze HP (ed) Handbook of paleoichthyology. vol. 3B. Verlag Dr. Friedrich Pfeil, München, pp 1–193Google Scholar
  10. Compagno LJV (1984) FAO species catalogue. vol. 4. sharks of the world. an annotated and illustrated catalogue of shark species known to date. Part 1. Hexanchiformes to Lamniformes. FAO Fisheries Synopsis, No.125, vol. 4. FAO, RomeGoogle Scholar
  11. Compagno LJV (1988) Sharks of the order Carcharhiniformes. Princeton University Press, PrincetonGoogle Scholar
  12. Compagno LJV (2016) Sharks. In: Carpenter KE, De Angelis N (eds) The living marine resources for Eastern Central Atlantic. volume 2: bivalves, gastropods, hagfishes, sharks, batoid fishes, and chimaeras. FAO Species Identification Guide for Fisheries Purpose. FAO, Rome, pp 1123–1336Google Scholar
  13. Compagno LJV, Niem VH (1998) Several family accounts. In: Carpenter KE, Niem VH (eds) The living marine resources of the western central pacific. Cephalopods, crustaceans, holothurans and sharks. FAO species identification guide for fishery purposes. FAO, Senckenbergiana Biologica, Rome, 2, pp 687–1396Google Scholar
  14. Compagno LJV, Ebert DA, Smale MJ (1989) Guide to the sharks and rays of Southern Africa. Struik, Cape TownGoogle Scholar
  15. Compagno LJV, Ebert DA, Cowley PD (1991) Distribution of offshore demersal cartilaginous fish (Class Chondrichthyes) off the West coast of Southern Africa, with notes on their systematics. S Afr J Mar Sci 11(1):43–139CrossRefGoogle Scholar
  16. Compagno LJV, Dando M, Fowler S (2005) Sharks of the world. Harper Collins Publishers, LondonGoogle Scholar
  17. Viana ST de F, de Carvalho MR (2016) Redescription of Squalus acutipinnis Regan, 1908, a valid species of spiny dogfish from Southern Africa (Chondrichthyes: Squaliformes: Squalidae). Copeia 104(2):539–553Google Scholar
  18. Viana ST de F, de Carvalho MR, Gomes UL (2016) Taxonomy and morphology of species of the genus Squalus Linnaeus, 1758 from the Southwestern Atlantic Ocean (Chondrichthyes: Squaliformes: Squalidae). Zootaxa 4133(1):1–89Google Scholar
  19. Deynat PP, Séret B (1996) Le revêtement cutané des raies (Chondrichthyes, Elasmobranchii, Batoidea). I: morphologie et arrangement des denticules cutanés. Ann Sci Nat Zool 17(2):65–83Google Scholar
  20. Duffy CAJ, Last PR (2007) Redescription of the Northern Spiny Dogfish Squalus griffini Phillipps, 1931 from New Zealand. In: Last PR, White WT, Pogonoski JJ (eds) Descriptions of new Dogfishes of the Genus Squalus (Squaloidea: Squalidae). CSIRO Marine and Atmospheric Research Paper, Hobart, pp 91–100Google Scholar
  21. Ebert DA (2013) Deep-sea cartilaginous fishes of the Indian Ocean. vol 1 – sharks. FAO Species Catalogue for Fisheries Purposes no.8. FAO, RomeGoogle Scholar
  22. Ebert DA (2015) Deep–sea cartilaginous fishes of the Southeastern Atlantic Ocean. FAO Species Catalogue for Fishery Purposes, No.9. FAO, RomeGoogle Scholar
  23. Ebert DA, Stehmann M (2013) Sharks, batoids and chimaeras from the North Atlantic. FAO Species Catalogue for Fisheries Purposes, no.7. FAO, RomeGoogle Scholar
  24. Ebert DA, White WT, Goldmann KJ, Compagno LJV, Daly-Engel TS, Ward RD (2010) Resurrection and redescriptions of Squalus suckleyi (Girard, 1854) from the North Pacific, with comments on the Squalus acanthias subgroup (Squaliformes: Squalidae). Zootaxa 2612(1):22–40Google Scholar
  25. Ebert DA, Fowler S, Compagno LJV (2013) Sharks of the world: a fully illustrated guide. Wild Nature Press, LondonGoogle Scholar
