Reviews in Fish Biology and Fisheries 11: 171–179, 2002.

© 2002 Kluwer Academic Publishers. Printed in the Netherlands.

171

The origin of the salmonid fishes: marine, freshwater . . . or neither?

R.M. McDowall
National Institute of Water and Atmospheric Research, PO Box 8602, Christchurch, New Zealand (E-mail:
R.McDowall@NIWA.CRI.NZ)

Accepted 10 April 2002

Contents
Abstract page 171
Introduction 171
Diadromy in Salmonidae
An historical perspective on opinions regarding the origins of the Salmonidae 172
The geochronological place of diadromous species in salmonid history 174
Diadromy and phylogenetic analysis 174
Origin of Salmonidae in a cladistic context 175
Relationships of salmonids in the context of variability of diadromous life cycles 176
Conclusion 177
Acknowledgements 178
References 178

Key words: diadromy, freshwater, marine, origins, Salmonidae

Abstract

The evolutionary origins of the salmonid fishes, whether in freshwater or the sea, have been debated for
centuries. Early views favoured a group of marine ancestry invading freshwaters; more recently, there was a shift
towards a freshwater ancestry, on grounds that a return to freshwater to spawn indicates the ancestral biome.
Salmonids are widely believed to share an ancient common ancestry with the northern hemisphere Osmeridae
and southern hemisphere Retropinnidae and Galaxiidae. Salmonidae are diadromous, as are Osmeridae,
Retropinnidae and Galaxiidae. This suggests that diadromy is an ancient behavioral phenomenon across all
these groups, that the shared common ancestry of these groups was also diadromous, and that the ancestry of
Salmonidae was neither marine nor freshwater, but was amongst diadromous fishes. This begs the question of
whether this common ancestor was marine or freshwater, a question for which an answer seems likely to be elusive.

Introduction ancestry during more recent decades (see below).

