Molecular genetic evidence suggests Long Island as the geographic origin for the present population of bay scallops in Barnegat Bay, New Jersey.ABSTRACT We have used molecular genetic methods to examine the question of the geographic origins of the newly returned Argopecten irradians populations in Barnegat Bay (BB), New Jersey. Using PCR PCR polymerase chain reaction. PCR abbr. polymerase chain reaction Polymerase chain reaction (PCR) to amplify specific polymorphic microsatellite See miniaturized satellite. regions for bay scallop scallop or pecten, marine bivalve mollusk. Like its close relative the oyster, the scallop has no siphons, the mantle being completely open, but it differs from other mollusks in that both mantle edges have a row of steely blue "eyes" and , we have genetically compared the 2004 and 2005 BB populations to those from Long Island (LI), New York, and North Carolina (NC). Our studies indicate that the 2004 and 2005 BB populations are genetically similar with some allelic frequency differences. Five of the eight loci studied are identical for marker size among BB, LI, and NC populations. The $336 locus demonstrates polymorphic sequences of 138 and 158 basepairs in the NC population that are not observed in LI or BB. The C 1832 locus appears identical (122 basepairs) between LI and BB, but demonstrates polymorphisms (132 or 142 basepairs) in the NC population. Additionally, the NC group manifests two further alleles in the M26 locus (135 and 149 basepairs) not seen in BB or LI. These results, along with genetic distance and mean estimated gene flow calculations, support a physical transfer of the Long Island bay scallop larvae Larvae, in Roman religion Larvae: see lemures. down the Atlantic coast to the transition regions around Barnegat Bay. KEY WORDS: bay scallops, Argopecten irradians, microsatellite markers, DNA fingerprinting, sequence length polymorphisms, phylogenetic analysis INTRODUCTION Bay scallops (Argopecten irradians Lamarck) are a common estuarine es·tu·a·rine adj. 1. Of, relating to, or found in an estuary. 2. Geology Formed or deposited in an estuary. Adj. 1. estuarine - of or relating to or found in estuaries estuarial species along the eastern and southern coasts of North America from Massachusetts to Texas. Three clear subspecies subspecies, also called race, a genetically distinct geographical subunit of a species. See also classification. (A. i. irradians, A. i. concentricus, A. i. amplicostatus) have been characterized within this geographic range based on shell morphometrics Generally, morphometrics (from the Greek: "morph," meaning shape or form, and "metron”, meaning measurement) comprises methods of extracting measurements from shapes. In most cases applied to biological topics in the widest sense. and molecular phylogenetics phy·lo·ge·net·ics n. The study of phylogeny. (Clarke 1965, Blake & Graves 1995, Blake et al. 1997, Marelli et al. 1997, Bologna et al. 2001). The bay scallop is prized both commercially and recreationally throughout its range and represents an important species in ecology and commerce. Historically, bay scallops were abundant and commercially fished in New Jersey, USA. The first available landing records were collected in 1956, when 52,300 bushels were harvested with an estimated value of $287,000. Continued success of scallop populations for the next 12 y yielded 317,000 bushels valued at over $1 million (Ford 1997). Subsequently, commercial bay scallop harvests were only recorded for 1973 and 1974. Despite its local importance, little information exists on the ecology and population structure of New Jersey bay scallops, particularly because of the collapse of the commercial fishery. In recent years, it was generally believed that bay scallops no longer occurred in New Jersey waters. The observation of numerous scallops in 1998 in Little Egg Harbor Little Egg Harbor is a brackish bay along the coast of southeast New Jersey. It was originally called Egg Harbor by the Dutch sailors because of the eggs found in nearby gull nests. Tributaries
