LIFE HISTORY PATTERNS AND
BIOGEOGRAPHY: AN
INTERPRETATION OF DIADROMY
IN FISHES1
Lynne R. Parenti2
ABSTRACT
Diadromy, broadly defined here as the regular movement between freshwater and marine habitats at some time during their
lives, characterizes numerous fish and invertebrate taxa. Explanations for the evolution of diadromy have focused on ecological
requirements of individual taxa, rarely reflecting a comparative, phylogenetic component. When incorporated into
phylogenetic studies, center of origin hypotheses have been used to infer dispersal routes. The occurrence and distribution of
diadromy throughout fish (aquatic non-tetrapod vertebrate) phylogeny are used here to interpret the evolution of this life
history pattern and demonstrate the relationship between life history and ecology in cladistic biogeography. Cladistic
biogeography has been mischaracterized as rejecting ecology. On the contrary, cladistic biogeography has been explicit in
interpreting ecology or life history patterns within the broader framework of phylogenetic patterns. Today, in inferred ancient
life history patterns, such as diadromy, we see remnants of previously broader distribution patterns, such as antitropicality or
bipolarity, that spanned both marine and freshwater habitats. Biogeographic regions that span ocean basins and incorporate
ocean margins better explain the relationship among diadromy, its evolution, and its distribution than do biogeographic regions
centered on continents.
Key words: Antitropical distributions, biogeography, diadromy, eels, fishes, global biogeographic regions, life history,
migration.
?? The extremely slow growth of the larvae of the
European eel is. . . an adaptation to the prolonged
journey [to spawning grounds in the Sargasso Sea]. It is
scarcely possible to understand this unique phase in the
life cycle of the European eel on the hypothesis that the
geographical conditions were formerly the same as now
exist. But if Wegener?s theory [of continental drift] be
accepted, the explanation is simple. As the coasts slowly
receded from one another the larval life of what became
the European species was more and more prolonged by
natural selection in correspondence with the greater
distance to be traversed.??
?Fulton (1923: 360)
Animals that migrate between marine and freshwa-
ter at some time during their life are diadromous, a
term coined by American ichthyologist George S.
Myers (1949a). Several categories of diadromy were
defined or redefined (Myers, 1949a, b; McDowall,
1988)?anadromy, catadromy, and amphidromy?
collectively referred to as diadromy herein. Some
invertebrate taxa, such as atyid crustaceans and
neritid gastropods, are also diadromous (e.g., Myers
et al., 2000).
Thomas Wemyss Fulton (1855?1929) was a
Scottish fisheries biologist best remembered today
for his classic The Sovereignty of the Sea (Fulton,
1911), a treatise on the development of the notion of
territorial waters. Fulton?s life began in 1855, a few
years before Charles Darwin?s publication of On the
Origin of Species in 1859, and ended in 1929, just as
Alfred L. Wegener published the fourth edition of Die
Entstehung der Kontinente und Ozeane, the proposal of
a theory of continental drift. In his letter to Nature,
quoted above, Fulton described a hypothesis for the
evolution of catadromy in the European eel, Anguilla
anguilla (L.), which incorporated modern concepts of
panbiogeography. First, the earth and its biota evolved
together, a tenet of botanist Le?on Croizat (1958).
Second, global biogeographic regions should be
centered on ocean basins, not continents (Croizat,
1958; Craw et al., 1999: figs. 6?13). Third, following
from the first two, migration patterns evolve in concert
with geology, and these changes lead to lineage
differentiation, here changes in length of larval
period, and ultimately to what we recognize as
1 I thank Richard L. Mayden, Peter H. Raven, and P. Mick Richardson for the invitation to present this paper in the 52nd
Annual Systematics Symposium of the Missouri Botanical Garden, ??Reconstructing Complex Evolutionary Histories: Gene-
species Trees, Historical Biogeography, and Coevolution,?? and colleagues at the Symposium, in particular Kevin Conway, for
discussion. John R. Grehan and James C. Tyler provided references. Malte C. Ebach, Anthony C. Gill, David G. Smith, and
Victor G. Springer read and commented on a draft of the manuscript and provided references.
2 Division of Fishes, Department of Vertebrate Zoology, Smithsonian Institution, P.O. Box 37012, National Museum of
Natural History, Room WG-12, MRC 159, Washington, D.C. 20013-7012, U.S.A. parentil@si.edu.
doi: 10.3417/2006051
ANN. MISSOURI BOT. GARD. 95: 232?247. PUBLISHED ON 18 JUNE 2008.
speciation. When Fulton referred to ??. . . what became
the European species. . . ,?? he postulated that there
had been a widespread northern Atlantic lineage that,
along with geological evolution of the Atlantic Ocean,
differentiated to become the European eel in the
eastern North Atlantic and the American eel, A.
rostrata (Lesueur), in the western North Atlantic.
Anguilla anguilla and A. rostrata are hypothesized
sister lineages (Lin et al., 2001a, 2005; see below).
Other early 20th-century biologists adopted earth
history to explain the evolution of migration patterns.
