Alewives Alosa pseudoharengus, copyright David J. Stang.

Belongs within: Clupeiformes.
Contains: Anodontostomatini, Alosa (Alosa), Sardinella, Clupea.

The Clupeidae, herrings, are mostly small fishes that are most diverse in coastal marine waters with some species entering fresh water. Members of the subfamilies Alosinae and Dorosomatinae have a distinct median notch or cleft in the front of the upper jaw whereas the upper jaw is rounded in front view in the Clupeinae and Pellonulinae (Munroe et al. 1999).

Many kinds of herring
Published 7 August 2016
The original herring: Baltic herrings Clupea harengus membras, copyright Riku Lumiaro.

The subject of today’s post is something that I’m sure that you’ve all encountered at one time or another. It’s a group of animals that features highly in the world’s food supply. Some of you may be grat fans of these animals and seek them out on a regular basis; others may not be so enthused. They go by many names: herring, sardines, sprats, shad… but all are members of the fish family Clupeidae.

For the most part, clupeids are a prime example of what I think of as ‘fishy’ fish: that is, fish that look exactly how the majority of people imagine a fish to look (as opposed, say, to some of those deep-sea jobs that are all teeth and poor muscle tone). They are most diverse in marine waters of the continental shelf though many spend part or all of their lives in fresh water. Most form schools, sometimes very large ones; it is this tendency to congregate that makes them such an important part of the food chain for humans and other predators. The clupeids themselves are mostly micro-predators, feeding on minute plankton. Most are medium-sized to small fish with large species getting up to a couple of feet in length*. Conversely, species of the south-east Asian freshwater genus Sundasalanx (on which more below) reach maturity at only 15 mm in length.

*Bond (1996) makes the remarkable statement that “Palonia castelnaudi, a freshwater herring of South America, reaches at least 1.5 m (Dr. Barry Chernoff, personal communication)“. Not only have I been unable to find another reference to a clupeid of this size, I have been unable to confirm the existence of a species of this name. The same reference gives a maximum length for the Chirocentridae as 3.5 m; a quick search online suggests the correct figure is less than a third of that.

Another commercially significant species: sardines Sardina pilchardus, photographed by Alessandro Duci.

The exact circumscription of the Clupeidae has varied over time. It is the largest family in a clade called the Clupeoidei which is well defined by characters such as a reduction in the lateral line and the presence of the recessus lateralis, a channel running through the pterotic bone between the swim bladder and the inner ear. Other families within the Clupeoidei are the Engraulidae (anchovies), Pristigasteridae (ilishas) and Chirocentridae (wolf herrings). While each of the other families is fairly distinctive, the Clupeidae lack clear uniting features of their own and have tended to be defined as ‘the rest’. Historically, some authors have united some or all of the other families within the Clupeidae, or recognised clupeid subgroups as their own additional families.

It therefore would not have come as too much of a surprise when a molecular phylogenetic analysis of the Clupeoidei by Lavoué et al. (2013) did not identify the Clupeidae as a monophyletic group. Instead, both the Pristigasteridae and Chirocentridae were nested within the Clupeidae. What is more, not one of the five subfamilies currently recognised within the clupeids was monophyletic either. Instead, Lavoué et al. found six distinct sublineages within the clupeids; each of these was individually well supported but the broader relationships between them were not. Four of these potentially formed a clade that may correspond to a restricted Clupeidae. However, members of the ‘Dussumieriinae’ (which differ from other clupeids in the shape of their pelvic scutes) formed two external lineages: one was potentially the sister group to all other clupeoids except the Engraulidae whereas the round herring genus Etrumeus was weakly placed as sister to the Chirocentridae. To the best of my knowledge, no-one has yet suggested a formal reclassification of the clupeoids as a result of such studies, but it seems likely that we will either see the Clupeidae expanded to include the chirocentrids and pristigasterids, or restricted to exclude the dussumieriines. Again, either one of these options would align with alternative classifications used in the past.

The paedomorphic Sundasalanx microps, copyright Michael Lo.

Also of note in recent studies on clupeid phylogeny is the position of the south-east Asian freshwater genus Sundasalanx. When first described in 1981, this genus was not recognised as a clupeid or even as a clupeoid. Instead, it was originally placed in the fish order Osmeriformes, the smelts, together with another fish genus Salanx. Members of these two genera are indeed similar in appearance: they are tiny and transparent, looking overall like whitebait but never growing into a larger adult. However, a study of the morphology of Sundasalanx in 1997 lead to the conclusion that the shared features of Salanx and Sundasalanx were actually convergences resulting from both exhibiting paedomorphy, becoming reproductively mature while still effectively in the larval stage. A relationship of Sundasalanx to the clupeoids was suggested instead and this was later corroborated by molecular analyses (Ishiguro et al. 2005). In fact, Sundasalanx is nested well within the Clupeidae, even in the family’s restricted sense. Recent years have seen something of a surge in descriptions of paedomorphic fish (many of which were previously mistaken for juveniles of related taxa). Lavoué et al. (2008) recorded another paedomorphic clupeoid from marine waters of south-east Asia that the identified by molecular analysis as related to the dussumieriines, but to the best of my knowledge this species remains unnamed.