1. pertaining to larvae. 2. larvate. larval migrans see cutaneous and visceral larva migrans. origin for the recolonization Re`col`o`ni`za´tion n. 1. A second or renewed colonization. . It was the goal of this present research to (a) investigate the genetic population structure of the these newly arisen Barnegat Bay (BB) scallops from 2004 and 2005 using species-specific microsatellite loci (Roberts et al. 2005) and (b) compare allelic frequencies among the Barnegat Bay, Long Island, NY (LI), and North Carolina (NC) bay scallop populations with the intent of establishing the geographic origin of the current BB population. MATERIALS AND METHODS Scallop Tissue Samples Bay scallops (Argopecten irradians Lamarck) were collected within extensive eelgrass beds (July 2004 and August, 2005) in Little Egg Harbor, NJ, USA (39[degrees]35'N, 74[degrees]14'W), which is located in the central portion of the MidAtlantic Bight bight, broad bend or curve in a coastline, forming a large open bay. The New York bight, for example, is the curve in the coast described by the southern shore of Long Island and the eastern shore of New Jersey. The term bight may also refer to the bay so formed. . The Long Island scallops were collected in 2005 by Dr. Brad Peterson (Long Island University) from Peconic Bay. North Carolina scallops were collected in 2005 by Wayne Cuthrell and Marc Hamric (North Carolina Division of Marine Fisheries) in Bogue Sound off the coast of Morehead City, NC. Scallops were returned live to our laboratory and dissected. Adductor muscle Noun 1. adductor muscle - a muscle that draws a body part toward the median line adductor skeletal muscle, striated muscle - a muscle that is connected at either or both ends to a bone and so move parts of the skeleton; a muscle that is characterized by tissue was extracted from individuals and stored frozen at -80[degrees]C for later DNA extraction. Long Island and North Carolina samples were shipped overnight on ice to New Jersey. DNA Extraction DNA DNA: see nucleic acid. DNA or deoxyribonucleic acid One of two types of nucleic acid (the other is RNA); a complex organic compound found in all living cells and many viruses. It is the chemical substance of genes. was extracted from fifteen 2004 and thirteen 2005 Barnegat Bay scallops. Twenty Long Island and 20 North Carolina scallops were also used. The DNA was obtained from 0.5-0.8 g of adductor muscle tissue. The tissue was stored frozen at -80[degrees]C and homogenized ho·mog·e·nize v. ho·mog·e·nized, ho·mog·e·niz·ing, ho·mog·e·niz·es v.tr. 1. To make homogeneous. 2. a. To reduce to particles and disperse throughout a fluid. b. in an ice-cooled 1.5 mL microfuge tube with a plastic DNasc/RNase-free micropestle. The DNeasy extraction kit (Qiagen Corp., Valencia, CA) was used for DNA extraction following the manufacturer's directions. Primers and PCR Amplification Eight microsatellite primer sets were developed for bay scallops by Roberts et al. (2005). These primers were synthesized for us by Invitrogen Corp (Carlsbad, CA). The PCR amplification conditions generally followed the directions of Roberts et al. (2005). The only protocol alteration was an increase in the annealing annealing (ənēl`ĭng), process in which glass, metals, and other materials are treated to render them less brittle and more workable. temperature to 60[degrees]C used with the GP63 locus to reduce the emergence of triplets. All PCR amplification was performed in a Mastercycler gradient thermocycler (Eppendorf, Inc., Westbury, NY). Microsatellite Fragment Analysis DNA fragments were electrophorctically separated in a high-resolution manner and analyzed by the methods used in our previous studies (Vander Zwan et al. 2000, Campanella et al. 2003, Provan & Campanella 2003, Campanella et al. 2004). The PCR products were imaged with Scion computer software (Scion, Inc., Frederick, MD). Molecular weights were analyzed and calculated with Collage Version 4.0 (Image Dynamics Corporation, Surrey, BC, Canada). Population Genetic Calculations Chord distances (Cavalli-Sforza & Edwards 1967) were computed using the Microsat program (Minch et al. 1995). The neighbor-joining radial tree was calculated employing the chord distance data into the Phylip subroutine A group of instructions that perform a specific task. A large subroutine might be called a "module" or "procedure." Subroutine is somewhat of a dated term, but it is still quite valid. Neighbor (Felsenstein 1993), and the tree itself was generated by Treeview (Page 1996). The mean estimated gene flow (Nm) (Slatkin & Barton 1989) was determined using Popgene v1.32 (Yeh et al. 1997, Yeh & Boyle 1997). RESULTS The 2004 and 2005 Barnegat Bay populations show no differences in molecular weights in all eight microsatellite markers observed (Table 1), although allele frequency does differ between the two populations. We used a chord distance calculation (Cavalli-Sforza & Edwards 1967), which makes no model assumptions for