title for learn-about section on whelks & relatives in A SNAIL'S ODYSSEY
  Reproduction & development

photograph of colour morphs of whelk Nucella lamellosa from a rock/sand beach near White Rock, British Columbia

A adults, whelks crawl little and, with no free-living larvae, gene flow is consequently low. Thus, snails in widely spaced communities may have intrinsically different population characteristics, i.e., the potential is high for population differences over their geographical range.



Different colour morphs of whelks Nucella lamellosa
collected from a small beach in the Strait of Georgia, British
Columbia. The presence of so many varieties in an area of
only a few square meters is not usual for this species 1X

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Dispersal, genetic heterozygosity, & glacial refugia

  Topics in this section on reproduction & development include dispersal, heterozygosity, & glacial refugia, considered here, and EGG PRODUCTION & ENCAPSULATED DEVELOPMENT, HATCHLING ECOLOGY, NURSE EGGS, and IMPOSEX, considered in other sections.

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Research study 1

map showing sampling sites for whelks Nucella lamellosa in a study  of genetic diversityA study of genetic heterozygosity in the whelk Nucella lamellosa in northern Washington and southern British Columbia employs 2 different geographic scales. This first is along 100m of beach near Bellingham, Washington; the second, along 1000km of shoreline extending from San Juan Islands, Washington to entral Oregon.  2286 specimens are collected from 27 intertidal locations and analysed for 2 polymorphic allozymes Pep-2 and Pgm used as population markers.  At one extreme of the geographic scales used the authors find significant allele-frequency differences among 12 breeding populations, each comprising about 120 individuals, sampled on a single low tide along a continuous 100m boulder beach.  Overall, their results, both small- and large-scale, show the largest population subdivision yet reported for a marine gastropod, thus supporting the postulate that gastropods with limited larval and adult dispersals may have genetically fragmented populations.  Grant & Utter 1988 Malacologia 28: 275.

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Research study 2

map showing collection sites in study of distributions of whelks Kelletia kelletiidensity data for whelks Kelletia kelletii from Monterey Bay, California to Baja CaliforniaKelletia kelletii is a buccinid whelk inhabiting west-coast shores from California to Baja California whose range recently has been expanding northwards, past its historic limit at Point Conception to sites as far north as Monterey Bay (see map).  The extent to which this expansion is related to climate change and other factors is examined by researchers from UC Santa Barbara and UC Los Angeles.  The authors compare historic and contemporary distribution patterns in relation to seawater seawater temperature data for Monterey-Bay shorelines over timetemperature and water-circulation data, and find several points of interest.  First, contemporary populations have lower densities and less regular size-frequency distributions than historical ones,  suggesting a decrease in successful recruitment.  Second, the transition point between historical and contemporary populations corresponds with the confluence point of 2 major ocean currents at Point Conception  (see histogram), but warming trends beginning in the 1970s may have favoured the range expansion northwards.  The data suggest that changing temperature and oceanographic barriers have been major influences in both historical and contemporary patterns in population structure and northern range expansion in the species.  Zacherl et al. 2003 J Biogeogr 30: 913.

NOTE  determined from examination of published records of prehistoric shell middens

NOTE  the collison of currents creates convergent flow, thus presenting a potential barrier to transport of larvae north of the Point

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Research study 3

map showing distribution of haplotypes in the whelk Nucella Lamellosa along the west coast of North AmericaDespite having no free-swimming larva for dispersion, Nucella lamellosa inhabits a range on the west coast of North America at least 1000km north of the southern limit of glacier cover during the last glacial period 14-20,000yr ago.  The explanation for this is either survival in northern refugia during the period of glaciation or post-glacial colonisation by juveniles and/or adults.  As for the latter, crawling would have taken much longer than 20,000yrs, although rafting on algae is a possibility.  Studies on haplotype1 differences among individuals from several populations of N. lamellosa along the west coast from Oregon to Alaska indicate that the answer is most likely continuous habitation of refugia2 during the glacial period. 

Haplotype frequencies at different locations along the coast are shown for Nucella lamellosa on the map. Twenty-five haplotypes are present at 10 sites along the coast, but only one of these, haplotype 6, is common to most sites.  Twenty of the haplotypes are specific to certain sites, with most occurring in Alaska.  Note the disjunction3 in haplotype types and distributions between southern Alaska and northern British Columbia. Although commonly thought of as a continuous sheet of ice during the last glacial period, it is more likely that the majority of coastal glaciers existed as discrete structures separated by areas of uncovered land.  The inter-digitation of ice and land may have disrupted species’ ranges, leading to genetic differentiation.  Marko 2004 Molecular Ecol 13: 597.

NOTE1  lit. “single type” G., referring to a set of genes in each chromosome which tend to be transmitted as a unit to the next generation.  Haplotypes are defined here from mtDNA nucleotide-sequence data from the first subunit of cytochrome c oxidase.