The question of whether fishes of the family
Salmonidae have a marine or freshwater ancestry is
a venerable one that has been discussed often and
for decades. It has attracted the attention of some
great ichthyologists of the past, and continues to
attract comment. Opinions, summarized across nearly
two centuries, seem evenly divided over the answer,
though perhaps support has shifted from a majority
early belief in a marine origin, towards a freshwater
The question has been so intriguing probably because
of the apparently deep-seated (and usually correct)
notion that freshwater fishes are generally unable to
make a transition to marine habitats (or marine fishes
make the reverse transition – though this seems to have
been of less interest) and yet salmon and trout can
move, seemingly freely, between these two biomes.
That there has been so much debate about salmonid
origins is perhaps enigmatic, given lesser interest in
the same question for other fish that move equally
172
freely between marine and freshwater habitats – such
as lampreys (Petromyzontidae, Geotriidae), sturgeons
(Acipenseridae), anguillid eels (Anguillidae), smelts
(Osmeridae, Retropinnidae), shads (Clupeidae), and
so on (see McDowall, 1988).
Tchernavin (1939, p. 120) addressed the question
for salmonids, suggesting that the many questions
arising from consideration of the salmon life history
“finally resolve into the cardinal one: the origin of
salmon. One of its two present haunts must have
been the original habitat of the salmon’s ancestors”
(original italics). Thorpe (1982) concurred, consid-
ering the “central question” to be: “did the salmonids
originate in the sea or in fresh water?” Stearley
(1992, p. 662) thought this a “long-standing contro-
versy”, and found no “unanimity of opinion on the
ancestral condition and subsequent historical trends in
the life histories for the Salmonidae.” Balon (1968,
p. 6), addressing the question at a much narrower
focus, asked: “What is the original form – brown
or sea trout?” and concluded that “anadromous trout
have evolved gradually from an unknown fresh-water
ancestor.” But, in an apparent contradiction (p. 9), he
also thought that “the recent form of salmon and trout
are obviously descendants of the anadromous stock,
irrespective of their possibly marine or fresh-water
distribution.”
In this discussion paper, I revisit the question
of salmonid origins, adding an additional, perhaps
peculiar, rider that salmonid origins may be neither
freshwater nor marine.
Diadromy in Salmonidae
Diadromy is a general term introduced to the ichthy-
ological lexicon by American ichthyologist George
Myers (1949). Diadromous fish are those that migrate
on a regularly-timed, consistent, and most often oblig-
atory basis between freshwaters and the sea. Myers
defined several sub-categories of diadromy, including
the already long used terms anadromy and catadromy,
and he coined the new term amphidromy. Each
term covers different sub-categories of diadromy (see
McDowall, 1999 for refined definitions). In general,
any salmonids that are diadromous fall into the sub-
category anadromous (but see Stearley, 1992).
Anadromy has the following general features:
Spawning is in freshwater, and the newly hatched
larvae remain there for periods varying from a
few days to a few months (occasionally a few
years). They feed and grow through some juvenile
phases (varying among species) in freshwater
before they emigrate to sea. Most feeding and
growth take place at sea, usually over several
years. Approaching maturity, the adults return to
freshwater, and penetrate up rivers to breeding
grounds, where they spawn.
A common feature of anadromous fishes is
homing (a return to their natal habitat), to spawn,
a habit that may have synergies with anadromy, in
enabling them to re-locate and exploit historically
productive spawning habitats (McDowall, 2001).
A review of diadromous fishes (McDowall, 1988)
showed that there are about 227 species, of which
110 are anadromous, with 27 of these being salmonids
(including Coregonus, which is sometimes placed in a
separate family). Anadromy is thus a well-represented
behavior amongst salmonids and they constitute a
significant proportion of all known anadromous fishes.
There are many non-anadromous salmonids, and
similarly, many non-anadromous stocks of otherwise
anadromous species. Given these interrelated facts,
and the additional fact that salmonids are amongst the
most important and well-studied of all fish, owing to
their high profile in both commercial and recreational
fisheries, perhaps it is no surprise that the question of
their origins has commanded so much attention.
An historical perspective on opinions regarding
the origins of the Salmonidae
Opinion about salmonid origins, whether marine or
freshwater, date back at least to the early 19th century,
leading Tchernavin (1939, p. 121) to despair after
reading the available literature. A very early English
commentator (Pennant, 1776), is cited by Day (1887)
as apparently having a neutral view, simply stating that
“the salmon is a fish that lives both in the salt and fresh
waters, quitting the sea at certain seasons for the sake
of depositing its spawn in security.” Fleming (1828,
p. 179) is said by Day to have considered the salmon
a “migratory fish from the sea”, perhaps implying a
marine origin, though this is unclear. And Day (1887)
reported Parnell (1838, p. 279) as arguing that “there
is no doubt that the true abode of the salmon is in the
sea, for as soon as it has entered the rivers it begins
to deteriorate in condition.” Day (1887) and Frank
Buckland, according to Day, held contrary views. For
example, Day (1887) cites Buckland (1873) as consid-
ering Atlantic salmon (Salmo salar) “a sea fish proper;
nevertheless, this fish ascends the rivers and streams