the genetic distance estimate (Table 2). The chord distances indicate the genetic separation between the two BB populations and reflect the differences in allelic frequency. Although allele frequency differed among the four scallop populations, five of the eight loci studied are identical for marker size among BB, LI, and NC (Table 1). The $336 locus evidences polymorphic fragments of 138 and 158 basepairs in the NC population that are not observed in the LI or BB populations. The C1832 locus appears identical (122 bp) between LI and BB, but demonstrates polymorphisms (132 or 142 bp) in the NC population not present in BB or LI. Additionally, the NC group manifests two further alleles at the M26 locus (135 and 149 bp) not seen in BB or LI (Table 1). The genetic distances were used to generate a radial, neighbor-joining tree (Fig. 1). The North Carolina population is positioned at the end of a long branch, whereas LI falls into a clade clade Cladus, subtype Genetics A branch of biological taxa or species that share features inherited from a common ancestor; a single phylogenetic group or line. See Inheritance, Species. with the 2004/2005 Barnegat Bay groups, indicating a closer relationship among the latter cluster. The calculated genetic distance and neighbor-joining tree both support the hypothesis of a physical transfer of the Long Island bay scallop larvae down the Atlantic coast to the Atlantic Bight transition regions around Barnegat Bay. DISCUSSION Several molecular discriminatory systems have been developed to characterize bay scallop populations in the last several years, including: allozyme (Beaumont & Zouros 1991, Marelli et al. 1997, Bologna et al. 2001), mitochondrial mitochondrial pertaining to mitochondria. mitochondrial RNAs a unique set of tRNAs, mRNAs, rRNAs, transcribed from mitochondrial DNA by a mitochondrial-specific RNA polymerase, that account for about 4% of the total cell RNA that (Blake & Graves 1995, Gjetvaj et al. 1992) and Random Amplified Polymorphic DNA (RAPD RAPD Randomly Amplified Polymorphic DNA RAPD relative afferent pupillary defect (ophthalmology; aka Marcus-Gunn Pupil) ) (Beaumont 2000) markers. Unfortunately, all of these markers are inadequate for population studies because of problems relating to data interpretation, laboratory use, inadequate genotype distinction, or limited availability of polymorphisms. Roberts et al. (2005) microsatellite loci have none of these weaknesses and offer clear, easily scored polymorphisms that are quickly and directly obtained. We have taken advantage of these new markers to examine the phylogeographic origins of the recently discovered populations of bay scallops in New Jersey. [FIGURE 1 OMITTED] We have found population genetic evidence that bay scallops previously extant in New Jersey waters were most recently recruited from New York and not North Carolina. Bologna et al. (2001) established that the BB scallops sampled in 1998 were genetically similar to those of North Carolina. We hypothesize hy·poth·e·size v. hy·poth·e·sized, hy·poth·e·siz·ing, hy·poth·e·siz·es v.tr. To assert as a hypothesis. v.intr. To form a hypothesis. that this apparent discrepancy with our present results can be resolved by invoking a "subspecies shift" having occurred in Barnegat Bay. The previous New Jersey population appears to have been completely eradicated between 2000 and 2002 by brown tides. Brown tide has been shown to negatively impact the physiology of bivalves by reducing their feeding and respiration rates (Gainey & Shumway 1991, Bricelj & Lonsdale 1997). As a result, gonadal gonadal pertaining to or arising from a gonad. See also testicular, ovarian. gonadal cords cords formed by epithelial cells which migrate from the mesonephric tubules in the embryo to the gonadal ridge and establish the indifferent mass does not increase for reproduction, and the potential for population collapse increases. During the 1999 2002 brown-tide events, scallop density decreased and the species was absent from sampling regimes in 2001-2003 (Bologna unpubl, data). Our data support that the replacement population did not originate from the South, but was colonized Colonized This occurs when a microorganism is found on or in a person without causing a disease. Mentioned in: Isolation instead from the Long Island, NY population. New Jersey resides at the midAtlantic Bight in a marine transition zone, the perfect location to be potentially affected by species transfer from either direction. To invoke a subspecies