Increasing globalization has spread invasive marine organisms, but it is not well understood why some species invade more readily than others. It is also poorly understood how species' range limits are set generally, let alone how anthropogenic climate change may disrupt existing species boundaries. We find a quantitative relationship that determines if a coastal species with a benthic adult stage and planktonic larvae can be retained within its range and invade in the direction opposite that of the mean current experienced by the larvae (i.e. upstream). The derivation of the retention criterion extends prior riparian results to the coastal ocean by formulating the criterion as a function of observable oceanic parameters, focusing on species with obligate benthic adults and planktonic larvae, and quantifying the effects of iteroparity and longevity. By placing the solutions in a coastal context, the retention criterion isolates the role of 3 interacting factors that counteract downstream drift and set or advance the upstream edge of an oceanic species' distribution. First, spawning over several seasons or years enhances retention by increasing the variation in the currents encountered by the larvae. Second, for a given population growth rate, species with a shorter pelagic period are better retained and more able to spread upstream. And third, prodigious larval production improves retention. Long distance downstream dispersal may thus be a byproduct of the many propagules often necessary to ensure local recruitment and persistence of a population in an advective environment.

In a single well-mixed population, equally abundant neutral alleles are equally likely to persist. However, in spatially complex populations structured by an asymmetric dispersal mechanism, such as a coastal population where larvae are predominantly moved downstream by currents, the eventual frequency of neutral haplotypes will depend on their initial spatial location. In our study of the progression of two spatially separate, genetically distinct introductions of the European green crab (Carcinus maenas) along the coast of eastern North America, we captured this process in action. We documented the shift of the genetic cline in this species over 8 y, and here we detail how the upstream haplotypes are beginning to dominate the system. This quantification of an evolving genetic boundary in a coastal system demonstrates that novel genetic alleles or haplotypes that arise or are introduced into upstream retention zones (regions whose export of larvae is not balanced by import from elsewhere) will increase in frequency in the entire system. This phenomenon should be widespread when there is asymmetrical dispersal, in the oceans or on land, suggesting that the upstream edge of a species' range can influence genetic diversity throughout its distribution. Efforts to protect the upstream edge of an asymmetrically dispersing species' range are vital to conserving genetic diversity in the species.invasive species | marine genetics | phylogeography | physical oceanography N ovel genetic material can appear in a population through mutation, migration, or long-distance dispersal events which may be human-mediated. In a single well-mixed population, the evolution of the frequency of novel neutral alleles will be governed by random genetic drift, not their initial spatial distribution (1). However, spatial structure and complexity can alter this expectation. In a metapopulation linked by migration, alleles introduced into source populations are more likely to persist than those that are introduced into sinks (2-4). Many metapopulations are embedded in complex spatial systems with a preferential direction of migration ("asymmetric dispersal"). In these systems, little is known about the equilibrium frequency of novel alleles or how this frequency depends on the location where these new lineages appear.Asymmetric dispersal is common where propagules are carried long distances by wind or water. In atmospheric, riverine, and oceanic flows, there is usually a predominant flow direction (downstream or downwind) that biases dispersal, and eddies or weather systems that slow or reverse such flow add a stochastic (and potentially upstream) component of migration. For example, many terrestrial plant species have propagules that can be dispersed by the winds (5), and a recent study of the spatial patterns of diversity in moss, liverwort, and lichen flora in the Southern Hemisphere were found to be best explained by the predominantly downwind dispersal (6). Further evidence that asymmetric dispersal can structure a species' genetic patter...

Understanding the processes that develop and maintain diversity in coastal communities is an important challenge, particularly given the conservation and management needs of these ecosystems. Such diversity patterns include not only species distributions and interactions, but also variation in genetic diversity. Alongshore variations in allele frequency along coastal oceans are controlled by interactions between physical and biological forces. Here we model these interactions in terms of Lagrangian descriptors of nearshore physical oceanography, the life history dynamics of an individual species and habitat quality. This model allows us to define population boundaries within the species range as a function of ocean circulation, spatial habitat variability and larval characteristics. From this, we can find quantitative criteria for the persistence of regions of alongshore genetic variation. Our results show quantitatively that the existence of phylogeographic patterns in species with high dispersal capacity along uniform coasts with typical mean currents is surprising, and requires either strong selection gradients, alongshore variation in ocean currents and/or habitat quality, or a mixture of both. Our model suggests that marine reserves and the harvesting of marine recources can dramatically modify spatial gradients in genetic diversity.

Aim  To determine timing, source and vector for the recent introduction of the European green crab, Carcinus maenas (Linnaeus, 1758), to Newfoundland using multiple lines of evidence.Location  Founding populations in Placentia Bay, Newfoundland, Canada and potential source populations in the north‐west Atlantic (NWA) and Europe.Methods  We analysed mitochondrial and microsatellite genetic data from European and NWA populations sampled during 1999–2002 to determine probable source locations and vectors for the Placentia Bay introduction discovered in 2007. We also analysed Placentia Bay demographic data and shipping records to look for congruent patterns with genetic analyses.Results  Demographic data and surveys suggested that C. maenas populations are established and were in Placentia Bay for several years (c. 2002) prior to discovery. Genetic data corroboratively suggested central/western Scotian Shelf populations (e.g., Halifax) as the likely source area for the anthropogenic introduction. These Scotian Shelf populations were within an admixture zone made up of genotypes from both the earlier (early 1800s) and later (late 1900s) introductions of the crab to the NWA from Europe. Placentia Bay also exhibited this mixed ancestry. Probable introduction vectors included vessel traffic and shipping, especially vessels carrying ballast water.Main conclusions  Carcinus maenas overcame considerable natural barriers (i.e., coastal and ocean currents) via anthropogenic transport to become established and abundant in Newfoundland. Our study thus demonstrates how non‐native populations can be important secondary sources of introduction especially when aided by human transport. Inference of source populations was possible owing to the existence of an admixture zone in central/western Nova Scotia made up of southern and northern genotypes corresponding with the crab’s two historical introductions. Coastal vessel traffic was found to be a likely vector for the crab’s spread to Newfoundland. Our study demonstrates that there is considerable risk for continued introduction or reintroduction of C. maenas throughout the NWA.

Abstract. A steady state cross-shelf density gradient of a wind-free coastal ocean undergoing winter time cooling is found for cooling and geometries which do not vary in the along-shelf direction. The steady state cross-shelf density gradient exists even when the average density of the water continues to increase. The steady state density gradient can be attained in less than a winter for parameters appropriate to the mid-Atlantic Bight. The cross-shelf eddy-driven buoyancy fluxes which cause this steady state gradient are found to depend critically on bottom friction and bottom slope, and the coastal polyna solutions of Chapman and Gawarkiewicz [1997] are significantly modified by this dependence in the limit of polynas with a large alongshore extent. Bottom friction retards the cross-shelf propagation of eddies, so that the buoyancy transport is no longer carried by self-advecting eddy pairs but mixed across the shelf by interacting eddies. The eddy interaction changes the length scale of the eddies until it is the lesser of the Rhines arrest scale or an analogous frictional arrest scale. The estimates of the steady state cross-shelf density gradient are found to compare well with numerical model results.