Notable among ornithologists was Albert Wolfson of
Northwestern University, Illinois, who, in 1948, wrote
a lead article in Science, ??Bird Migration and the
Concept of Continental Drift,?? which correlated the
increasing distances birds traveled with the separation
of land masses and their latitudinal shift. Nearly four
decades later, Wolfson (1986) vividly recounted the
negative and positive reaction to his correlation of
bird migration and continental drift.
Many modern explanations for the evolution of
migration and diadromy conflict with those of Fulton
and Wolfson because, for the most part, they reject the
coupled evolution of life and earth, at global and local
scales, and rely exclusively on ecological and
evolutionary models. For example ??. . . the relative
productivity or growth advantage of sea and freshwater
habitats appears to be the key to [diadromy?s]
evolution. The productivity differential can probably
explain why fish migrate across the sea?freshwater
boundary, predict their direction of movement and
account for where in the world diadromous species
occur?? (Gross, 1987: 21). Earth history was separated
from biological history, in part as a return to center of
origin explanations for geographic distributions as
outlined by Matthew (1915) in his influential, often
reprinted ??Climate and Evolution?? (see Nelson &
Ladiges, 2001). Equally influential was Myers, who
classified fishes in ecological categories for the
purposes of biogeography. A decade before he defined
diadromy, Myers (1938) proposed divisions of fresh-
water fish taxa based on their ability to tolerate
saltwater: freshwater fishes, including primary divi-
sion freshwater fishes (no salt tolerance), secondary
division freshwater fishes (some salt tolerance), and
peripheral or migratory fishes (diadromous) versus
marine fishes (salt tolerant). These categories have
been incorporated into fish biogeography and are still
used broadly today (e.g., McDowall, 1988; Helfman et
al., 1997; Berra, 2001). (The category of primary
division freshwater fishes was modified by paleontol-
ogist Colin Patterson [1975], who recognized archae-
olimnic taxa, those inferred to have originated and
always lived in freshwaters, and telolimnic, those
living in freshwater now, but not necessarily through-
out their history. These two categories have been used
almost exclusively by paleoichthyologists [e.g., Hilton,
2003].) Following Myers?s ecological classification,
the distribution of primary division freshwater fishes
has been interpreted with respect to earth history (viz.
Darlington, 1957), whereas the distribution of sec-
ondary division freshwater fishes, peripheral fishes,
and marine fishes has largely not (e.g., Briggs, 1974,
1995; but see Springer, 1982; Parenti, 1991; Mooi &
Gill, 2002; Nelson, 2004).
Accepting Myers?s ecological categories precludes
the comparison of distribution patterns among dia-
dromous, freshwater, and marine fishes: ??It is plainly
evident that a fish which can swim through sea water
from one river mouth to another is not of much use in
studies of terrestrial zoogeography?? (Myers, 1938:
343). And so it was believed by Myers and
contemporaries (e.g., Herre, 1940) and subsequent
generations of biogeographers. Adhering to Myers?s
philosophy, biogeography of marine and freshwater
fishes on a global scale remains largely independent.
Textbooks and courses have traditionally drawn the
line, contrasting Zoogeography of the Sea (Ekman,
1953) and Marine Zoogeography (Briggs, 1974) with
Zoogeography of Fresh Waters (Ba?na?rescu, 1990,
1992, 1995), for example. More general texts, such
as Zoogeography: The Geographical Distribution of
Animals (Darlington, 1957), Global Biogeography
(Briggs, 1995), or Biogeography, an Ecological and
Evolutionary Approach (Cox & Moore, 2005), likewise
have divided their attention between marine and
freshwater taxa.
The arguments for dispersal (e.g., Gross, 1987;
McDowall, 1970, 1988, 2001, 2002; Berra et al.,
1996; Keith, 2003), vicariance (e.g., Rosen, 1974;
Croizat et al., 1974), or some combination of the two
(e.g., Choudhury & Dick, 1998) as explanations for
the distribution of diadromous taxa, especially the
antitropical salmoniform fishes, have been debated
extensively. Likewise, broad geographic ranges have
been interpreted as explicit evidence of dispersal
(e.g., McDowall, 2001, 2002), of vicariance (e.g.,
Rosen, 1974), or of either dispersal or vicariance
(Leis, 1984). Fulton?s explanation for Anguilla
Schrank distribution (above) will not convince a
dispersalist to embrace vicariance or a vicariance
biogeographer to adopt dispersalist explanations, as
each represents a particular view of the relationship
between the organism and the environment in forming
biogeographic distributions.