Systematics of Clupeidae

Characters (from Munroe et al. 1999): Typically fusiform fishes, oval in cross-section, and sometimes strongly compressed; complete series of scutes along abdomen (pelvic scute always present); mouth terminal, usually with two supramaxillae, and small or minute jaw teeth. No spiny rays in dorsal fin; single dorsal fin, which is short and near midpoint of body. Pectoral fins set low on body; pelvic fins just anterior to, below, or just posterior to dorsal-fin base; anal fin short (usually less than 28 soft rays) with origin well posterior to vertical through base of posteriormost dorsal-fin ray; caudal fin always deeply forked. Scales cycloid, adherent and of moderate size (about 35 to 68 in lateral series). Eyelids usually with vertical opening in middle. Lateral line absent. Scales without posterior striations. Colour typically blue-green on back and silvery on flanks; with variable darker markings including spot behind gill cover, spots along flanks, spot at dorsal-fin origin, and dark pigmentation on part of dorsal, pectoral, anal, and caudal fins.

| |--Pellonula leonensisEA03
| |--ClupeichthysMWN99
| |--HyperlophusMWN99
| | |--H. translucidus McCulloch 1917MWN99
| | `--H. vittatus (Castelnau 1875)MWN99
| `--CoricaMWN99
| |--C. laciniata Fowler 1935MWN99
| `--C. soborna Hamilton-Buchanan 1822MWN99
|--Dorosomatinae [Dorosomidae]MWN99
| |--AnodontostomatiniNR73
| |--Clupanodontini [Clupanodontidae]NR73
| | |--Konosirus Jordan & Snyder 1900 [=Nealosa Herre & Myers 1931]NR73
| | | `--*K. punctatus (Temminck & Schlegel 1846)NR73, MWN99 (see below for synonymy)
| | `--Clupanodon Lacepède 1803 [=Thrissa Rafinesque 1815; Clupanodontidae, Clupanodontini]NR73
| | |--*C. thrissa (Linnaeus 1758)NR73 (see below for synonymy)
| | |--C. oglina (Lesueur 1818) [=Megalops oglina]NR73
| | `--C. pseudohispanicusF16
| `--Dorosoma Rafinesque 1820 [Dorosomatini]NR73
| |--D. (Dorosoma)NR73
| | |--D. (D.) cepedianum (Lesueur 1818) [=Megalops cepediana; incl. *D. notata Rafinesque 1820]NR73
| | |--D. (D.) anale Meek 1904NR73
| | |--D. (D.) chavesi Meek 1907NR73
| | `--D. (D.) smithi Hubbs & Miller 1941NR73
| `--D. (Signalosa Evermann & Kendall 1898)NR73
| `--D. (S.) petenense (Günther 1866) (see below for synonymy)NR73
| |--Hilsa kelee (Cuvier 1829)MWN99 (see below for synonymy)
| |--TenualosaMWN99
| | |--T. ilisha (Hamilton-Buchanan 1822)MWN99 [=Hilsa ilishaB96]
| | |--T. macrura (Bleeker 1852) [=Alausa macrurus, Hilsa macrura, Macrura macrura]MWN99
| | |--T. reevesii (Richardson 1846) (see below for synonymy)MWN99
| | |--T. thibaudeaui (Durand 1940)MWN99
| | `--T. toli (Valenciennes 1847) [=Hilsa toli; incl. Alausa ctenolepis Bleeker 1852]MWN99
| `--Alosa [incl. Caspialosa]FWA06
| | i. s.: A. alabamaeB96
| | A. fintaP61
| | ‘Caspialosa’ kessleriP61
| | |--C. k. kessleriP61
| | `--C. k. volgensisP61
| | A. pontica (Eichwald 1838)SE08
| |--A. (Alosa)FWA06
| `--A. (Pomolobus)FWA06
| |--A. (P.) aestivalis (Mitchill 1814)FWA06, M02
| |--A. (P.) chrysochlorisFWA06
| |--A. (P.) mediocris (Mitchill 1814)FWA06, M02
| `--A. (P.) pseudoharengus (Wilson 1811)FWA06, M02
| |--E. elongata Wongratana 1983MWN99
| `--E. thoracata (Valenciennes 1847)MWN99 (see below for synonymy)
| |--A. clupeoides Bleeker 1849 [=Sardinella clupeoides]MWN99
| |--A. leiogaster (Valenciennes 1847) [=Sardinella leiogaster; incl. Clupea okinawensis Kishinouye 1908]MWN99
| |--A. perforatumJR10
| `--A. sirm (Walbaum 1792)MWN99 (see below for synonymy)
| |--S. caeruleaCH97
| |--S. melanostictusIM04
| |--S. neopilchardus (Steindachner 1879)MWN99
| |--S. ocellataW81
| `--S. sagaxB96
|--H. blackburni (Whitley 1948)MWN99
|--H. castelnaui (Ogilby 1897)MWN99
|--H. collettei Wongratana 1987MWN99
|--H. dispilonotus (Bleeker 1852) [=Harengula dispilonotus]MWN99
|--H. gotoi Wongratana 1983MWN99
|--H. koningsbergeri (Weber & de Beaufort 1912) [incl. Harengula maccullochi Whitley 1931]MWN99
|--H. lippa (Whitley 1931)MWN99
|--H. punctatusMWN99
`--H. quadrimaculatus (Rüppell 1837)MWN99

