Reviews in Fish Biology and Fisheries 9: 71–87, 1999.
© 1999 Kluwer Academic Publishers. Printed in the Netherlands.
Fish haemoglobins: the order Clupeiformes
Martino Rizzotti & Flavio Gioppato
Department of Biology, University of Padova, Via U. Bassi 58 B, 35131 Padova, Italy
Accepted 21 September 1998
Abstract page 71
Haemoglobin multiplicity 74
Systematic survey 74
Engraulidae, subfamily Engraulinae
Pristigasteridae, subfamily Pelloninae
Clupeidae, subfamily Clupeinae
Clupeidae, subfamily Alosinae
Clupeidae, subfamily Dorosomatinae
Phylogenetic remarks 84
The haemoglobin systems of the order Clupeiformes have been studied by several researchers in 41 species
belonging to three out of its ﬁve families. Most of them were investigated in the native form using electrophoretic
methods, and a few were also examined from the functional point of view. Both approaches corroborate the
widespread view that acidic and basic haemoglobin components, which are structurally and functionally distinct,
may be present in teleost ﬁsh. However, the former are always present, whereas the latter are often lacking,
depending on the taxonomic group. Both kinds of components are found in families Clupeidae and Pristigasteridae,
but only acidic ones in Engraulidae.
Most electrophoretic patterns show high multiplicity, and chieﬂy concern the acidic components. Ontogenetic
variation was described in three species. Individual variants were also observed in other species, although some of
these might be due to ontogenetic variations rather than genetic polymorphism.
Key words: clupeiformes, haemoglobin, multiplicity, ontogeny, phylogeny, polymorphism, Root effect
The order Clupeiformes is the only living order of
the subdivision Clupeomorpha (Teleostei). It includes
357 species which have been grouped into ﬁve fami-
lies (Table 1). From the classic survey on ﬁsh
haemoglobins by Riggs (1970) to recent ones (Brit-
tain, 1991; Di Prisco and Tamburrini, 1992; Inger-
mann, 1992; Weber, 1992; Pérez et al., 1995), little
or no attention has been paid to this order, despite
its worldwide distribution and great economic value.
Because its importance is well known, this inatten-
tion cannot be due to an underestimation. Rather, the
reason is that most data (both old and new) on its
haemoglobins are scattered among a large number of
short papers or are mixed with information on other
ﬁsh orders. This paper is an attempt to ﬁll this gap.
Haemoglobin systems may be characterized from
structural and functional standpoints. Initial work on
the contents of the red blood cells of a particular