Cladistic biogeography (viz. Nelson & Platnick,
1981; Humphries & Parenti, 1986, 1999; Crisci et al.,
2003) puts primary emphasis on the phylogenetic
relationships among organisms, not their physiological
or ecological requirements, to discover distribution
Volume 95, Number 2 Parenti 233
2008 Life History Patterns and Biogeography
patterns. Cladistic biogeographers do not ignore
ecology (viz. Wiens & Donoghue, 2004), but, rather,
are explicit in interpreting ecology or life history
patterns within the broader framework of phylogenetic
patterns (e.g., Sparks & Smith, 2005). Detailed
reviews and commentaries on the systematics, biology,
and distribution of diadromous fishes, largely within
the dispersalist framework, have been provided by
New Zealand ichthyologist Robert M. McDowall (e.g.,
1970, 1987, 1988, 1992, 1993, 1997a, b, 2001, 2002,
2003). I have argued previously that the distribution
of each genus of diadromous gobies of the subfamily
Sicydiinae (family Gobiidae) may be described by the
ocean basins in which it lives and, further, allopatry
may be recognized at the generic level: the eastern
Pacific and Atlantic Sicydium Valenciennes does not
overlap with its sister genus Sicyopterus Gill in the
Indian Ocean and western and central Pacific
(Parenti, 1991; Parenti & Thomas, 1998).
My purpose here is to reinterpret the phylogeny and
global distribution of fishes, both freshwater and
marine, within a cladistic biogeographic framework to
ask if, and if so, how, the biology and ecology of
diadromous fishes are related to their distribution
patterns and to competing models of earth history.
DIADROMY AND CLASSIFICATION OF FISHES
Table 1 lists the major groups of living aquatic
vertebrates that we call fishes, the estimated number
of species in each taxon, the number of species that
live almost exclusively in freshwater, and the total
number of species that enter freshwater during some
phase of their lives (modified from Nelson, 2006: 4?
5). Diadromous species are included in the last
column, which also enumerates those species that do
not undergo regular habitat migrations but are
euryhaline, such as cyprinodontiform killifishes (see,
e.g., Rosen, 1973). An estimated 225 species are
diadromous (McDowall, 1992; Nelson, 2006). Not all
diadromous species are obligately so (McDowall,
2001), precluding a precise enumeration of such taxa.
A large proportion of diadromous species is
concentrated among inferred basal lineages: lampreys
(Petromyzontiformes), sturgeons (Acipenseriformes),
eels (Elopomorpha), herring (Clupeomorpha), and
salmon (Protacanthopterygii), as noted by McDowall
(1988, 1993) and others, and many species in these
lineages are euryhaline. Many, but not all, diadromous
fishes are also antitropical or bipolar, distributed in
the northern boreal and/or the southern austral zone,
and absent from the tropics, again as noted by
McDowall (1988, 1993) and others. Salmoniforms and
osmeriforms (Protacanthopterygii; Rosen, 1974: fig.
45), lampreys (Petromyzontiformes; Berra, 2001: 6, 8,
12) and some clupeomorphs (Nelson, 1986) have
bipolar distributions, whereas sturgeons (Acipenser-
idae; Choudhury & Dick, 1998; Berra, 2001: 42) and
sticklebacks (Gasterosteidae; Berra, 2001: 351) are
boreal, for example.
EPICONTINENTAL SEAS: THE SETTING FOR BONY
FISH EVOLUTION
Living actinopterygian (ray-finned) fish clades are
the bichirs (Polypteriformes), sturgeons and paddle-
fishes (Acipenseriformes), gars (Lepisosteiformes),
bowfins (Amiiformes), and the bony fishes (Teleostei;
see Nelson, 2006). Relationships among these line-
ages have been debated by both morphologists and
molecular biologists (e.g., Nelson, 1969a; Patterson,
1973; Wiley & Schultze, 1984; Inoue et al., 2003)
and, for the purposes of this discussion, may be
summarized by the cladogram of Figure 1.
Polypteriformes (e.g., Britz, 2004) live in African
freshwaters. Fossil bichirs of Africa and South
America are of Middle Cretaceous age (Nelson, 2006).
Acipenseriformes (Grande & Bemis, 1991) com-
prise the anadromous and freshwater circumboreal
sturgeons, Acipenseridae, and the paddlefishes, Poly-
odontidae. There are two extant species of paddlefish,
one in freshwater in North America and the other in
freshwater and estuarine habitats in the Yangtze River
and associated areas of China. Fossil paddlefishes
date to the Lower Cretaceous from China (Berra,
2001). Sturgeons are an old lineage, known from at
least the Middle Jurassic (175 million years ago [Ma];
Choudhury & Dick, 1998).
Lepisosteiformes includes one living family, Lepi-
sosteidae (Wiley, 1976), which comprises two extant
genera with seven species living in fresh, brackish, or
marine waters in North and Central America and
Cuba. Fossil gars are more broadly distributed
throughout North and South America, Europe, Africa,
and India (Wiley, 1976: fig. 69); they date from at
least the Early Cretaceous (Janvier, 2007).
Amiiformes (Grande & Bemis, 1998, 1999) are
represented by one living species, Amia calva L.,
distributed in freshwater habitats in North America. It is
the single, living representative of the Halecomorphi,
known from marine and fossil taxa dating to the Jurassic.