Clupeidae incertae sedis:
Sprattus sprattusB96
Palonia castelnauiB96
Amazonsprattus scintillaB96
Ethmalosa fimbriataEA03
Opisthonema Gill 1862M02, BR05
`--O. oglinum (LeSueur 1818)M02
|--E. teres (DeKay 1842)M02
`--E. whiteheadiM02
|--C. multispinis Hardenberg 1941M58
`--C. papuensis (Ramsay & Douglas-Ogilby 1886) [=Corica papuensis]M58
Kowala venulosa (Weber & de Beaufort 1912) [=Clupeoides venulosus]M58
|--S. neopilchardus Steindachner 1879 (see below for synonymy)YT27
`--S. pilchardusBM03
|--B. gunteriP61
|--B. patronusM02
|--B. smithiM02
`--B. tyrannus (Latrobe 1802)M02
|--H. abbreviataG75
|--H. antipoda (Hector 1872) (see below for synonymy)YT27
|--H. humeralisF15
|--H. koningsbergeri (Weber & de Beaufort 1912) [=Clupea (Harengula) koningsbergeri]M58
|--H. macrolepisG75
|--H. macrophthalmusF15
|--H. ovalis (Bennett 1830) (see below for synonymy)M58
`--H. schrammii (Bleeker 1849) [=Alausa schrammii]M58
Sierrathrissa leonensisMWN99
Thrattidion noctivagusMWN99
Knightia vetusta Grande 1982P93
Stolothrissa tanganicae Regan 1907B50, KC-L22

Inorganic: Harengula zunsai minilorientalis Okamura 1987O87

Amblygaster sirm (Walbaum 1792)MWN99 [=Clupea sirmM58, Sardinella sirmMWN99; incl. Sardinops dakini Whitley 1937MWN99, Sardinella leiogastroides Bleeker 1854MWN99, C. pinguis Günther 1872MWN99, Harengula pinguisM58]

*Clupanodon thrissa (Linnaeus 1758)NR73 [=Clupea thrissaM58, Dorosoma thrissaNR73, Konosirus thrissaJR10, Meletta thryssaNR73, *Thrissa thrissaNR73; incl. Clupanodon haihoensis Oshima 1926MWN99, Chatoessus maculatus Richardson 1846MWN99, Dorosoma maculatumNR73, D. maculatusNR73, Chatoessus osbeckii Valenciennes 1848NR73, Clupanodon osbeckiiNR73, Dorosoma osbeckiiNR73, Clupea triza Linnaeus 1758MWN99, Dorosoma trizaNR73]

Dorosoma (Signalosa) petenense (Günther 1866) [=Meletta petenensis, Signalosa petenensis; incl. *S. atchafalayae Evermann & Kendall 1898]NR73

Escualosa thoracata (Valenciennes 1847)MWN99 [incl. Clupea coval Cuvier 1829M58, Kowala covalMWN99, C. huae Tirant 1883MWN99, Meletta lile Valenciennes 1847MWN99, C. macrolepis Steindachner 1879MWN99]

Harengula antipoda (Hector 1872) [=Clupea spratus var. antipodum, Amblygaster antipodus, C. antipoda; incl. C. holodon Regan 1916]YT27

Harengula ovalis (Bennett 1830) [=Clupea ovalis; incl. C. kunzei Macleay 1880, Sardinella kunzei, C. moluccensis Günther 1880, C. (Harengula) moluccensis, Harengula punctata Whitley 1927, C. venenosa Meyer 1885]M58

Hilsa kelee (Cuvier 1829)MWN99 [=Macrura keleeMWN99; incl. Alosa brevis Bleeker 1848MWN99, M. brevisM58, Clupea durbanensis Regan 1906MWN99, M. durbanensisMWN99, Alausa kanagurta Bleeker 1852MWN99, Hilsa kanagurtaMWN99, Alosa malayana Bleeker 1866MWN99, Clupea platygaster Günther 1868MWN99]