replacement, an assessment of the potential larval transport mechanism is essential. Sverdrup et al. (1942) identified the southerly coastal currents along the MidAtlantic region of the United States, which provides the net transport of water masses from New York to New Jersey. Additionally, the potential tidal and Eckman transport into the coastal bays adjacent to the current provide a plausible mechanism for larval delivery into Barnegat Bay (see Epifanio & Garvine 2001). As such, the most germane possibility is transport from New York to New Jersey and this is demonstrated in our results. This extinction/replacement scenario brings up the general question of the stability of populations. How stable is a marine population living in a very dynamic environment? It is possible that the New Jersey bay scallop populations have gone extinct and been replaced many times by various source populations, because it appears to have happened at least once. We have no information on whether this cycle has occurred before in this instance, but this phenomenon has been observed for other populations (Keough & Chernoff 1987, Tettelbach & Wenczel 1993, Colson & Hughes 2004). However, the results from Bologna et al. (2001) showing a close relationship with North Carolina populations, coupled with the southerly coastal currents (Epifanio & Garvine 2001), suggests that this is the first occurrence of this phenomena in A. irradians from New Jersey. Further evidence supporting this hypothesis is obtained from our mean estimated gene flow (Nm) analysis. We calculated the Nm values (Slatkin & Barton 1989) between the various populations and found that gene flow is much higher between the BB populations and LI than between BB and NC (Table 3). Although we are employing limited population sample sizes, these Nm values may indicate a higher degree of genetic interaction between the LI and BB populations. An internal control for this conclusion can be seen in the expectedly small gene flow value between North Carolina and Long Island (Table 3). This small estimate is consistent with the results of Blake and Graves (1995) who used mitochondrial DNA polymorphisms to examine geographically isolated populations along the Eastern Seaboard. Understanding the basic population structure of any commercially exploited species is important for determining the viability of these stocks as well as the potential for interbreeding interbreeding crossbreeding, as between half-breds. among stocks. Currently, there is insufficient genetic information regarding bay scallops. The population information that we have gathered here will be essential for fisheries enhancement and aquaculture aquaculture, the raising and harvesting of fresh- and saltwater plants and animals. The most economically important form of aquaculture is fish farming, an industry that accounts for an ever increasing share of world fisheries production. (e.g., stock selection), but it will also provide a foundational assessment of New Jersey bay scallops. Given the potential of recurrent brown-tides causing broad scale extirpation ex·tir·pa·tion n. The surgical removal of an organ, part of an organ, or diseased tissue. ex  tir·pate in this species, the results of this
research will be broadly applicable to scientists, fisheries managers,
and commercial harvesters.
ACKNOWLEDGMENTS The authors thank Dr. Brad Peterson and Tina Moore for their contributions of time and effort in obtaining Long Island and North Carolina bay scallops (respectively) for us. They also thank Lisa Campanella for her help in manuscript editing. This publication was supported by the National Sea Grant College sea grant college n. A college or university that receives government grants for oceanographic research. Program of the U.S. Department of Commerce's National Oceanic and Atmospheric Administration Noun 1. National Oceanic and Atmospheric Administration - an agency in the Department of Commerce that maps the oceans and conserves their living resources; predicts changes to the earth's environment; provides weather reports and forecasts floods and hurricanes and under NOAA NOAA abbr. National Oceanic and Atmospheric Administration Noun 1. NOAA - an agency in the Department of Commerce that maps the oceans and conserves their living resources; predicts changes to the earth's environment; Grant #NA060AR4170086 (JJC JJC Joliet Junior College (Illinois) JJC John Jay College JJC Juvenile Justice Clearinghouse JJC Jurong Junior College (Jurong, Singapore) , PAXB). The views expressed herein do not necessarily reflect the views of any of those organizations. NJSG-07-662. LITERATURE CITED Beaumont, A. R. & E. Zouros. 