Evolutionary consequences of microhabitat: population-genetic structuring in kelp- vs. rock-associated chitons

Rafting has long been invoked as a key marine dispersal mechanism, but biologists have thus far produced little genetic evidence to support this hypothesis. We hypothesize that coastal species associated with buoyant seaweeds should experience enhanced population connectivity owing to rafting. In particular, invertebrates strongly associated with the buoyant bull-kelp Durvillaea antarctica might be expected to have lower levels of population-genetic differentiation than taxa mainly exploiting nonbuoyant substrates. We undertook a comparative genetic study of two codistributed, congeneric chiton species, assessing population connectivity at scales of 61-516 km, using ≥ 186 polymorphic AFLP loci per species. Consistent with predictions, population-genetic differentiation was weaker in the kelp-associated Sypharochiton sinclairi than in the rock-associated S. pelliserpentis. Additionally, while we found a significant positive correlation between genetic and oceanographic distances in both chiton species, the correlation was stronger in S. pelliserpentis (R(2) = 0.28) than in S. sinclairi (R(2) = 0.18). These data support the hypothesis that epifaunal taxa can experience enhanced population-genetic connectivity as a result of their rafting ability.     

Phylogeography of the snakeskin chiton Sypharochiton pelliserpentis (Mollusca: Polyplacophora) around New Zealand: are seasonal near‐shore upwelling events a dynamic barrier to gene flow?

We present a phylogeographic study of the New Zealand and Australian intertidal chiton Sypharochiton pelliserpentis that was conducted to ascertain levels of population connectivity and to investigate the effect of previously hypothesized general phylogeographic boundaries. The analysis incorporated both cytochrome oxidase subunit 1 sequence data (approximately 700 bp) and RFLP data from 29 populations around New Zealand, and from one Australian population, for a total of N = 472. The major population structure observed was a strong disjunction between northern and southern populations (Φ_ST = 0.47), with the genetic breaks located at Cloudy/Clifford Bay and Farewell Spit, at the northern tip of the South Island. This finding corresponds with a common phylogeographic barrier observed in a number of other marine invertebrates, highlighting its significance and ubiquity. A third barrier to gene flow was identified between Spirits Bay and Ahipara, around the northern tip of the North Island. All three of these areas that exhibit significant population disjunctions have strong near‐shore upwelling along with water current movement offshore, and these features are prevalent during the time of year when S. pelliserpentis spawns. That these seasonal hydrographic patterns contribute to the population structuring of S. pelliserpentis is supported by comparison with other phylogeographic studies of marine invertebrates, where the strength of this barrier seems to correlate with spawning season. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 552–563.     

The Bering Strait connection: dispersal and speciation in boreal macroalgae

A large number of boreal seaweeds have either sibling species or conspecific populations of a single species in the North Pacific and North Atlantic Oceans. This pattern is thought to have arisen from the dispersal between the two oceans through the Arctic Ocean after the opening of the Bering Strait in the mid-to-late Miocene or earliest Pliocene and from subsequent vicariant speciation as the Arctic Ocean froze and Bering Strait closed intermittently during glacial periods. Recent molecular studies of species in all three major seaweed phyla reveal patterns of vicariance. However, a number of lines of evidence point to differences in origins of these clades; some appear to be Pacific in origin whereas others appear to be derived from Atlantic stock. Different origins can be explained by recent stratigraphic finds that push the first Cenozoic opening of the Bering Strait back from 3.1–4.1 to 4.8–7.4 Ma (million years ago). Northern hemisphere ocean circulation models suggest that water flow would have been from the North Atlantic–Arctic south through the Bering Strait prior to the closure of the Panamanian Isthmus c. 3.5 Ma in contrast to the northward flow from the Pacific into the Arctic and North Atlantic, which developed after the closing of the Isthmus. Despite these differences in timing of the two invasions, there are no significant differences in levels of relationships among species with a North Atlantic origin compared with species with a North Pacific origin based on currently available data. More work is required to understand vicariance in seaweeds, especially in deciphering when a speciation event has occurred.     

