Molecular Ecology (2004) 13, 2143–2156 doi: 10.1111/j.1365-294X.2004.02225.x
© 2004 Blackwell Publishing Ltd
Blackwell Publishing, Ltd.
Strong genetic clines and geographical variation in gene
flow in the rocky intertidal barnacle Balanus glandula
ERIK E. SOTKA,* JOHN P. WARES,† JOHN A. BARTH,‡ RICHARD K. GROSBERG † and
STEPHEN. R. PALUMBI *
* Stanford University, Department of Biological Sciences, Hopkins Marine Station, Pacific Grove, CA 93950, † Center for Population
Biology, Division of Biological Sciences, 1 Shields Drive, University of California, Davis, CA 95616, ‡ College of Oceanic and
Atmospheric Sciences, Oregon State University, 104 Ocean Administration Building, Corvallis, OR 97331, USA
Abstract
A long-standing issue in marine biology is identifying spatial scales at which populations
of sessile adults are connected by planktonic offspring. We examined the genetic continuity of
the acorn barnacle Balanus glandula, an abundant member of rocky intertidal communities
of the northeastern Pacific Ocean, and compared these genetic patterns to the nearshore
oceanography described by trajectories of surface drifters. Consistent with its broad dispersal
potential, barnacle populations are genetically similar at both mitochondrial (cytochrome
oxidase I) and nuclear (elongation factor 1-alpha) loci across broad swaths of the species’
range. In central California, however, there is a striking genetic cline across 475 km of coastline
between northern and southern populations. These patterns indicate that gene flow within
central California is far more restricted spatially than among other populations. Possible
reasons for the steep cline include the slow secondary introgression of historically separated
populations, a balance between diversifying selection and dispersal, or some mix of both.
Geographic trajectories of oceanic drifters closely parallel geographical patterns of gene
flow. Drifters placed to the north (Oregon; ∼44°N) and south (Santa Barbara, California;
∼34° N) of the cline disperse hundreds of kilometres within 40 days, yet over the long-term
their trajectories never overlapped. The lack of communication between waters originating
in Oregon and southern California probably helps to maintain strong genetic differentiation
between these regions. More broadly, the geographical variation in gene flow implies that
focusing on species-level averages of gene flow can mask biologically important variance
within species which reflects local environmental conditions and historical events.
Keywords: California Current, cline, dispersal, marine invertebrate, mtDNA sequences, nuclear
sequences, selection
Received 22 December 2003; revision received 25 March 2004; accepted 7 April 2004
Introduction
Virtually all species of bottom-dwelling marine organisms
are distributed in patches that are linked by the dispersal
of planktonic larvae (Thorson 1950). As such, larval dispersal
plays a fundamental role in the ecology and evolution
of marine organisms and their biotic interactions (see re-
views in Caley et al . 1996; Botsford et al . 2001; Grosberg &
Cunningham 2001; Strathmann et al . 2002). Genetic estimates
of the spatial extent of larval dispersal generally support
a positive correlation between a species’ ability to disperse
— as measured by larval lifespan — and the actual distances
they travelled (Bohonak 1999). However, a number of more
recent studies on fish and invertebrates with long-lived
planktonic larvae indicate that planktonic larvae can be
locally retained (e.g. Jones et al . 1999; Swearer et al . 1999;
Barber et al . 2000; Taylor & Hellberg 2003), suggesting that
the realized dispersal of marine organisms is probably less
extensive in distance than estimates based on life-history
alone predict.
More importantly, there is growing recognition that
realized dispersal is probably more complicated than can
be estimated from limited sampling from a small portion
Correspondence: Erik E. Sotka. Fax: 831-655-6215; E-mail:
sotka@stanford.edu