in order to deposit its eggs, for unlike other sea fish,
it does not breed in the sea.” But their compatriot,
British Museum ichthyologist Albert Günther (1866,
p. 3) described salmon as a freshwater fish that has
adopted the habit of descending to the sea to feed.
C.T. Regan (1911, p. 24), also at the British Museum
(during the early 1900s), and one of the more notable
of early 20th century ichthyologists, thought that
salmonids “may be regarded as marine fishes [that]
are establishing themselves in fresh water.” Meek
(1916, p. 115) agreed, presenting arguments “why
the evolution [of salmonids] is assumed to be from
marine to fresh-water conditions. . . . The reason . . .
is that the Atlantic salmon and the trouts are more
closely related than the Atlantic and Pacific salmons,
that in other words the salmons preceded the trouts.”
Boulenger (1917), who was at the British Museum
with Regan, also supported a marine origin. Thus,
opposing views amongst contemporary biologists are
evident from early times, although there appears to
have been a minor gathering of “authority” around
a marine ancestry in the late 19th and early 20th
centuries.
Similarly divergent opinions continued into
modern ichthyological history. Unlike many earlier
authorities discussed above, Tchernavin (1939)
decided on a freshwater origin. His analysis of the
marine/freshwater ancestry of salmon (Salmo salar)
is certainly the most cited and carefully developed
perspective from the mid 20th century until recent
times. That being so, it is worth summarising
Tchernavin’s argument:
1. Many genera are wholly freshwater but none is
entirely marine, and the great likeness between
freshwater and migratory forms of salmonids
suggests recent evolution of migratory forms.
2. Those salmonids that are the least marine are
the most primitive, the trout (which Tchernavin
wrongly regarded as non-migratory) being more
primitive than the salmon (migratory).
3. The distribution of salmonids resembles that of
freshwater plants and animals.
4. The migratory salmon expends great effort in
getting into freshwater to spawn, finding condi-
tions congenial for the activity of eggs and sperm,
for fertilisation, and for development and early
life of the young; moreover persistence, genera-
tion after generation, of salmon returning to fresh
water indicates their origins.
173
5. Existence of a homing migration to fresh water
indicates the origin [source] of the migratory
habit.
6. Young Salmonidae show “amazing” resemblance
to the adults of freshwater forms.
Various subsequent commentators have followed
Tchernavin’s view, for various reasons (e.g., Vlad-
ykov, 1963). Hoar (1976, p. 1235) carefully re-
evaluated Tchernavin’s arguments and concluded that
“they still make sense and nothing seems to have been
discovered in the past half century that invalidates
the hypothesis of a freshwater origin.” Hoar (1976,
p. 1236) further argued that additional information on
distribution, morphology, biochemistry, physiology,
and behavior “seem to have added support to most of
[Tchernavin’s] arguments.”
While some found Tchernavin’s arguments
persuasive, Thorpe (1982, p. 88) described them as
“argued forcibly and authoritatively” but (Thorpe,
1988) “with more conviction than evidence” – I
think with some validity. Not one of the points listed
above from Tchernavin (1939) seems, to me, to
indicate a freshwater ancestry. Salmonids are, of
course, well-adapted to spawning and early growth
in fresh water – where they have been spawning and
undergoing early life for millions of years. And why
should what salmonids do now indicate what their
ancient ancestors did before them? Thus, Tchernavin’s
(1939) argument rests substantially on false inference
from contemporary importance to historical priority
(Gould, 1983).
It is also unclear why a migratory life history
should be derived from a non-migratory one, rather
than the reverse. This is especially true when it
is recalled that loss of diadromous migrations has
recurred repeatedly in diverse species of extant
Salmonidae (and other diadromous families). If this
is indicative of what happened in salmonid history,
Tchernavin’s (1939) assumption of migratory species
being recently derived is difficult to accept.
Thorpe (1982) presented an explicit argument in
favour of a marine ancestry, and as such his work
also needs discussion. Thorpe (1982, p. 89) observed
that there is a shift from diadromy (anadromy) to
landlocked/freshwater populations in some salmonid
species, and argued that therefore:
the evolutionary trend in the family appears to
be toward total dependence on fresh water, and
away from dependence on the sea. It is then appro-
priate to view the Salmonidae as relatively prim-
174
itive teleosts of probable marine pelagic origin”
