Taxonomic status of seven nominal species of the anchovy genus Stolephorus described by Delsman (1931), Hardenberg (1933), and Dutt and Babu Rao (1959), with redescriptions of Stolephorus tri (Bleeker 1852) and Stolephorus waitei Jordan and Seale 1926 (Clupeiformes: Engraulidae)

  • Harutaka HataEmail author
  • Sébastien Lavoué
  • Hiroyuki Motomura
Full Paper


Examination of the original descriptions and available type specimens of Stolephorus insularis Delsman 1931 and subsequently proposed subspecies (Stolephorus insularis insularis Delsman 1931, Stolephorus insularis bataviensis Hardenberg 1933, Stolephorus insularis baweanensis Hardenberg 1933, and Stolephorus insularis oceanicus Hardenberg 1933), and Stolephorus baganensis baganensis Delsman 1931, Stolephorus baganensis megalops Delsman 1931, Anchoviella baganensis bengalensis Dutt and Babu Rao 1959, and Stolephorus waitei Jordan and Seale 1926, in addition to a large number of non-type specimens representing an extensive geographic range, confirmed the validity of S. baganensis, S. bataviensis, S. baweanensis, S. bengalensis, S. oceanicus, and S. waitei. Stolephorus i. insularis and S. b. megalops were recognized as junior synonyms of Stolephorus tri (Bleeker 1852). All of the valid species, including S. tri, are rediagnosed together with detailed color descriptions, with neotype designations for S. baganensis, S. bataviensis, S. baweanensis, S. bengalensis, S. oceanicus, S. i. insularis, and S. b. megalops and clarification of their taxonomic history. To complement our morphological observations, we built a large phylogenetic tree for 93 specimens of Stolephorus collected from the South China Sea belonging to ten species using two mitochondrial markers, cytochrome b and cytochrome oxidase I. Our phylogenetic results support the genetic distinctiveness of each of the ten species examined and offer the first hypothesis on their relationships.


Morphology Molecules Taxonomy Phylogenetics Neotype 



We thank R. Arrindel (AMNH), M. McGrouther, A. Hay, and S. Reader (AMS), K.-T. Shao and S.-P. Huang (ASIZP), O. Crimmen and J. Maclaine (BMNH), A. Suzumoto (BPBM), D. Catania (CAS), S. Kimura (FRLM), K. Hartel (MCZ), R. Causse and P. Pruvost (MNHN), H.-C. Ho and K. Koeda (NMMB), G. Shinohara, M. Nakae, and K. Kuriiwa (NSMT), W.-J. Chen and M. Lin (NTUM), K. Miyamoto (OCF), and J. Williams, K. Murphy, S. Raredon, and D. Pitassy (USNM) for opportunities to examine specimens of Stolephorus. We also thank Y. Haraguchi and other volunteers, and students of KAUM for curatorial assistance and G. Hardy (Ngunguru, New Zealand) for reading the manuscript and providing help with English. This study was supported in part by the Sasakawa Scientific Research Grant from the Japan Science Society (28-745); a Grant-in-Aid from the Japan Society for the Promotion of Science for JSPS Fellows (DC2: 29-6652); JSPS KAKENHI Grant Numbers JP19770067, JP26241027, JP24370041, JP23580259, and JP26450265; the JSPS Core-to-Core Program: B Asia-Africa Science Platforms; the “Biological Properties of Biodiversity Hotspots in Japan” project of the National Museum of Nature and Science, Tsukuba, Japan; “Establishment of Research and Education Network on Biodiversity and Its Conservation in the Satsunan Islands” project of Kagoshima University adopted by the Ministry of Education, Culture, Sports, Science and Technology, Japan; and the “Island Research” project by Kagoshima University. Malaysian specimens were collected during the JSPS Asian Core Program, “Establishment of Research and Education Network on Coastal Marine Science in Southeast Asia”, supported by the Ministry of Higher Education (Government of Malaysia), University Putra Malaysia, and University Malaysia Terengganu. Vietnamese specimens were collected with the support of the Institute of Marine Environment and Resources (Haiphone) and the Ha Long Bay Management Department (Ha Long), with permission for their use granted by the Biodiversity Conservation Agency, Ministry of Natural Resources and Environment (Hanoi). SL is grateful to Mr Sahat Ratmuangkhwang (Andaman Coastal Research Station for Development, Kasetsart University) and Dr Prachya Musikasinthorn (Faculty of Fisheries, Bangkok, Kasetsart University) for organizing fish survey in Thailand. SL’s contribution was supported by research grants MOST103-2119-M- 002-019-MY3 and MOST106-2119-M-002-032 from the Ministry of Science and Technology of Taiwan.


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Copyright information

© The Ichthyological Society of Japan 2019

Authors and Affiliations

  • Harutaka Hata
    • 1
    Email author
  • Sébastien Lavoué
    • 2
  • Hiroyuki Motomura
    • 3
  1. 1.Center of Molecular Biodiversity ResearchNational Museum of Nature and ScienceTsukubaJapan
  2. 2.School of Biological SciencesUniversiti Sains MalaysiaPenangMalaysia
  3. 3.The Kagoshima University MuseumKagoshimaJapan

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