The ancestral habitat for bony fishes is inferred
from the phylogeny and distribution of living and
fossil basal actinopterygian taxa to be shallow
epicontinental seas and their freshwater margins,
including river deltas (Fig. 1). A reconstructed
widespread ancestral range was illustrated for a
subgroup of halecomorph fishes by Grande and Bemis
(1999: fig. 6), who outlined the distribution on a map
of present-day geography contrasted with that of the
234 Annals of the
Missouri Botanical Garden
Early Cretaceous, 118 Ma. The present-day, disjunct
transatlantic distributions are continuous when drawn
across Cretaceous epicontinental seaways.
An area cladogram, or areagram, of halecomorphs
indicating habitat, marine or freshwater (Fig. 2A), was
used by Grande and Bemis (1999: fig. 7) to infer a
marine ancestry for the group. This hypothesis results
from optimizing habitat or, at the least, inferring that
the habitat of the basal taxon is the ancestral habitat.
It rests on the assumptions that the habitat has
changed (e.g., marine to freshwater), even when there
may be no evidence for such a transformation, and
that one or the other taxon of a sister group pair may
be identified as basal (see Parenti, 2006; Santos,
2007; also see below). Without these assumptions, the
ancestral habitat may be reconstructed as epiconti-
nental seas, spanning marine and freshwater habitats
(Fig. 2B). Likewise, bichirs in South America and
Africa also represent a transatlantic distribution, that
is, a remnant of the broader distribution pattern.
Table 1. Classification of living aquatic vertebrates (non-tetrapods) or fishes, estimated number of species in each taxon,
number of those that live almost exclusively in freshwater, and total number of those that enter freshwater at some time during
their life (from Nelson, 2006: 4?5; Stiassny et al., 2004). The last column includes all of the species in the second column, plus
those that are migratory (diadromous or euryhaline).
Taxon
No. of
species
No. of
freshwater species
Total no. of species
that enter freshwater
Myxiniformes 70 0 0
Petromyzontiformes 38 29 38
Elasmobranchii 937 24 40
Chimaeriformes 33 0 0
Osteichthyes
Sarcopterygii
Coelacanthiformes 2 0 0
Ceratodontiformes 6 6 6
Actinopterygii
Polypteriformes 16 16 16
Acipenseriformes 27 14 27
Lepisosteiformes 7 6 7
Amiiformes 1 1 1
Teleostei
Osteoglossomorpha 220 220 220
Elopomorpha 857 6 33
Clupeomorpha 364 79 85
Ostariophysi 7980 7847 7858
Protacanthopterygii 356 127 152
Esociformes 10 10 10
Stomiiformes 391 0 0
Ateleopodiformes 12 0 0
Aulopiformes 236 0 0
Myctophiformes 246 0 0
Lampridiformes 21 0 0
Polymixiiformes 10 0 0
Paracanthopterygii 1340 21 24
Stephanoberyciformes 75 0 0
Zeiformes 32 0 0
Beryciformes 144 0 0
Gasterosteiformes 278 21 43
Synbranchiformes 99 96 99
Perciformes 10,033 2040 2335
Mugiliformes 72 1 7
Atherinomorpha 1552 1304 1352
Scorpaeniformes 1477 60 62
Pleuronectiformes 678 10 20
Tetraodontiformes 357 14 22
TOTALS 27,977 11,952 12,457
Volume 95, Number 2 Parenti 235
2008 Life History Patterns and Biogeography
Paddlefishes reflect transpacific relationships, as do
other North American taxa (Grande, 1994a, b).
A lineage with marine and freshwater representa-
tives (Fig. 3A) may be interpreted to have a marine
and freshwater ancestral distribution (Fig. 3B). In a
likelihood approach to reconstructing ancestral ranges
of lineages, Ree et al. (2005: fig. 6c) also reconstruct a
widespread ancestral range equivalent to that of
Figure 3B when dispersal is considered unlikely, that
is, when no a priori process is specified to explain the
pattern (see also Ebach, 1999; Santos, 2007).
??Ecologically variable ancestors?? are analogous to
??geographically widespread?? ancestors (see Hardy,
2006: 13). (The method of Ree et al. [2005] does not
correspond to other cladistic biogeographic methods,
in part because it defines an area as a discrete
geographic unit, not the organic areas of endemism
that are defined by the ranges of the organisms that
inhabit them [see Harold & Mooi, 1994].)
Our notion of freshwater and marine fishes comes,
in large part, from the existence of speciose taxa that
are today nearly all freshwater, such as the Ostar-
iophysi (catfish, characins, minnows, and relatives) or
Osteoglossomorpha (bony tongues), or from deep-sea
marine orders, such as the Myxiniformes (hagfishes),
Chimaeriformes (chimaeras), Stomiiformes (dragon-
fishes), Aulopiformes (lizardfishes), Myctophiformes
(lanternfishes), and so on (Table 1). Osteoglossomor-
pha is the sister group to all other teleosts, all extant
species of which live exclusively in freshwater
(Nelson, 1969b, 2006; Table 1). It is not historically
a freshwater lineage: marine Paleocene-Eocene fos-
sils, such as the genus {Brychaetus Agassiz, are
hypothesized to be nested in the osteoglossomorph
cladogram in the family Osteoglossidae (Hilton,
2003). The oldest osteoglossomorph fossils are Middle
Mesozoic, and Ba?na?rescu (1995: 1162) acknowledged
their long history and widespread ancestral distribu-
tion by proposing that Osteoglossomorpha is Pangean.