  26. Garrick JAF (1960) Studies on New Zealand Elasmobranchii. Part XII. The species of Squalus from New Zealand and Australia; and a general account and key to the New Zealand Squaloidea. T Roy Soc New Zeal 88(3):519–557Google Scholar
  27. Garrick JAF (1961) A note on the spelling of the specific name of immaculate spiny dogfish, Squalus blainvillei (Risso, 1826). T Roy Soc New Zeal 88(4):843Google Scholar
  28. Heemstra PC, Heemstra E (2004) Coastal fishes of Southern Africa. National Inquiry Service Centre (NISC) and South African Institute for Aquatic Biodiversity (SAIAB), GrahamstownGoogle Scholar
  29. Herman J, Hovestadt-Euler M, Hovestadt DC (1989) Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites in living supraspecific taxa of Chondrichthyes fishes. Part A: Selachii. No 3: Order Squaliformes – Families Echinorhinidae, Oxynotidae and Squalidae. Bull de l’Institute Royal des Sciences Naturalle de Belgique, Biologie 59(1):101–157Google Scholar
  30. Iglésias SP (2012) Chondrichthyans from the North-eastern Atlantic and the Mediterranean (A natural classification based on collection specimens, with DNA barcodes and standardized photographs). Volume I, Provisional version 06Google Scholar
  31. Jungersen HFE (1899) On the appendices genitales in the greenland shark Somniosus microcephalus (Bl. Schn.) and other selachians. The Danish Ingolf-Expedition, 2, Bianco Luno, CopenhagenGoogle Scholar
  32. Kousteni V, Kasapidis P, Kotoulas G, Megalofonou P (2016) Evidence of high genetic connectivity for the longnose spurdog Squalus blainville in the Mediterranean Sea. Medit Mar Sci 17(2):371–383Google Scholar
  33. Krefft G (1968) Knorpelfische (Chondrichthyes) aus dem tropischen Ostatlantik. Atlantide Rep 10:33–81Google Scholar
  34. Last PR, White WT, Pogonoski JJ (2007a) Descriptions of new dogfishes of the genus Squalus (Squaloidea: Squalidae). CSIRO Marine and Atmospheric Research Paper, HobartGoogle Scholar
  35. Last PR, White WT, Stevens J (2007b) Part 5 - New species of Squalus of the highfin megalops group from the Australasian region. In: Last PR, White WT, Pogonoski JJ (eds) Descriptions of New Dogfishes of the Genus Squalus (Squaloidea: Squalidae). CSIRO Marine and Atmospheric Research Paper, Hobart, pp 39–53Google Scholar
  36. Last PR, Marshall LJ, White WT (2007c) Part 8 - Squalus nasutus sp. nov, a new long-snouted spurdog of the S. japonicus group from the Indian Ocean. In: Last PR, White WT, Pogonoski JJ (eds) Descriptions of New Dogfishes of the Genus Squalus (Squaloidea: Squalidae). CSIRO Marine and Atmospheric Research Paper, Hobart, pp 83–90Google Scholar
  37. Linnaeus C (1758) Systema Naturae. Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decimal, reformata.. Tomus I, Sweden, HolmiaeGoogle Scholar
  38. Macleay W (1881) A descriptive catalogue of Australian fishes. Part IV. P Linn Soc N S W 6(2):202–387Google Scholar
  39. Marouani S, Chaâba R, Kadri H, Saidi B, Bouain A, Maltagliati F, Last PR, Séret B, Bradai MN (2012) Taxonomic research on Squalus megalops (Macleay, 1881) and Squalus blainvillei (Risso, 1826) (Chondrichthyes: Squalidae) in Tunisian waters (Central Mediterranean Sea). Sci Mar 76(1):97–109CrossRefGoogle Scholar
  40. Muñoz-Chápuli R, Ramos F (1989) Morphological comparisons of Squalus blainvillei and S. megalops in the Eastern Atlantic, with notes on the genus. JPN J Ichthyol 36(1):6–21Google Scholar
  41. Myagkov NA, Kondyurin VV (1986) Dogfishes, Squalus (Squalidae) of the Atlantic Ocean and comparative notes on the species of this genus from other regions. J Ichthyol 26(6):1–18Google Scholar