. . . meaning that “. . . the fresh water phases of
the life histories of salmonids are enabled by
specific adaptations, which are superimposed on
a basically marine organism.
Thorpe thus viewed salmonids as evolving away
from a diadromous (anadromous) (marine/freshwater)
life to a purely freshwater one – reflecting Regan’s
(1911) view that salmonids may be regarded as marine
fish establishing themselves in freshwater. Thorpe’s
argument seems, to me, to also be erroneous on at
least two grounds. Firstly, though not necessarily
most importantly, Thorpe’s (1982) argument (as with
Tchernavin’s) rests on false inference from contem-
porary importance to historical priority (Gould, 1983).
Secondly, he appears to be arguing that future evolu-
tionary directions of salmonids were determined by
the direction of their evolutionary past, thus implying
that the evolution of salmonids has momentum and
direction, projecting adaptation forwards. This is an
essentially orthogenetic argument, a form of evolution
that is generally rejected by conventional evolutionary
biologists (Mayr, 1982).
Thorpe (1982) and McCormick and Saunders
(1987) listed Balon (1968) as one contrary author
favouring a marine ancestry. Thorpe (1982: 93)
summarized Balon’s (1980) hypothesis of salmonid
evolution as “evolution of life history strategies
through penetration of fresh water by a pelagic marine
fish, and progressive restriction of life-history to the
freshwater habitat”, but this does not seem to be
Balon’s position. Balon (1980, p. 552) wrote that
“evidence of the evolution of reproductive guilds . . .
seems to support [the] assumption . . . that earlier
forms [of salmonids] were related to the oceanic pela-
gial by their reproductive strategy”, which Thorpe
(1982) took to mean “evolution of salmonid life
history strategies through penetration of fresh water
by a pelagic marine fish.” It seems to me that Thorpe
misinterpreted Balon’s suggestion that freshwater
populations of salmonids evolved from anadromous
populations, as Balon’s support for a marine ancestry.
The geochronological place of diadromous species
in salmonid history
Two distinctly contrasting views have been expressed
regarding when the first diadromous salmonids
evolved. Tchernavin (1939), believing in a fresh-
water origin, argued for development of sea-migratory
stocks of salmonids “during the Glacial period”,
so that “divergence into freshwater and migratory
forms occurred only recently.” Clemens (1953) and
Neave (1958) similarly suggested that Oncorhynchus
diverged from Salmo as recently as Pleistocene glacial
times. The logic for these assertions is, at best,
elusive.
Apart from the fact that the amount of diversific-
ation and geographical spread represented by modern
diadromous salmonids seems intuitively inconsistent
with such a recent origin, these notions are clearly fals-
ified by the fossil record. Cavender and Miller (1972)
described Smilodonichthys rastrosus, a huge Pliocene
salmonid fossil from western North America, (now
included in Oncorhynchus – Stearley and Smith, 1993,
p. 23), and thought this fish diadromous (anadromous)
based on the site from which the fossil came, the
likelihood that it was a pelagic filter feeder (it had
numerous gill rakers), its large premaxillary teeth, and
internal organisation of the dermal bone (cavernulous
providing potential storage of fats – p. 37).
Cavender and Miller (1972, p. 39) also reported
“Fragmentary remains of what appears to be an
extinct species of Oncorhynchus” from western North
America (of Pliocene age), leading them to conclude
that the genus is “at least as old as middle Plio-
cene.” Moreover, Stearley (1993, p. 23) cited several
other Tertiary fossils, the most telling of which (with
regard to a supposed Pleistocene origin of diadromous
salmonids), being a Miocene fossil of chum salmon,
Oncorhynchus keta, from western North America.
Stearley and Smith (1993, p. 23) found that “these
fossils of Oncorhynchus document abundant evidence
for a minimum age of 6 million years for the modern
species of Pacific salmons and trouts”, and accept-
ance of this age certainly lays to rest any notion
that a Pleistocene diversification produced these taxa.
There seems good evidence for probably anadromous
salmonids as early as the Miocene, far earlier than
speculated by Tchernavin (1939) and those who have
followed him.
Diadromy and phylogenetic analysis
Diadromy is a behavioral trait, and there is a view that,
as behavioral traits tend to be both non-quantitative
and malleable, they are of dubious value in phylogen-
etic analysis. However a converse view has prevailing
acceptance, viz. that when applied with care, behavi-
oral traits can be used in phylogenetic analysis with