Osteoglossomorpha, therefore, has both marine and
freshwater representatives and is a freshwater group
today because of widespread extinction of marine
taxa. If the majority of species in a taxon are marine
and just a handful are freshwater, it is often assumed
that the group originated in marine habitats and
several taxa dispersed to freshwater (e.g., Myers,
1938; Berra, 2001; Tsukamoto et al., 2002). The
reverse is also assumed. Such an assumption is not an
analysis of biogeographic data, but an implicit and
untestable center of origin hypothesis (viz. Croizat et
al., 1974).
Both Nelson (1973) and Patterson (1975) optimized
habitat on areagrams to infer habitat of origin of
osteoglossomorphs and came to different conclusions.
For example, Patterson (1975: 162) asserted that
??. . . if the conventional view of the history of
Figure 1. Hypothesized relationships among living actinopterygian clades, following Inoue et al. (2003) and Grande and
Bemis (1999). The ancestral distribution is inferred to be throughout epicontinental seas and their freshwater margins.
236 Annals of the
Missouri Botanical Garden
Figure 2. ?A. Areagram of halecomorph habitats from Grande and Bemis (1999), who inferred a marine ancestry for the
group. M/FW means distribution in marine and freshwater habitats; M (FW) means distribution mostly in marine habitats with
some representatives in freshwater. ?B. Reconstruction of nodes to infer a widespread marine/freshwater ancestral habitat,
interpreted here as epicontinental seas and their freshwater margins.
Volume 95, Number 2 Parenti 237
2008 Life History Patterns and Biogeography
Gondwana is accepted, marine origin of the Osteo-
glossomorpha is indicated.?? Nelson (1973: 9), who
read Patterson in manuscript, countered that ??. . . the
totality of evidence concerning the relationships and
distribution of osteoglossomorphs, both fossil and
Recent, indicates that {Brychaetus is secondarily
marine,?? that is, that osteoglossomorphs originated in
freshwaters.
Even though Nelson later dismissed such arguments
(e.g., Nelson, 1974; Nelson & Ladiges, 2001), they
persist (see Parenti, 2006). Optimization of habitat on
an areagram has been applied broadly to interpret the
ecological and distributional history of lineages (see,
e.g., Brooks & McLennan, 1991; McLennan, 1994).
Optimization of habitat is often justified by the
principle of parsimony, yet, in practice, can require
Figure 3. Hypothetical areagram representing taxa in marine (M) and freshwater (FW) habitats. ?A. Optimization of
nodes to infer a marine center of origin. ?B. Reconstruction of nodes to infer a widespread marine/freshwater ancestral
habitat.
238 Annals of the
Missouri Botanical Garden
multiple hypotheses of switching between habitats
(e.g., Baker, 1978). The distributional history of gobioid
fishes, for example, was hypothesized by Thacker and
Hardman (2005: 869): ??. . . gobioids arose in freshwater,
from a marine ancestor, then returned to marine
habitats once or many times.??
Whether diadromy or euryhalinity is a primitive or
derived life history pattern for fishes has been debated
seemingly endlessly (Tchernavin, 1939; Denison,
1956; Patterson, 1975; Griffith, 1987; McDowall,
1988, 1993, 1997a; Johnson & Patterson, 1996; Bemis
& Kynard, 1997; Waters et al., 2000; see especially
review in Janvier, 2007). The distribution of fishes
throughout epicontinental seas, including along their
margins, as proposed for basal actinopterygians, is in
accord with numerous explanations for the evolution of
solely freshwater or marine taxa from widespread
diadromous or euryhaline ancestors. Suppression of
the marine or of the freshwater life history phase of a
migratory lineage could lead to evolution of solely
freshwater or marine fishes, respectively, as speculated
by Patterson (1975: 168?169). In addition to evolution
of development, the roles of extinction and earth history
have been recognized: isolation or stranding through
changes in sea level or other vicariant events can result
in a former euryhaline species being restricted to
freshwater, for example (e.g., Choudhury & Dick, 1998;
Waters et al., 2000; Heads, 2005a), a phenomenon
termed ??ecological stranding?? (Craw et al., 1999).
Widespread extinction of the earth?s biota since the
Upper Mesozoic has left just remnants of these formerly
widespread distributions, ??the ring on the bathtub?? to
paraphrase Heads (1990: 225). Isolation in one habitat
or the other by stranding does not require or imply
invasion of that habitat (see also Heads, 2005a). Some
taxa persist in freshwater, others in marine habitats,
and some in both.