NOTE2  at least one such refugium has been identified as existing through the last glaciation period on the east side of Graham Island in Haida Gwai, British Columbia, indicated by a blue dot on the map. Warner et al. 1982 Science 218: 675. 

NOTE3   similar disjunctions are described for other west-coast benthic marine invertebrates in the ODYSSEY, including sea cucumbers: LEARN ABOUT SEA CUCUMBERS: GENETIC DRIFT   None of these other invertebrates has a free-living planktonic larval phase

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Research study 4

map showing collection sites for genetics study on glaciation effectsIn a later contribution to the interesting topic of glaciation effects on distributions of rocky-shore marine invertebrates in the northeastern Pacific, the same author with colleagues from Clemson University and Simon Fraser University collect and analyse mitochondrial DNA data sets for 12 invertebrate species1 in 4 phyla, including whelks Nucella lamellosa and N. ostrina (see RS3 above) Their findings provide strong support for a “hit and miss” pattern of glaciation effects on survival and distribution of rocky shore marine invertebrates on this coast. Thus, results for 4 species2 (2 seastars and 2 winkles) show demographic patterns consistent with large population expansions occurring at the time of the last glacial maximum (20,000ybp), such as would be expected from recent colonisation. In contrast, 6 invertebrate species3 (2 mussels, 2 barnacles, 1 seastar, and 1 chiton) show genetic patterns consistent with long-term stability in populations, consistent with persistence during the last glacial maximum. The remaining 2 invertebrates, N. lamellosa and N. ostrina show spatially variable demographic histories. The study is interesting for several reasons, the most notable being a big swing-shift in our belief that glaciation effects on this coast were fatally exterpative to a certain defined southern latitude; rather, it now seems that long-term survival through the last glaciation event occurred in discontinuous geographical pockets. In this regard, the situation for coastal marine species differs from that found in comparable terrestrial studies, where regional persistence in northern cryptic refugia appears to have been less common. Marko et al. 2010 Mol Ecol 19: 146.

NOTE1 2 species of prickleback fishes Xiphister are included in the study, but not mentioned further here

NOTE2 seastars: Pisaster ochraceus, Evasterias troschelii; winkles: Littorina sitkana, L. scutulata

NOTE3 mussels: Mytilus californianus, M. trossulus; barnacles: Balanus glandula, Semibalanus cariosus; seastar: Patiria miniata; chiton: Katharina tunicata

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Research study 5

In a study related to that of Research Study 2 above, a consortium of researchers from several west-coast and one Hawai’ian university investigate in an ingenious way gene flow in relation to ocean currents using the buccinid whelk Kelletia kelletii as the model.  Simply put, if you want to determine the effect of inter-population distance on genetic structure in the adults of a marine species, then you follow the genes carried in the disseminative propagules, the larvae, along the dispersal track defined by the ocean currents carrying these larvae.  Now, experimentally speaking, this would be difficult, if not impossible to do, so the researchers do the next best thing and that is to simulate larval dispersal from 10 Kelletia-inhabited sites in shallow areas along the Santa Barbara Channel using a computer model of direction and intensity of the local ocean currents.  Since currents and eddies don’t move in  straight lines, the data obtained from genetic analysis of snails at the 10 sites located along the projected tracks of currents provides far more accurate information on extent of genetic connectivity than would be gained strictly from measure of Euclidian (straight-line) distances.  In fact, use of relative oceanographic distances in this study explains almost 50% of the variance in measured pairwise genetic differences in adult Kelletia among the 10 sites over a scale of about 120 linear km (see schematic)  The authors interpret patterns of genetic structure of populations that correspond with patterns of ocean circulation, a concept encapsulated in the term “seascape genetics”.  The idea for the study is not new, but the strength of the data obtained for Kelletia is so convincing that, in the words of the authors, the methodology “represents a novel approach to characterize population connectivity at small spatial scales”, and thus has great potential value for conservation and fisheries management.  White et al. 2010 Proc Roy Soc B 277: 1685.

schematic showing probabilities of genetic connectivity between populations of whelks Kelletia kelletii in the Santa Barbara ChannelNOTE  a model of near-surface ocean flow is developed from data on ocean temperature, salinity, currents, and winds, and is “seeded” with virtual larvae at 448 25km-2 coastal grid locations each day for a 2mo period in summer (corresponding with the seasonal spawning time of Kelletia).  The species has a 40-60d pelagic larval duration

NOTE  the researchers evaluate genetic polymorphism at 9 microsatellite loci using established methodology

3 types of pairings are used in the analyses: between sites on the islands
(red), between sites on the mainland (green), and  between sites across
the Channel (blue).  Line thicknesses correlate with probability of
dispersal over a single generation from one site to another .  Some
pairings not joined may seem to have good connectivity based on
the current patterns , but larval settlement may actually be limited
owing to high offshore advection of larvae in fast-flowing currents

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