The arrival of propagules of marine macroalgae in the intertidal zone

The composition and abundance of macroalgal propagules contained in sea water arriving at intertidal rocky shores was estimated monthly at Pelancura, central Chile, from June to December, 1984. Samples from surface water and from water running off rocky platforms with mixed algal vegetation were cultured in laboratory conditions and examined for development of sporelings. Thirty eight macroalgal entities grew in the cultures, 75% of them with opportunistic life-styles. The total number of sporelings was similar in the surface water of sites separated by 300 m. Marked variability in composition and number of sporelings was, however, observed between samples taken simultaneously at one site and between different months, suggesting patchiness in the dispersal of propagules. Marked differences were established in the ratio run-off water/surface water in the number of sporelings, which could be related to the dispersal shadows of the various taxa. Sporelings of late successional algae showed marked spatial and temporal variations in number. Some taxa of opportunistic algae were characterized by the development of several thousand sporelings per litre of sea water while in others the number of sporelings was two or three orders of magnitude less, suggesting differences in reproductive effort.     

Pacific Coral-reef Fishes: The Implications of Behaviour and Ecology of Larvae for Biodiversity and Conservation, and a Reassessment of the Open Population Paradigm

The two-phase life history of most marine fishes and invertebrates has enormous implications for dispersal, population connectivity, and resource management. Pelagic dispersal larvae of marine animals traditionally thought to ensure that populations are widespread, that chances of local extinction are low, and that marine protected areas (MPA) can easily function to replenish both their own populations and those of unprotected areas. Traditionally, dispersal is considered to depend primarily on two variables: pelagic larva duration and far-field currents. These conclusions arise from the open population paradigm and are usually accompanied by a simplifying assumption: larvae are distributed passively by far-field currents. Unfortunately, they ignore the complex reality of circulation and hydrological connectivity of reefs, and do not consider newly-demonstrated behavioural capabilities of coral-reef fish larvae. Far-field circulation varies with depth and often excludes water bodies where propagules are released, and this has important implications for predicting trajectories of even passive larvae. However, larvae are not passive: late-stage larvae of coral-reef fishes can swim faster than currents for long periods, can probably detect reefs at some distance, and can actively find them. This behaviour is flexible, which greatly complicates modelling of larval fish trajectories. Populations at ecological (as opposed to evolutionary) scales are probably less open and more subdivided than previously assumed. All this means that dispersal predictions based solely on far-field water circulation are probably wrong. An emerging view of larval-fish dispersal is articulated that takes these new data and perspectives into account. This emerging view shows that re-evaluation of traditional views in several areas is required, including the contribution of larval-fish biology and dispersal to biodiversity patterns, the way reef fishes are managed, and the way in which MPA are thought to operate. At evolutionary and zoogeographic scales, reef-fish populations are best considered to be open.     

Radio-telemetric monitoring of dispersing stag beetles: implications for conservation

Migratory movements of the endangered stag beetles Lucanus cervus (18 males, 38 females) were monitored radio-telemetrically for three reproductive periods (2003–2005). The aim of the study was to estimate the migratory range of free-ranging individuals as a measure of connectivity among neighbouring populations for future conservation measures. Miniature transmitters of c . 350 mg (battery life: 10–15 days) were attached externally to the pronotum. Transmitter/beetle mass ratio was 12.8% on average (7.1–28.0%). Male dispersal behaviour consisted of frequent flights directed to sites with reproductive females and rarely of on-ground movement. Total displacement distance recorded was up to 2065 m, the maximum distance of a single flight being 1720 m. Flights always began at elevated structures such as trees and shrubs and took place in an air temperature range of 11–27°C. Within this range, temperature did not influence flight distance. Female dispersal behaviour consisted mostly of a single flight, followed by mating and consequent ground movements towards oviposition sites. Total displacement distance recorded was up to 762.6 m, the maximum distance of a single flight being 701 m. Climatic constraints of flights were the same as in males. Modelling the dispersal behaviour suggests that about 1% of males are capable of maintaining gene flux among nest sites within a radius of about 3 km. However, the colonization of new nest sites depends on the dispersal ability of females and amounts to less than 1 km. Thus, isolated populations (distance to the next population greater than 3 km) have an increased probability of local extinction.     