1991. Genetics of scallops. In: S. E. Shumway, editor. Scallops: biology, ecology and aquaculture. Amsterdam: Elsevier. pp. 585-623. Beaumont, A. 2000. Genetic considerations in transfers and introductions of scallops. Aquacult Int. 8:493-512. Blake, S. G. & J. E. Graves. 1995. Mitochondrial DNA variation in the bay scallop, Argopecten irradians (Lamarck), and the calico scallop. Argopecten gibbus (Dall). J. Shellfish Res. 14:79-85. Blake, S. G., N. J. Blake, M. J. Oesterling & J. E. Graves. 1997. Genetic divergence and loss of diversity in two cultured populations of bay scallop, Argopecten irradians (Lamarck, 1819). J. Shellfish Res. 16:55-58. Bologna, P. A. X., A. E. Wilbur & K. W. Able. 2001. Reproduction. population structure, and recruitment limitation in a bay scallop (Argopecten irradians Lamarck) population from New Jersey, USA. J. Shellfish Res. 20:89-96. Bricelj, V. M. & D. J. Lonsdale. 1997. Aurecoccus anophagefferens: causes and ecological consequences of brown tides in U.S. midAtlantic coastal waters. Limnol. Oceanogr. 42:1023-1038. Campanella, J. J., V. Singh, N. Kumburis, J. Smalley & Z. Thakkar. 2004. A Molecular polymorphic analysis of Brassicaceae outgroups for use in phylogeny studies with Arabidopsis thaliana. In Vivo 25:14-25. Campanella, J. J., V. Singh, N. Kumburis, K. Vargas & A. Rios. 2003. Molecular Evidence Supporting the Asian Origin Hypothesis of Arabidopsis. In Vivo 25:4-11. Cavalli-Sforza, L. L. & A. W. F. Edwards Anthony William Fairbank Edwards (born 1935) is a British statistician, geneticist, and evolutionary biologist. He is a Life Fellow of Gonville and Caius College and retired Professor of Biometry at the University of Cambridge, and holds both the ScD and LittD degrees. A pupil of R. . 1967. Phylogenetic analysis: models and estimation procedures. Am. J. Hum. Genet genet: see civet. . 19:233-257. Clarke, A. H. 1965. The scallop superspecies Aequipecten irradians (Lamarck). Malacologia 2:161-188. Colson, I. & R. N. Hughes. 2004. Rapid recovery of genetic diversity of dogwelk (Nucella lapillus la·pil·lus n. pl. la·pil·li A small, solidified fragment of lava. [Latin, diminutive of lapis, stone. L.) populations after local extinction and recolonization contradicts predictions from life-history characteristics. Mol. Ecol. 13:2223-2233. Epifanio, C. E. & R. W. Garvine. 2001. Larval transport on the Atlantic Continental Shelf of North America: a review. East. Coast. 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Harmful Algae algae (ăl`jē) [plural of Lat. alga=seaweed], a large and diverse group of primarily aquatic plantlike organisms. These organisms were previously classified as a primitive subkingdom of the plant kingdom, the thallophytes (plants that 3:305-320. Gainey, L. F. & S. E. Shumway. 1991. The physiological effect of Aureococcus anophagefferens ("brown tide") on the lateral cilia cilia /cil·ia/ (sil´e-ah) sing. cil´ium [L.] 1. the eyelids or their outer edges. 2. the eyelashes. 3. of bivalve bivalve, aquatic mollusk of the class Pelecypoda ("hatchet-foot") or Bivalvia, with a laterally compressed body and a shell consisting of two valves, or movable pieces, hinged by an elastic ligament. mollusks. Biol. Bull. 181:298-306. Gjetvaj, B., D. I. Cook & E. Zouros. 1992. Repeated sequences and large-scale size variation of mitochondrial DNA: a common feature among scallops (Bivalvia: Pectinidae). Mol. Biol. Evol. 9:106-124. Keough, M. J. & H. Chernoff. 1987. Dispersal and Population Variation in the Bryozoan bryozoan Aquatic invertebrate of the phylum Bryozoa (“moss animals”), members (called zooids) of which form colonies. Each zooid is a complete and fully organized animal. Species range in size from a one-zooid “colony” small enough (less than 0. Bugula neritina. Ecology 68:199-210. Marelli, D. C., W. G. Lyons, W. S. Arnold & M. K. Krause. 1997. Sub-specific status of Argopecten irradians concentricus (Say, 1822) and of the bay scallops of Florida. Nautilus 110:42-44. Minch. E., A. Ruiz-Linares, D. Goldstein, M. Feldman & L. L. CavalliSforza. 