useful results (Dobson, 1985; McLennan et al., 1988;
de Quieroz and Wimberger, 1993; Paterson et al.,
1995; McDowall, 1997).
Diadromy poses problems for phylogenetic
analysis, particularly though not exclusively amongst
salmonids. For example, many diadromous salmonids
also have non-diadromous populations – landlocked
populations establish and forsake migrations to
and from the sea, sometimes replacing these
with migrations to and from lakes. Perhaps best
known are kokanee populations of sockeye salmon
(Oncorhynchus nerka) in western North America
(Foerster, 1968; Foote et al., 1994; Wood and
Foote, 1996), but there are numerous other examples
amongst salmonids (and elsewhere amongst the
related Osmeroidei – McDowall, 1988). Rounsefell
(1958), for instance, actually ranked salmonids of
western North America according to the degree to
which their diadromy is facultative.
It is known that in some normally diadromous
species, a small proportion of individual fish may
be non-diadromous (they fail to migrate to sea),
though they may participate in reproduction with
their diadromous con-specifics at spawning time. This
may occur in both sexes, but is reported especially
for males of species of Salmo, Oncorhynchus and
Salvelinus (Balon, 1968; Saunders et al., 1982; Silver-
stein and Hershberger, 1992; Jonsson and Jonsson,
1993; Unwin et al., 1999; Healey et al., 2000).
The presence of these few non-migratory indi-
viduals among the many became evident when
several diadromous salmonids were introduced to
New Zealand in the late 1800s and early 1900s.
Diadromous stocks of sockeye salmon, Atlantic
salmon, and rainbow trout (O. mykiss) were trans-
located to New Zealand, but in all three instances
the outcome was acclimatization of entirely non-
migratory stocks. There must have been some indi-
viduals amongst the translocated fish that had a genetic
tendency to not migrate, and these formed the ancestry
of populations that became established (McDowall,
1990, 1994). Instances are known of latitudinal vari-
ation in the extent to which populations of a species
are diadromous, typically with a greater tendency for
diadromy at higher (cooler latitudes), as with intro-
duced brown trout (Salmo trutta) in New Zealand
(McDowall, 1990).
Conversely, there are records of non-diadromous
populations of diadromous species being translocated
to new locations and reverting to diadromy, indic-
ating that some individuals in such non-migratory
175
populations have retained the instinct to migrate to
sea (Northcote, 1992; Jonsson and Jonsson, 1993).
Nordeng (1983) has shown this for Arctic char
(Salvelinus alpinus) and other instances include brown
trout when non-migratory stocks were translocated in
Europe (Thorpe, 1987), and when introduced to the
Falkland Islands (Nolan, 1993).
With all this within-species variability, it appears
that diadromy is a trait of limited utility in phylo-
genetic analysis. Despite this, one notable aspect of
diadromy is that it has evolved relatively rarely across
the diversity of fishes. Diadromy is known in only 36
of 482 families of fishes (McDowall, 1988; Nelson,
1994). Most diadromous fish species are found in only
a few families – in the listing of McDowall (1988),
there are 11 sturgeons, 27 salmonids, 11 salangids,
30 clupeids, 15 anguillid eels, 15 mugilid mullets,
64 gobies. Thus 173 of the 227 species, or 76% of
species listed in McDowall (1988) are in only seven
families. Diadromy may have evolved several times
in some of these families but, even so, it is clear
that diadromy is a trait that has evolved relatively
rarely. This rarity, together with variability in the pres-
ence/absence of diadromy within species discussed
above, suggests that, at best, diadromy may be of
phylogenetic interest when comparing higher taxa that
do, or do not, exhibit diadromy. The evolution of
diadromy may be a novel and phylogenetically signifi-
cant event, and its presence in two or more species in
a family that is otherwise non-diadromous may hint at
relationships of those where it is present. But, owing
to within and between taxon variability, differences in
the nature of the diadromy (or its presence/absence
in species being compared) seem of minimal phylo-
genetic value when exploring the relationships of
congeneric or confamilial species.
Origin of Salmonidae in a cladistic context
Few of the authors discussed earlier have examined
the question of salmonid relationships by identifying
common derived characters, i.e., from a cladistic
perspective. Stearley and Smith (1993, p. 5) recog-
nised that Regan (1914) used “shared specializa-
tions as evidence of relationship as later formalized
by Hennig (1966)”, whereas (p. 4) they found that
Tchernavin (1939) used “shared primitive characters
for diagnosing genera”, which provide no indica-
tions of relationships. Thus, virtually all early studies
searched for characters that indicate a marine or fresh-