*Konosirus punctatus (Temminck & Schlegel 1846)NR73, MWN99 [=Chatoessus punctatusNR73, Clupanodon punctataNR73, Cl. punctatusNR73, Dorosoma punctatumNR73; incl. Ch. aquosus Richardson 1846 (n. o.)NR73]

Sardinia neopilchardus Steindachner 1879 [=Clupanodon neopilchardus, Clupea neopilchardus, Amblygaster neopilchardus]YT27

Tenualosa reevesii (Richardson 1846) [=Hilsa reevesii, Macrura reevesii; incl. Clupea sinensis Linnaeus 1758 (n. d.)]MWN99

*Type species of generic name indicated


[BM03] Barreiros, J. P., T. Morato, R. S. Santos & A. E. de Borba. 2003. Interannual changes in the diet of the almaco jack, Seriola rivoliana (Perciformes: Carangidae) from the Azores. Cybium 27 (1): 37–40.

[B96] Bond, C. E. 1996. Biology of Fishes 2nd ed. Saunders College Publishing: Fort Worth.

[BR05] Bouchet, P., & J.-P. Rocroi. 2005. Classification and nomenclator of gastropod families. Malacologia 47 (1–2): 1–397.

[B50] Brooks, J. L. 1950. Speciation in ancient lakes (concluded). Quarterly Review of Biology 25: 131–176.

[CH97] Castro, P., & M. E. Huber. 1997. Marine Biology 2nd ed. WCB McGraw-Hill: Boston.

[EA03] Écoutin, J.-M., & J.-J. Albaret. 2003. Relation longueur-poids pour 52 espèces de poissons des estuaires et lagunes de l’Afrique de l’Ouest. Cybium 27 (1): 3–9.

[FWA06] Faria, R., S. Weiss & P. Alexandrino. 2006. A molecular phylogenetic perspective on the evolutionary history of Alosa spp. (Clupeidae). Molecular Phylogenetics and Evolution 40 (1): 298–304.

[F15] Fowler, H. W. 1915. Cold-blooded vertebrates from Florida, the West Indies, Costa Rica, and eastern Brazil. Proceedings of the Academy of Natural Sciences of Philadelphia 67 (2): 244–269.

[F16] Fowler, H. W. 1916. The fishes of Trinidad, Grenada, and St. Lucia, British West Indies. Proceedings of the Academy of Natural Sciences of Philadelphia 67 (3): 520–548.

[G75] Grant, E. M. 1975. Guide to Fishes. The Co-ordinator-General’s Department: Brisbane (Australia).

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Ishiguro, N. B., M. Miya, J. G. Inoue & M. Nishida. 2005. Sundasalanx (Sundasalangidae) is a progenetic clupeiform, not a closely-related group of salangids (Osmeriformes): mitogenomic evidence. Journal of Fish Biology 67: 561–569.

[JR10] Jordan, D. S., & R. E. Richardson. 1910. Check-list of the species of fishes known from the Philippine archipelago. Bureau of Printing: Manila.

[KC-L22] Kmentová, N., A. J. Cruz-Laufer, A. Pariselle, K. Smeets, T. Artois & M. P. M. Vanhove. 2022. Dactylogyridae 2022: a meta-analysis of phylogenetic studies and generic diagnoses of parasitic flatworms using published genetic and morphological data. International Journal for Parasitology 52: 427–457.

Lavoué, S., M. Miya, A. Kawaguchi, T. Yoshino & M. Nishida. 2008. The phylogenetic position of an undescribed paedomorphic clupeiform taxon: mitogenomic evidence. Ichthyol. Res. 55: 328–334.

Lavoué, S., M. Miya, P. Musikasinthorn, W.-J. Chen & M. Nishida. 2013. Mitogenomic evidence for an Indo-west Pacific origin of the Clupeoidei (Teleostei: Clupeiformes). PLoS ONE 8(2): e56485. doi:10.1371/journal.pone.0056485.

[L58] Linnaeus, C. 1758. Systema Naturae per Regna Tria Naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Laurentii Salvii: Holmiae.

[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 no. 1).

[M02] Munroe, T. A. 2002. Herrings. Family Clupeidae. In: Collette, B. B., & G. Klein-MacPhee (eds) Bigelow and Schroeder’s Fishes of the Gulf of Maine 3rd ed. pp. 111–160. Smithsonian Institute Press: Washington.

[MWN99] Munroe, T. A., T. Wongratana & M. S. Nizinski. 1999. Clupeidae. Herrings (also, sardines, shads, sprats, pilchards, and menhadens). 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. 1775–1821. FAO: Rome.

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[YT27] Young, M., & G. M. Thomson. 1927. Occurrence of pilchards and sprats in New Zealand seas. Transactions and Proceedings of the New Zealand Institute 57: 314–319.

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