Belongs within: Clupeiformes.
I think ikan
Published 15 September 2025
It seems that anchovies are a divisive fish. For some, their presence in a dish incites revulsion, to the degree that such a reaction has become a comic cliché. Personally, I adore the salty, oily flavour of anchovies, whether used as a pizza topping, to add body to a sauce, or even just eaten on their own. Whatever your own opinions on anchovies, one thing you may not have been aware of is that they are a diverse bunch, with numerous recognised genera and species. Among these are the members of the genus Stolephorus.
Stolephorus species are small anchovies found in coastal regions (to about 30 metres water depth) of the tropical Indo-Pacific. Most are less than ten centimetres in length when mature; the Indian anchovy S. indicus is a comparative giant that may reach fifteen centimetres (Wongratana et al. 1999). Identification of anchovies, even to genus level, can be a challenging business, but Stolephorus is characterised by the presence of needle-like prepelvic scutes, no postpelvic scutes, a relatively short anal fin, and a relatively long isthmus muscle on the underside of the jaw that reaches to the gill membrane at the front and covers the urohyal bone (this last feature is the primary distinction between Stolephorus and the similar Encrasicholina with which it is often confused). Close to fifty Stolephorus species have been described to date, and new ones continue to appear. Some species historically thought widespread (such as the aforementioned S. indicus) have proven to encompass multiple geographically distinct species.
Like other anchovies, Stolephorus species typically form schools for protection*. Stolephorus schools tend to prefer more muddy estuarine environments than other anchovies, congregating close to the seabed (Dalzell & Lewis 2024). Spawning occurs in brackish waters close to shore; eggs of Stolephorus lack buoyant oil globules so cannot develop in more open waters. Stolephorus anchovies may also migrate between brackish and saltwater habitats in search of food sources, of which copepods make up the greater part.
*Though how protective these schools are is, of course, a question of context (Dalzell & Lewis 2024). A study of predator responses to Commerson’s anchovy Stolephorus commersonii found that schools formed an effective defense against the mackerel tuna Euthynnus affinis, which was unable to target individual fish among the tight mass. However, bull sharks Carcharhinus leucas, for which individual anchovies would normally be too small a snack to bother with, would suddenly take interest when presented with a large fishy ball!
Commercially, Stolephorus species are regarded as less significant than more pelagic anchovies of the genus Engraulis. However, Stolephorus and Encrasicholina anchovies are commonly fished around eastern Africa and southern Asia. They may be used as the base of fish sauces, or eaten dried and salted (for instance, as the ikan of ikan bilis). They may also be used as live bait for tuna, or ground into fish meal. Because of the difficulty of distinguishing taxa, and the fact that they are more important to local than industrial fisheries, it is hard to say just which species are important where. The same challenges affect assessments of population levels and environmental threats. For instance, the species Stolephorus advenus is known from just a single specimen taken off the Coburg Peninsula in Australia’s Northern Territory, but whether this dearth of material reflects actual rarity or mere oversight remains uncertain. Whatever commercial significance they may have, Stolephorus anchovies are certain to also be significant as a food source for larger animals with which they share their habitat. It would be worthwhile to know how they are dealing with the challenges of a changing environment.
Systematics of Stolephorus
<==Stolephorus
|--S. advenus Wongratana 1987WMN99
|--S. andhraensis Babu Rao 1966WMN99
|--S. apiensis (Jordan & Seale 1906)WMN99 [=Anchovia apiensisM58]
|--S. baganensis Hardenberg 1933 (see below for synonymy)WMN99
|--S. brachycephalus Wongratana 1983WMN99
|--S. carpentariae (de Vis 1883) [incl. S. tysoni Wongratana 1983]WMN99
|--S. chinensis (Günther 1880) [=Anchoviella chinensis]WMN99
|--S. commersonii Lacepède 1803WMN99 (see below for synonymy)
|--S. dubiosus Wongratana 1983WMN99
|--S. gracilis [incl. S. japonicus]JR10
|--S. indicus (van Hasselt 1823)WMN99 [=Engraulis indicusWMN99; incl. E. samam inan Thiollière 1857M58]
|--S. insularis Hardenberg 1933 [incl. S. insularis oceanicus Hardenberg 1933]WMN99
|--S. multibranchus Wongratana 1987WMN99
|--S. nelsoni Wongratana 1987WMN99
|--S. pacificus Baldwin 1984WMN99
|--S. ronquilloi Wongratana 1983WMN99
|--S. tri (Bleeker 1852) [=Engraulis tri]M58
|--S. waitei Jordan & Seale 1926 [=Anchoviella waitei; incl. S. insularis bataviensis Hardenberg 1933]WMN99
`--S. zollingeri (Bleeker 1849) [=Engraulis zollingeri]M58
Stolephorus baganensis Hardenberg 1933 [=Anchoviella baganensis; incl. S. baganensis bengalensis (Dutt & Babu Rao 1959), S. baganensis macrops Hardenberg 1933]WMN99
Stolephorus commersonii Lacepède 1803WMN99 [=Anchovia commersoniiM58, Anchoviella commersoniiWMN99, Stolephorus commersonianusWMN99; incl. S. rex Jordan & Seale 1926WMN99]
*Type species of generic name indicated
References
Dalzell, P. J., & A. D. Lewis. 2024. The fisheries biology and ecology of anchovy genera Stolephorus and Encrasicholina in the Indo-West Central Pacific region. Fisheries Centre Research Reports 32 (1): 1–106.
[JR10] Jordan, D. S., & R. E. Richardson. 1910. Check-list of the species of fishes known from the Philippine archipelago. Bureau of Printing: Manila.
[M58] Munro, I. S. R. 1958. The fishes of the New Guinea region: a check-list of the fishes of New Guinea incorporating records of species collected by the Fisheries Survey Vessel “Fairwind” during the years 1948 to 1950. Papua and New Guinea Agricultural Journal 10 (4): 97–369 (reprinted: 1958. Territory of Papua and New Guinea Fisheries Bulletin 1).
[WMN99] Wongratana, T., T. A. Munroe & M. S. Nizinski. 1999. Engraulidae. Anchovies. In: Carpenter, K. E., & V. H. Niem (eds) The Living Marine Resources of the Western Central Pacific vol. 3. Batoid fishes, chimaeras and bony fishes part 1 (Elopidae to Linophrynidae) pp. 1698–1753. FAO: Rome.