176
water ancestry using “indirect” methods (Stearley,
1992, p. 622), the characters having no obvious indic-
ation of why a marine or freshwater origin should
be indicated – there is no reason why contemporary
character states should indicate which of these is
true, as Tchernavin (1939, p. 127) admitted when
noting that “the distribution of an animal does not
always follow the milieu inhabited by its ancestors”
(though he appears to have taken no notice of his
own warning). For example, it is widely suggested
that because salmonid reproductive products survive
and result in fertilisation only in freshwater, and
that because egg development takes place there, a
freshwater origin is indicated (Tchernavin, 1939;
McDowall, 1988), and this argument looks good,
as long as you don’t look too carefully. Of course
salmonid eggs and sperm function best in freshwater
if this is where salmonid reproduction has taken
place for the past several (many?) millions of years,
and this scarcely argues for a freshwater origin at
some long-past time. The present character state does
not prevent some change of reproductive medium in
the past, the possibility of which is explicitly indi-
cated by the fact that although most species in the
related family Osmeridae spawn and develop in fresh-
water (like Salmonidae), three species of osmerid,
in three different genera spawn and develop at sea.
Two of these genera (Hypomesus and Spirinchus) have
both marine and freshwater spawning species (McAl-
lister, 1963). Given the varied spawning media in
Osmeridae, assumptions that Salmonidae originated
in freshwater because they spawn and develop there
are fraught with risk. Unlike these marine osmerids,
no salmonids have adopted entirely marine life history
strategies, pink salmon (Oncorhynchus gorbuscha)
being closest with their spawning, sometimes within
tidal influence in low elevation rivers (Heard, 1991).
Stearley (1992) addressed the origin of the
subfamily Salmoninae in an explicitly cladistic
analysis of the constituent genera and species – he
considered (p. 622) that a “good understanding of
salmonid phylogenetic relationships is of extreme
import for understanding the evolution of their life
histories”, hoping that, structured upon a proper
framework, “the sequence of branching relationships
of taxa . . . a sequential elaboration of salmonid beha-
viors may be discernible.” I agree.
Stearley (1992, p. 625) stated that he constructed
his phylogenetic relationship using (amongst other
taxa) the osmerid genera Thaleichthys and Spirin-
chus as outgroups (Stearley and Smith, 1993, p. 7,
specified only Thaleichthys), since these are regarded,
by some, as the most basal (plesiomorphic) of
osmerids. He addressed the “span of morphology
defining ancestral states for salmonid characters”, and
used the phylogeny generated for interpreting the
history/evolution of diadromous migratory strategies
in the subfamily. He argued that in Salmoninae, in
particular, a series of non sea-migratory genera such
as Brachymystax, Acantholingua, Salmothymus and
Platysalmo are more plesiomorphic sister-groups of
diadromous (anadromous) apomorphic genera such as
Salmo, Salvelinus and Oncorhynchus. On this basis,
Stearley (1992) argued for a freshwater origin of
salmonines because the genera listed above as “prim-
itive” are non-migratory. Greenwood (1993, p. 374)
described Stearley’s (1992) account as a “new and
convincing solution to the old problem of marine
versus freshwater ancestry for salmonoids”, though
in fact Stearley (1992, p. 623) wrote explicitly of
his “focus on the life histories of the salmonine
subfamily”, which is rather different (emphasis added
in both quotes).
Relationships of salmonids in the context of
variability of diadromous life cycles
Although Stearley (1992) anchors his morpholo-
gical phylogenetic analysis in outgroups, rather than
carrying this approach through into comparable iden-
tification of outgroup character states in his behavioral
analysis, he uses the phylogenetic relationships to
determine patterns of behavioral evolution amongst
the taxa. This forms the basis for his conclusion
that salmonines have a freshwater ancestry. Stearley’s
(1992) argument for a freshwater origin of salmonines
is suspect for several reasons:
1. As discussed above, diadromy is a notori-
ously variable or reversible character trait within
species, and that being so, specification of non-
diadromy in salmonines as plesiomorphic and
diadromy apomorphic is unjustified.
2. Stearley (1992, p. 627) argues that “Primitive
salmonid sistergroups – Prosopium, Thymallus
and the archaic trouts [those listed above
as the primitive salmonines] are restricted to
freshwaters”, whereas “Advanced salmonine
and coregonine clades – Coregonus, Stenodus,
Salvelinus, Hucho, Salmo and Oncorhynchus
all contain sea-run species”. He regarded non-
migratory Salvelinus and Hucho species as