  42. Naylor GJP, Caira JN, Jensen K, Rosana AM, White WT, Last PR (2012a) A DNA sequence-based approach to the identification of shark and rays species and its implication of global elasmobranch diversity and parasitology. B Am Mus Nat Hist 367(1):1–262CrossRefGoogle Scholar
  43. Naylor GJP, Caira JN, Jensen K, Rosana AM, Straube N, Lakner C (2012b) Elasmobranch Phylogeny: A mitochondrial estimate based on 595 species. In: Carrier JC, Musick JA, Heithaus ME (eds) Biology of Sharks and their relatives, 2nd edn. CRC Press, Boca Raton, pp 31–56CrossRefGoogle Scholar
  44. Regan CT (1908) A collection of fishes from the coasts of Natal, Zululand, and Cape Colony. Ann Natal Museum 1(3):241–255Google Scholar
  45. Risso A (1827) Histoire naturelle des principals productions de l’Europe méridionale, et particulièrement de celles des environs de Nice et des Alpes maritimes, vol. 3. FG Levrault, ParisGoogle Scholar
  46. Sabaj MH (2016) Standard symbolic codes for institutional resource collections in herpetology and ichthyology: an Online Reference. American Society of Ichthyologists and Herpetologists, Washington, DC. Version 6.5 (16 August 2016). Accessed 2 September 2016
  47. Schneider W (1990) FAO species identification sheets for fishery purposes. Field guide to the commercial marine resources of the Gulf of Guinea. prepared and published with the support of the FAO Regional Office for Africa. FAO, RomeGoogle Scholar
  48. Shirai S (1992) Squalean phylogeny: a new framework of “squaloid” sharks and related taxa. Hokkaido University Press, SapporoGoogle Scholar
  49. Springer VG, Garrick JAF (1964) A survey of vertebral numbers in sharks. Proc U S Nat Mus 116(3496):73–96CrossRefGoogle Scholar
  50. Tortonese E (1956) Fauna D’Italia. vol II – Lepdocaria, Cyclostomata, Selachii a cura di Enrico Tortonese. Edizioni Calderini, BolognaGoogle Scholar
  51. Vasil’eva ED, Sazonov NN, Verigina IA (2001) Pisces. In: Borissenko AV et al (eds) Types of vertebrates in the Zoological Museum of Moscow Univerisity. Moscow University Publisher, Moscow, pp 11–88Google Scholar
  52. Veríssimo A, Zaera-Perez D, Leslie R, Iglésias SP, Sèret B, Grigoriou P, Sterioti A, Gubili C, Barría C, Duffy C, Hernández S, Batjakas IE, Griffths AM (2016) Molecular diversity and distribution of Eastern Atlantic and Mediterranean dogfishes Squalus highlight taxonomic issues in the genus. Zool Scr. doi: 10.1111/zsc.12224 Google Scholar
  53. Weigmann S (2016) Annotated checklist of the living sharks, batoids and chimaeras (Chondrichthyes) of the world, with a focus on biogeographical diversity. J Fish Biol 88(3):837–1037CrossRefPubMedGoogle Scholar
  54. White WT, Iglésias SP (2011) Squalus formosus, a new species of spurdog shark (Squaliformes: Squalidae), from the Western North Pacific Ocean. J Fish Biol 79(4):954–968. doi: 10.1111/j.1095-8649.2011.03068.x CrossRefPubMedGoogle Scholar
  55. Whitehead PJP, Bauchot ML, Hureau JC, Nielsen J, Tortonese E (1984) Fishes of the North-Eastern Atlantic and the Mediterranean (FNAM). volume 1. UNESCO, ParisGoogle Scholar

Copyright information

© Senckenberg Gesellschaft für Naturforschung and Springer-Verlag Berlin Heidelberg 2017

Authors and Affiliations

  • Sarah T. de F. L. Viana
    • 1
  • Mark W. Lisher
    • 2
  • Marcelo R. de Carvalho
    • 1
  1. 1.Departamento de Zoologia, Instituto de BiociênciasUniversidade de São PauloSão PauloBrazil
  2. 2.South African Institute for Aquatic BiodiversityGrahamstownSouth Africa

Personalised recommendations