Testing the consistency of connectivity patterns for a widely dispersing marine species

Connectivity is widely recognized as an important component in developing effective management and conservation strategies. Although managers are generally most interested in demographic, rather than genetic connectivity, new analytic approaches are able to provide estimates of both demographic and genetic connectivity measures from genetic data. Combining such genetic data with mathematical models represents a powerful approach for accurately determining patterns of population connectivity. Here, we use microsatellite markers to investigate the genetic population structure of the New Zealand Rock Lobster, Jasus edwardsii, which has one of the longest known larval durations of all marine species (>2 years), a very large geographic range (>5500 km), and has been the subject of extensive dispersal modeling. Despite earlier mitochondrial DNA studies finding homogeneous genetic structure, the mathematical model suggests that there are source-sink dynamics for this species. We found evidence of genetic structure in J. edwardsii populations with three distinct genetic groups across New Zealand and a further Australian group; these groups and patterns of gene flow were generally congruent with the earlier mathematical model. Of particular interest was the consistent identification of a self-recruiting population/region from both modeling and genetic approaches. Although there is the potential for selection and harvesting to influence the patterns we observed, we believe oceanographic processes are most likely responsible for the genetic structure observed in J. edwardsii. Our results, using a species at the extreme end of the dispersal spectrum, demonstrate that source-sink population dynamics may still exist for such species.     

The influence of coastal topography, circulation patterns, and rafting in structuring populations of an intertidal alga

Understanding the dispersal processes that influence genetic structure in marine species requires estimating gene flow in a dynamic, fluid environment that is often poorly characterized at scales relevant to multiple dispersive stages (e.g. spores, gametes, zygotes, larvae, adults). We examine genetic structure in the marine alga Fucus vesiculosus L., which inhabits moderately exposed shores in the northern Atlantic but releases gametes only under sunny, calm conditions. We predicted genetic structure would correlate with coastal topography because weather frequently varies across coastal promontories on the Maine shore when F. vesiculosus is reproductive, which causes one side to experience high levels of water motion (= no gamete release) while one side is calm (= gamete release). Furthermore, we expected that the effect of low dispersal capacities of gametes and zygotes would result in spatial genetic structure over short distances. Using surface drifters, we characterized near-shore circulation patterns around the study sites to investigate whether directionality of gene flow was correlated with directionality of currents. We found significant genetic differentiation among sites sampled at two different peninsulas, but patterns of differentiation were unrelated to coastal topography and there was no within-site spatial structuring. Our genetic and near-shore circulation data, combined with an examination of gamete longevity, support the dependency of gene flow on storm-detached, rafting, reproductive adults. This study highlights the significance of rafting as a mechanism for structuring established populations of macroalgae and associated biota and demonstrates the importance of coupling population genetics' research with relevant hydrodynamic studies.     

NEW ZEALAND HERD STRANDING SITES DO NOT RELATE TO GEOMAGNETIC TOPOGRAPHY

Recent theories relating cetacean stranding sites to geomagnetic topography were tested for the New Zealand coastline. New Zealand herd strandings showed no relationship to either perpendicular geomagnetic contours or magnetic minima, nor did whales appear to be avoiding magnetic gradients. Single-live stranding sites also showed no geomagnetic relationship.     