“atavistic, reverting to the primitive (strictly
freshwater) condition possessed by more prim-
itive salmonine sister groups.” But there is
no reason why the primitive salmonine sister
groups (Brachymystax, Acantholingua, Salmo-
thymus and Platysalmo) should not also have
lost their formerly diadromous behaviors just
like these Salvelinus and Hucho species have (as
also have various species of Salmo and Onco-
rhynchus); Stearley (1992, p. 629) himself notes
that “Spontaneous abandonment of migratory life
cycles appears to occur regularly” in salmonines.
So, it is unclear why should this not have
occurred in the listed primitive salmonines whose
lack of diadromy is interpreted by Stearley as
plesiomorphic and indicating a freshwater origin
in salmonines.
3. In spite of the above proposed set of relation-
ships, looking more widely at the Salmonidae
(i.e., including coregonines and thymallines),
according to Stearley and Smith (1993) the most
plesiomorphic genera in the family are, Core-
gonus, Stenodus and Prosopium; species of Core-
gonus and Stenodus are diadromous (McDowall,
1988), so that diadromy is present in these
plesiomorphic genera, creating a conflict with
Stearley’s (1992) conclusion that absence of
diadromy in the most primitive salmonines indi-
cates a freshwater ancestry for salmonines.
4. Perhaps more importantly than any of the above,
relationships of the Salmonidae, and the signal
indicated about the family’s origins (marine
or freshwater), must be viewed in an even
broader context. Specifically, although details are
debated, there is quite wide and long-standing
acceptance that Salmonidae are a sister-group
of Osmeroidei+Galaxioidei (McDowall, 1969;
Rosen, 1974; Fink and Weitzman, 1982; Johnson
and Patterson, 1996 – family assignments and
arrangements vary, but I think the taxa involved
in these relationships are quite clearly under-
stood). That this view is accepted, at least in
part, by Stearley (1992) and Stearley and Smith’s
(1993) is shown by their choice of the osmerids
Spirinchus and Thaleichthys as being amongst the
outgroups for their salmonid phylogenetics.
If it is accepted that [Osmeroidei+Galaxioidei] is
the sister group of Salmonidae, then the life history
character states within the [Osmeroidei+Galaxioidei]
ought to be informative of the ancestral condi-
tion. Diadromy is widely recognised within osmeroid
177
and galaxioid families, including most species
of Osmeridae (including Plecoglossus), Salangidae
(both Osmeroidei) and Retropinnidae (including
Prototroctes) and Galaxiidae (including Aplochiton
and Lovettia) (both Galaxioidei), i.e., diadromy
is widely present in a recognised sister-group
of Salmonidae, suggesting that that diadromy
was a characteristic of the group/species ancestral
to the entire [Salmonidae+Osmeroidei+Galaxioidei]
complex. That being so, there are no grounds
for either regarding non-diadromous salmonids as
plesiomorphic (indicating the ancestral condition), or
for regarding the origins of Salmonidae as being in
fresh water (recall that although most of these taxa
are freshwater spawners, three osmerids are marine
spawners).
Conclusion
Prolonged debate has centered around whether the
Salmonidae had a marine or freshwater ancestry, with
strong arguments presented in favour of each. If it is
accepted that Osmeroidei+Galaxioidei are the sister-
group of Salmonidae, however, given the widespread
presence of diadromy in Osmeroidei+Galaxioidei,
then it has to be concluded that salmonids had a neither
marine nor freshwater origin, but are part of a diverse
group of fishes in which diadromy is an ancestral char-
acter state (McDowall, 1993). This conclusion, of
course, begs the question of whether the broader group
comprising Salmonidae+Osmeroidei+Galaxioidei had
a marine or freshwater origin. I know of no way this
broader question can be answered with any assurance
or finality. Flexibility and reversibility in the pres-
ence/absence of diadromy mean that we may never
know whether Salmonidae+Osmeroidei+Galaxioidei
had a marine or freshwater ancestry. Fyhn et al. (1999)
have explored origins of teleost fishes as a whole,
based on views that the radiation of teleosts began
in fresh water. They suggested that the “first trials
of colonising the saline ocean was by way of marine
excursions into the sea for feeding, and return to fresh
water for spawning.” As they point out, anadromous
fish “such as the salmonids still show this behavior.”
Perhaps this is a pointer to a freshwater ancestry of the
entire osmeroid/galaxioid/salmonid complex of fishes.
On the other hand, it might be argued that fishes of
marine origin invaded freshwater (and there are plenty
of precedents for this across the diversity of fishes),
found this a fruitful biome in which to live, and so
178
radiated and expanded to produce the diversity that we
see today.
Acknowledgements
I am grateful to John Thorpe, University of Glasgow,
Scotland, Eugene Balon, University of Guelph,
Guelph, Ontario, Canada, Don Jellyman, NIWA,
Christchurch, and also Gerald Smith, University of
Michigan, Ann Arbour, Michigan, U.S.A., and a
second anonymous referee, for thoughtful comment
on this paper.
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