Molecular Ecology (2003) 12, 385–394
© 2003 Blackwell Publishing Ltd
Blackwell Science, Ltd
Fine-scaled geographical population structuring in
a highly mobile marine species: the Atlantic cod
H. KNUTSEN,*† P. E. JORDE,*† C. ANDRÉ ‡ and N. CHR. STENSETH *†
*Flødevigen Marine Research Station, Department of Coastal Zone, Institute of Marine Research, N-4817 His, Norway, †Division of
Zoology, Department of Biology, University of Oslo, Oslo, Norway, ‡Tjärnö Marine Biological Laboratory, Strömstad, Sweden
Abstract
Compared with many terrestrial and freshwater environments, dispersal and interbreeding
is generally much less restricted in the marine environment. We studied the tendency for a
marine species, the Atlantic cod, to be sub-structured into genetically differentiated popu-
lations on a fine geographical scale. We selected a coastal area free of any obvious physical
barriers and restricted sampling to a 300-km region, well within the dispersal ability of this
species. Screening 10 polymorphic microsatellite loci in 6 samples we detected a weak, but
consistent, differentiation at all 10 loci. The average F
ST
over loci was small (0.0023) but
highly significant statistically, demonstrating that genetically differentiated populations
can arise and persist in the absence of physical barriers or great distance. We found no geo-
graphical pattern in the genetic differentiation and there was no apparent trend of isolation
by distance along the coastline. These findings lend support to the notion that low levels
of differentiation are due to passive transport of eggs or larvae by the ocean currents rather
than to adult dispersal, the latter being strongly dependent on distance.
Keywords: Gadus morhua, gene flow, genetic differentiation, local populations, microsatellites
Received 6 October 2002; revision received 7 November 2002; accepted 7 November 2002
Introduction
Populations constitute interbreeding units with more or
less autonomous dynamics and recruitment. In terrestrial
and freshwater environments, populations are often well
defined and distinct from each other, often separated
physically by barriers to mixing and interbreeding (see,
e.g. Avise 2000 for an extensive review). In the marine
environment, in contrast, physical barriers are often absent
and the continuous water instead represents a potential
means for dispersal, favouring intermixing of individuals
over the species range. Tides and ocean currents may
further act to mix passively drifting organisms, primarily
eggs and larvae, over appreciable distances (see Palumbi
2001). For these reasons, distinct populations are more
difficult to detect in the marine environment and for
many marine organisms it is unclear to what degree
distinct populations exist at all, or whether they are
organized into larger panmictic units (McQuinn 1997). The
distinction is crucial, in particular for heavily exploited
organisms such as many marine fish, because recruitment
and sustainability are properties of the population. Failure
to identify the population could lead to local over-
exploitation and subsequent decline, as has become
abundantly clear in recent decades (e.g. Atlantic cod in
Alaska, Hutchings 2000, and along the Swedish west coast,
Svedäng et al . 2001).
Genetic markers are valuable tools for analysing popu-
lation structure (see, e.g. Utter 1991 for a historical review
of the application of genetic markers to fish populations).
By characterizing the geographical distribution of allele or
haplotype frequencies, population sub-structuring can be
detected and local populations identified. Such applica-
tions of genetic markers have been very successful in
uncovering cryptic population structure in freshwater
(Allendorf et al . 1976; Jorde & Ryman 1996; Carlsson et al .
1999) and marine fish (Nesbø et al . 2000; Ruzzante et al .
2000; Hutchinson et al . 2001). The use of genetic markers
for population delineation requires a detectable level of
genetic differentiation, however, and this has presented
problems in studies of many marine organisms (Ward et al .
1994). In the marine environment many studies have failed
to detect statistically significant population structuring
Correspondence: H. Knutsen. Fax: +47-370-59001; E-mail:
halvor@imr.no