EARTH HISTORY MODELS
Plate tectonics is the model used most often to
interpret global distribution patterns with respect to
earth history (e.g., Rosen, 1974; Patterson, 1975;
Springer, 1982; Choudhury & Dick, 1998; Grande &
Bemis, 1999). The model specifies an Atlantic Ocean
expanding since the Mesozoic and supports vicariant
explanations for transatlantic taxa, yet leaves bioge-
ographers scrambling for similar explanations, such as
migration of allochthonous terranes, for the distribu-
tion of transpacific taxa (McCarthy, 2003: 1556). A
vicariance explanation of transpacific taxa finds
geological support in the still-maligned theory of
Expanding Earth, which specifies that the age of the
Pacific Ocean is roughly the same as that of the
Atlantic and Indian oceans (Shields, 1979, 1983,
1991, 1996; McCarthy, 2003, 2005). Biogeographic
sister areas across the Pacific have complementary,
matching geological outlines (McCarthy, 2003: fig. 3).
That there are competing models of earth history
endorses the view that biogeographic patterns should
be discovered, then interpreted with respect to these
models, rather than the models a priori specifying
biogeographic patterns (Ebach & Humphries, 2002).
Minimal divergence times between lineages have
been estimated using ages of fossils or a molecular
clock and used to test hypotheses of dispersal versus
vicariance (e.g., Rosenblatt & Waples, 1986; San-
mart??n et al., 2001; Burridge, 2002, to cite just three
examples). Minimal estimates from sequence data
have been interpreted as absolute, thus calling into
question earth history?Plate Tectonics or Expanding
Earth?interpretations of distribution patterns (e.g.,
de Queiroz, 2005). This interpretation has been
countered by Heads (2005b), Parenti (2006), and
others, continuing a long debate within biogeography
that was revived with the advent of molecular
techniques (see especially Nelson, 2004). Teleost as
well as basal actinopterygian lineages are old enough
to be interpreted within earth history models. Fossils
of living teleost lineages date from about 150 Ma in
the Upper Jurassic, and teleosts as a clade are
estimated to be at least of Triassic age (Janvier, 2007;
see also Benton & Donoghue, 2007). Differentiation is
also ancient: tetraodontiforms, nested within the
apomorphic percomorph clade, are represented by
fossils from the Upper Cretaceous of Slovenia, Italy,
and Lebanon, the oldest of which dates from 95 Ma
(Tyler & Sorbini, 1996).
DISTRIBUTION PATTERNS OF DIADROMOUS FISHES
Historical biogeographic patterns may be complex,
and are, by definition, hierarchical. They are
characterized by repeated elements that have become
the basis of broadly recognized global distribution
patterns (Croizat, 1958, 1964) that may be expressed
in a classification (Fig. 4): Atlantic, Indian Ocean,
(Pacific, Antitropical), or by fragments or composites
of these areas. One stated aim of historical biogeog-
raphy is to develop a standard language or nomen-
clature for global distribution patterns (e.g., Morrone,
2002) including their fragments. Fragmentation may
be caused by extinction, which has left remnants of
formerly complete distribution patterns, such as a lone
freshwater representative of a taxon once living
broadly in marine and freshwater habitats. These
remnants have been used to interpret, and misinter-
pret, the distributional history of taxa (see Heads,
2005a, b; Nelson & Platnick, 1981; Nelson & Ladiges,
1991, 1996; also see below).
Volume 95, Number 2 Parenti 239
2008 Life History Patterns and Biogeography
The distribution of antitropical or bipolar taxa has
been interpreted with regard to evolution of the
Pacific Basin (e.g., Humphries & Parenti, 1986, 1999;
Nelson, 1986). In a recent discussion of the evolution
of diadromy among fishes of the southern oceans,
McDowall (2002: 208) acknowledged but did not
discuss bipolarity. He divided fishes of New Zealand
into two kinds: those with close relatives throughout the
southern oceans, such as lampreys and galaxioids, both
groups bipolar, and those that he inferred, a priori, to be
part of another global pattern: ??Anguillid eels and
gobiid [fishes] have nothing in common with the
southern cool temperate fish fauna regarding historical
origins, distributions and relationships. They are
probably not Gondwanan and yet, anguillids and
gobiids express a series of relationships and distribu-
tions between the faunas of eastern Australia and New
Zealand that are essentially the same as those seen in
southern families?? (italics added) (McDowall, 2002:
212?213). Sharing a pattern implies sharing a history: I
examine the distribution of eels and gobies to ask if
they have anything in common with the southern cool
fish fauna?in other words, whether they conform to the
same global as well as local patterns.
PHYLOGENY AND DISTRIBUTION: EELS
Eels of the genus Anguilla are catadromous (Myers,
1949a): adults live in freshwater streams and migrate to
marine spawning grounds (Tesch, 1977; Smith, 1989a,
b). Phylogenetic relationships among the 19 extant taxa
(Fig. 5) were hypothesized using molecular data by two
research groups (Lin et al., 2001a, b; Aoyama et al.,
2001), and these two sets of analyses were contrasted
and compared with morphology (Lin et al., 2005). An
areagram of Anguilla taxa, with taxon names of
Figure 5 replaced by a brief description of distribu-
tional limits, is given in Figure 6. Another molecular
phylogeny based on whole mitochondrial genome
sequences was published by Minegishi et al. (2005).