1995. Microsat (version 2.0): a computer program for calculating various statistics on microsatellite allele allele (əlēl`): see genetics. allele Any one of two or more alternative forms of a gene that may occur alternatively at a given site on a chromosome. data. Stanford University Medical Center Stanford University Medical Center (Stanford Hospital & Clinics) is one of four hospitals affiliated with Stanford University and Stanford University School of Medicine, along with the Lucile Packard Children's Hospital, the Veteran's Administration Hospital in Palo Alto, and Santa , Stanford, CA, USA. Page, R. D. M. 1996. TREEVIEW: an application to display phylogenetic trees on personal computers. Comput. Appl. Biosci. 12:357-358. Provan, J. & J. J. Campanella. 2003. Patterns of Cytoplasmic cytoplasmic pertaining to or included in cytoplasm. cytoplasmic inclusions include secretory inclusions (enzymes, acids, proteins, mucosubstances), nutritive inclusions (glycogen, lipids), pigment granules (melanin, lipofuscin, Variation in Arabidopsis thaliana (Brassicaceae) revealed by polymorphic chloroplast chloroplast (klōr`əplăst', klôr`–), a complex, discrete green structure, or organelle, contained in the cytoplasm of plant cells. microsatellites. Syst. Bot. 28:578-583. Roberts, S., C. Romano & G. Gerlach. 2005. Characterization of EST EST electroshock therapy. EST abbr. electroshock therapy derived SSRs from the bay scallop, Argopecten irradians. Mol. Ecol. Notes 5:567-568. Slatkin, M. & N. H. Barton. 1989. A comparison of three indirect methods for estimating average levels of gene flow. Evolution Int. J. Org. Evolution 43:1349-1368. Sverdrup, H. U., M. W. Johnson & R. H. Fleming. 1942. The oceans: their physics, chemistry, and general biology. Old Tappan, NJ: Prentice-Hall. pp. 1087. Tettelbach, S. T. & P. Wenczel. 1993. The status of bay scallop Argopectens irradians (Lamark, 1819) populations in New York following the occurrence of "brown tide" algal blooms. J. Shellfish Res. 12:423-431. Vander Zwan, C., S. Brodie & J. J. Campanella. 2000. The Intraspecific in·tra·spe·cif·ic also in·tra·spe·cies adj. Arising or occurring within a species: intraspecific competition. Phylogenetics of Arabidopsis thaliana in World-Wide Populations. Syst. Bot. 25:47-59. Yeh, F. C., R.-C. Yang, T. J. B. Boyle, Z.-H. Ye & J. X. Mao. 1997. POPGENE, the user-friendly shareware for population genetic analysis. Molecular Biology and Biotechnology Centre, University of Alberta, Canada. Yeh, F. C. & T. J. B. Boyle. 1997. Population genetic analysis of co-dominant and dominant markers and quantitative traits. Belg. J. Bot. 129:15. JAMES J. CAMPANELLA, (1)* PAUL A. X. BOLOGNA, (1) LILIANA E. J. KIM (1) AND JOHN V. SMALLEY (2) (1) Montclair State University History Montclair State was established in 1908 as "Montclair Normal School" in response to a growing need for teachers. It was renamed "Montclair State Teachers College" in 1927, when it developed a program of educating secondary school teachers through a Bachelor of Arts , Department of Biology and Molecular Biology, 1 Normal A venue, Montclair, New Jersey 07043; (2) Bergen Community College Bergen Community College is an accredited, co-educational, two-year, public, community college located in Bergen County, New Jersey. Its primary campus is in Paramus where it was built on 9 holes of the Orchard Hills Golf Course, cutting Orchard Hills' size down in half. , Department of Science and Technology, 400 Paramus Road, Paramus, New Jersey Paramus (IPA: /pəˈræməs/) is a borough in Bergen County, New Jersey, United States. As of the United States 2000 Census, the borough population was 25,737. 07652 * Corresponding author. E-mail: james.campanella@montclair.edu
TABLE 1.
Microsatellite alleles and polymorphisms examined
in the bay scallop populations.
Marker BB 2004(15) BB 2005(13) LI(20) NC(20)
S336 128/168 * 128/168 128/168 128/138#/158#/168
G340 114/129 114/119 114/129 114/129
C1831 122/142 122/142 122 122/142
C1832 122 122 122 132/142#
GP63 200/223/285 200/223/285 200/223/285 200/223
N391 243/292 243/292 243/292 243/292
GL23 132 132 132 132
M26 120/157 120/157 120/157 120/135#/149#/157
Note: Polymorphisms are indicated with #.
* Molecular weights are in basepairs, the value in parenthesis = N,
polymorphisms in bold are not found in BB or LI populations, and the
"/" indicates heterologous allele sizes.
TABLE 2.
Chord Distance Matrix indicating the genetic
distance between the various bay scallop populations.
BB 2004 BB 2005 Ll NC
BB 2004 0.000
BB 2005 0.095 0.000
LI 0.129 0.108 0.000
NC 0.281 0.266 0.297 0.000
TABLE 3.
The mean estimated gene flow (Nm) matrix
indicating the Nm values between the various
bay scallop populations.
BB 2004 BB 2005 L1
BB 2004
BB 2005 18.2350
Ll 4.6550 4.8760
NC 1.0200 1.1258 0.8760
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