Molecular genetics of peripheral populations of Nova Scotian Unionidae (Mollusca: Bivalvia)

Peripheral populations of eight species of freshwater bivalves (Unionidae.) extending their geographic ranges into Nova Scotia, Canada, were examined electrophoretically to determine both the extent of genetic variability within such populations, and whether the hypothesized pathway of colonization across the Isthmus of Chignecto is reflected in patterns of genetic resemblance among these populations. The Nova Scotian species examined could be separated into two groups based on levels of observed heterozygosity and levels of variability in allele frequencies. The first group is characterized by low levels of heterozygosity and polymorphism compared with north-eastern American populations, and in the case of one species, Elliptio complanala, considerable variability in allele frequencies among populations occurring in similar habitats in different drainages. Populations of E. complanata from Nova Scotia can be differentiated from conspecific populations on the southern Atlantic Slope by possession of fast alleles at two loci. Multivariate analyses define subgroups within populations of E. complanata consistent with hypothesis that the species invaded Nova Scotia by way of the Isthmus of Chignecto, and then split into two groups, one of which colonized Cape Breton to the north and the other of which colonized southern areas of the Province. The second group of Nova Scotian species is characterized by little reduction in heterozygosity and polymorphism compared with values observed among north-eastern American conspecifics or congeners, little variability in allele frequencies from population to population, and little evidence to suggest that these species were dependent on the land bridge to invade the Province. The type of dispersal is hypothesized to be responsible, in part, for these differences: larvae of species in the first group rely on a parasitic attachment to fish with territorial habits limited to fresh water, and are thus likely to invade new drainages separated by salt water by chance, in small numbers, and in stepping-stone fashion. Species in the second group parasitize anadromous or saltwater tolerant hosts, are likely to be introduced into new habitats in greater numbers and/or receive greater amounts of gene flow subsequent to colonization, and seem less dependent on land-bridges to colonize new habitats.     

A model study on variations in larval supply: are populations of the polychaete <Emphasis Type="Italic">Owenia fusiformis</Emphasis> in the English Channel open or closed?

The polychaete Owenia fusiformis is one of the most ecologically important species in the muddy fine sand sediments in the English Channel where it is distributed in geographically separated populations. A vertically averaged Lagrangian hydrodynamic model integrating tidal residual currents and wind-induced currents was used to drive an advection–diffusion model for investigating the variability of larval transport in order to assess the self-seeding capabilities and the degree of connectivity between local populations. Three different types of environmental forcing (i.e. tidal forcing alone, tidal forcing coupled with either NE winds or SW winds) were applied to 19 distinct populations. Without wind influence, self-seeding is the principal mechanism involved in the renewal of most populations. However, larval retention ranged from under 1% up to 81% in relation to the adult habitat size and the mean velocity of tidal residual currents. Wind forcing had a strong influence on larval dispersal patterns by modifying the origin and densities of settlers as well as the degree of connectivity between populations. As a consequence, larval supply from distant populations generally exceeded local supply and the inter-annual variability of wind forcing induced large year-to-year variations in larval settlement rates. Larval exchanges occurred mainly between neighbouring populations and three groups of interconnected local populations were thereby identified. Within each group, settlement patterns were related to inter-annual variations in the direction and magnitude of larval exchanges. Electronic Publication     

Rafting in Antarctic Collembola

Darwin was an early exponent of the importance of 'occasional means of dispersal' in accounting for the present-day distribution of plants and animals. This study examined the implications of capture on the water surface of meltwater and seawater for the local and long-range dispersal of Antarctic springtails. Individuals of the maritime Antarctic collembolan Cryptopygus antarcticus , were floated on tap water and seawater at 0, 5 and 10°C. LT₅₀s on seawater were 34 (10°C), 65 (5°C) and 75 (0°C) days. On tap water, LT₅₀s were 69 (10°C), 126 (5°C) and 239 (0°C) days. Less than 20% escaped from the water surface. A significantly greater proportion of springtails moulted on tap water and viable offspring were produced on both tap water and seawater. Comparison across treatments of survival of moulting and non-moulting individuals found significantly greater survival in moulting animals for three of the treatment combinations. It is suggested that moult exuviae facilitate survival on the water film through the simultaneous provision of a flotation aid and a source of nourishment – that is, an 'edible raft'. A separate experiment measuring changes in haemolymph osmolality over time on tap water and seawater at 2 and 5°C found significant differences in all treatments. Causes of mortality are discussed in relation to osmoregulatory failure and starvation.