Anguilla species are associated with all continents,
except Antarctica, and do not live in the eastern
Pacific or South Atlantic (Berra, 2001). Distribution of
anguillid eels has largely been interpreted within a
dispersalist paradigm (e.g., Harden Jones, 1968) and,
recently, to conflicting ends (Heads, 2005a: 700): Lin
et al. (2001a, b) put the center of origin of eels in the
southwest Pacific and proposed dispersal eastward
across the Pacific and further across the Central
American Isthmus to enter the Atlantic Ocean,
whereas Aoyama et al. (2001), Aoyama and Tsuka-
moto (1997), and Tsukamoto et al. (2002) hypothe-
sized dispersal from a center of origin near Borneo,
westward through an ancient Tethys Sea to the
Atlantic Ocean. Both of these explanations were
rejected by Minegishi et al. (2005: 141), who
concluded that ??. . . the present geographic distribu-
tion [of eels of the genus Anguilla] could be attributed
to, for example, multiple dispersal events, multidi-
rectional dispersion, or past extinctions, etc.??
Figure 4. Areagram or classification of Croizat?s (1958) and Craw et al.?s (1999) global biogeographic patterns, following
Nelson and Ladiges (2001: fig. 8b).
240 Annals of the
Missouri Botanical Garden
Panbiogeographers have long endorsed the view
that the distribution and evolution of eels of the genus
Anguilla may be interpreted with respect to earth
history, specifically evolution of a Tethys Sea biota
(Croizat, 1958), and dismiss ad hoc dispersal
hypotheses (see Heads, 2005a). Cladistic biogeogra-
phers endorse the panbiogeographic view and add
relationships among areas, as interpreted from
phylogenetic hypotheses, as a framework for further
refinement of distributional history patterns. Rather
than optimizing areas, the information in the areagram
of Anguilla taxa (Fig. 6) may be summarized by a
variety of methods, such as reduced area?cladograms
or paralogy-free subtrees (Nelson & Ladiges, 1996), or
by comparing it to the cladistic summary of Croizat?s
(1958) global biogeographic patterns (Fig. 4). Notable
Figure 5. Cladogram of relationships of eels of the genus Anguilla, following Lin et al. (2001a). Branch lengths are
arbitrary. Anguilla malgumora, as used by Lin et al., is a synonym of A. borneensis, the species name used by Aoyama et al.
(2001) and Minegishi et al. (2005). Spelling of species names and authors follows Eschmeyer?s online catalog (http://www.
calacademy.org/research/ichthyology/catalog/), accessed on 19 October 2007.
Volume 95, Number 2 Parenti 241
2008 Life History Patterns and Biogeography
in the areagram of Anguilla (Fig. 6) are taxa
distributed along the western margin of the Pacific
Basin (shaded group 1), the Indian Ocean (shaded
group 2), antitropical distributions, and antitropical
taxa sister to Pacific taxa. Eels demonstrate one of the
three-area relationships in the global biogeographic
pattern (Fig. 4): Pacific (boreal, austral). Northwest
Pacific taxa (boreal) are closely related to those from
eastern Australia, Tasmania, New Guinea, and New
Caledonia (austral, with additional elements from the
western margin of the Pacific Basin; shaded group 3),
and these are, in turn, sister to a group of Pacific taxa.
Another clade is also antitropical: the North Atlantic
sister species (boreal) are closely related to a group of
southwest Pacific, New Zealand, and New Caledonian
taxa (austral; shaded group 4). Freshwater eels of the
genus Anguilla have not been characterized as
antitropical or bipolar, as antitropicality of fishes is
seen largely as a phenomenon of strictly marine taxa
(e.g., Nelson, 2006: 12). These remnants of antitropi-
cality would likely not be identified without a
hypothesis of phylogenetic relationships or without
Figure 6. Areagram of Anguilla taxa, with taxon names of Figure 5 replaced by brief description of distributional limits.
NG 5 New Guinea, NC 5 New Caledonia, LHI 5 Lord Howe Island. Shaded area 1: western margin of Pacific Basin; shaded
area 2: Indian Ocean; shaded area 3: antitropical; shaded area 4: antitropical.
242 Annals of the
Missouri Botanical Garden
the aim of identifying general patterns, rather than
hypothesizing individual distributional histories.
Are anguillid eels ancient, like other antitropical
taxa? Perhaps. A molecular clock estimate of 20 Ma
for the beginning of Anguilla speciation was rejected
by Minegishi et al. (2005: 141) as an underestimate
because of fossil evidence: anguilliforms date from the
Upper Cretaceous (113?119 Ma) and the genus
Anguilla from the Eocene (50?55 Ma) (Patterson,
1993), both minimum estimates of the age of these
lineages.
PHYLOGENY AND DISTRIBUTION: GOBIES
Gobioid fishes comprise over 2000 species distrib-
uted broadly in pantropical and temperate freshwater,
estuarine, and marine aquatic habitats (Nelson, 2006).
Because the vast majority of gobioids live in marine
waters, they have been characterized as a marine
group (e.g., Myers, 1938; Berra, 2001). Several
subgroups, such as Gobiomorphus Gill, and the gobiid
subfamily Sicydiinae are diadromous (e.g., Parenti &
Maciolek, 1993; Keith, 2003). Phylogenetic analysis
of sicydiine gobies and their relatives at the generic
level (e.g., Parenti, 1991; Parenti & Thomas, 1998)
has been offered as evidence that the repeated,
recognizable biogeographic areas for gobioids are
ocean basins, not continents (see also Springer, 1982;
Gill in Lundberg et al., 2000).
Hypotheses of relationships among gobioids (e.g.,
Thacker & Hardman, 2005: fig. 1) are preliminary, yet,
when combined with distribution, display remnants of
global patterns (Fig. 4): Rhyacichthyidae, the sister
group to all other gobioids, lives in the tropical western
Pacific (Berra, 2001: 458), whereas Odontobutidae,
sister to the remaining gobioids, exhibits a remnant
boreal distribution, living in freshwaters of the
northwest Pacific (Berra, 2001: 460). The southeast
Australian and New Zealand Gobiomorphus, a goby
genus that McDowall (2002: 213) characterized as
??probably not Gondwanan,?? is interpreted as having a
remnant of an austral distribution; it is sister to a large
gobioid lineage distributed throughout the pantropics.
The fossil record of gobiids is scant; they date from at
least the Middle Tertiary (Patterson, 1993).
Endemism of gobioids on islands or island groups is
high, notably throughout the Indo-Pacific (Mauge? et
al., 1992; Parenti & Maciolek, 1993, 1996; Larson,
2001; Donaldson & Myers, 2002). Endemism through-
out the marine realm is underappreciated in large part
because marine species are defined traditionally as
widespread (e.g., Gill & Kemp, 2002, fishes; Meyer et
al., 2005, gastropods). Endemism may be under-
recognized at both lower (species) and higher (genera)
taxonomic levels. The lack of genetic structure among
populations of the five freshwater or amphidromous
gobiid fish species living throughout Hawaiian
streams (e.g., Chubb et al., 1998) prompted McDowall
(2003) to conclude that they must have colonized
Hawaii by long-distance, chance dispersal. Alterna-
tive explanations are plausible. Four of the five
species are endemic to the Hawaiian Islands, as
McDowall (2003: 705) acknowledged. Traditional
island biogeography tells us that what inhabits an
island or island group is what is able to physiolog-
ically and physically stand a long ocean voyage from
the mainland (MacArthur & Wilson, 1967), although
islands, as well as continents, are inhabited today by
the remnants of once more diverse faunas. Islands and
coastal zones support life that is able to withstand the
wide swings in salinity, temperature, and so on (see
also Heads, 2005a), and chief among that life is the
diadromous or euryhaline taxon.
CONCLUSIONS
Biogeography of diadromous fishes is not about
whether they are capable of surviving in the seas.
They are, at least during some phase of their life
history. It is not about whether diadromous fishes have
marine or freshwater ancestors because all fish
lineages have both. Biogeography of diadromous
fishes is like that of all other taxa: a search for a
pattern of relationships among endemic areas that
conforms to or differs from general distributions of
taxa worldwide. Eels and gobies, phylogenetically
disparate and distinct diadromous taxa, reflect global
biogeographic patterns. One can acknowledge this
congruence, this conformity to a pattern, or one can
ignore it. Predictions that may be made about
distribution of diadromous taxa based on their life
history are not necessarily corroborated by their
phylogeny or their distribution patterns. The answer
to ??Why migrate??? is straightforward when earth
history is considered: migration evolves as the earth
evolves (see also Wolfson, 1948; Croizat, 1958;
Heads, 2005a). Migration of diadromous species is
??the result. . . not the cause?? of their evolution
(Wolfson, 1948: 30; see also Grehan, 2006). When
interpreting the evolution of diadromy in fishes, or of
any migration pattern, the relationship between
phylogeny and distribution must be considered along
with ecological patterns. Ecology, phylogeny, and
distribution are inseparable (see also Heads, 1985).
The ability to live in both freshwater and marine
habitats is interpreted as an ancestral life history
pattern for fishes. The evolution of diadromy is tied to
the evolution of ocean basins. Optimizing habitat on
the nodes of an areagram to invoke an ancestral
habitat, here marine or freshwater, is equivalent to
Volume 95, Number 2 Parenti 243
2008 Life History Patterns and Biogeography
invoking a center of origin for areas. A repeated,
general pattern among taxa that are not closely related
and/or that differ in ecological requirements is most
parsimoniously explained by the same events. There
should be no separate global historical biogeography
for marine and freshwater fishes. Proposing separate
explanations for each prevents discovery of general
patterns.
Today, we see but a remnant of the past distribution
of life on earth. Global biogeographic patterns are the
organizing framework for interpreting distributional
and ecological history of both marine and freshwater
taxa. Recognizing that anguillid eels have remnants of
antitropical patterns, for example, is a first step
toward accepting that their distribution may be
classified along with that of other taxa, rather than a
priori requiring a unique explanation.
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