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Low Genetic Diversity of the Putatively Introduced, Brackish Water Hydrozoan, Blackfordia Virginica (Leptothecata: Blackfordiida

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Low Genetic Diversity of the Putatively Introduced, Brackish Water Hydrozoan, Blackfordia Virginica (Leptothecata: Blackfordiida Low genetic diversity of the putatively introduced, brackish water hydrozoan, Blackfordia virginica (Leptothecata: Blackfordiidae), throughout the United States, with a new record for Lake Pontchartrain, Louisiana Author(s): Genelle F. Harrison , Kiho Kim , and Allen G. Collins Source: Proceedings of the Biological Society of Washington, 126(2):91-102. 2013. Published By: Biological Society of Washington DOI: http://dx.doi.org/10.2988/0006-324X-126.2.91 URL: http://www.bioone.org/doi/full/10.2988/0006-324X-126.2.91 BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/ terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder. BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 126(2):91–102. 2013. Low genetic diversity of the putatively introduced, brackish water hydrozoan, Blackfordia virginica (Leptothecata: Blackfordiidae), throughout the United States, with a new record for Lake Pontchartrain, Louisiana Genelle F. Harrison*, Kiho Kim, and Allen G. Collins (GFH) Smithsonian Institution, Museum Support Center, Department of Entomology, 4210 Silver Hill Road, Suitland, Maryland 20746-2863, U.S.A., e-mail: [email protected]; (KK) Department of Environmental Science, American University, Washington, D.C. 20016, U.S.A.; (AGC) National Systematics Laboratory of NOAA’s Fisheries Service and Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. Abstract.—Despite first being described from Virginia, the widely distributed brackish water hydrozoan Blackfordia virginica is often hypoth- esized to have been introduced from the Black Sea to the United States. However, the alternative view that B. virginica was introduced to the Black Sea also persists in the literature. This study investigates the population structure of B. virginica in the United States to assess the directionality and/or the number of introduction events. During 2009 and 2010, estuaries were sampled from Delaware to Louisiana for brackish water hydromedusae. Nineteen samples of Blackfordia virginica were collected from four localities, including a channel running between St. Catherines Island and Lake Pontchartrain, Louisiana, a region for which it had not been reported prior to this study. We PCR amplified and sequenced two mitochondrial markers (COI & 16S), and one nuclear marker (ITS1). We compared data from individuals collected on the east coast of the United States with individuals collected in California. This revealed low diversity (two haplotypes with a maximal p-difference of 0.03% for COI and just a single haplotype for 16S) and no unique haplotypes at any locality. Low genetic variability, shared haplotypes in disparate localities, and a lack of unique haplotypes in any population are consistent with a founder effect, suggesting a single introduction and subsequent spread throughout the United States. Keywords: founder effect, genetic variation, Hydrozoa, invasive species, medusa Species in the cnidarian class Hydrozoa a wide range of salinities, and are preda- are relatively inconspicuous and may, tors of fish larvae, crustaceans, and other therefore, be overlooked in invasion biol- planktonic and benthic organisms (Bouil- ogy. However, the ecological impact of lon et al. 2006). As a result, neglecting these organisms could be great because Hydrozoa could cause inaccuracies in they reproduce asexually, often survive in ecological assessments of introduced spe- * Corresponding author. cies (Bouillon et al. 2004). 92 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Blackfordia virginica Mayer, 1910 was train, Louisiana. We collected and se- first described from specimens taken in an quenced 19 individuals from four estuarine environment in the United States locations around the United States: Dela- (U.S.) near Hampton Roads, Virginia ware, Virginia, Louisiana, and California. (Mayer 1910). Within the U.S., it has We sequenced two mitochondrial (16S & subsequently been found in the Napa and COI) and one nuclear rDNA regions Petaluma Rivers in San Francisco Bay, (ITS1). Because samples of B. virginica California (Mills & Sommer 1995), Coos from the Black Sea were not available for Bay in Oregon (Mills & Rees 2000), as well sequencing for this study, a direct com- as Delaware Bay (Cronin et al. 1962) and parison of the two populations was not is presumed to have been introduced to the possible. Instead, genetic variation of B. United States from the Black Sea region virginica in the U.S. was compared with (Mills & Sommer 1995). The hypothesis that of other hydrozoan species. While not that B. virginica is native to the Black Sea a perfect test, a comparatively lower level and was introduced to Virginia through of genetic variation relative to other shipping traffic prior to its discovery there hydrozoan species suggests that popula- appears to stem from an observation of tions of B. virginica in the U.S. was high abundance of B. virginica in the founded on a single introduction event Mandra swamp on the Bulgarian coast and has subsequently spread. Further- (Mills & Sommer 1995). An alternative more, the genetic structure of B. virginica hypothesis is that B. virginica is native to in the U.S. documented here is an impor- Virginia, where it was first collected and tant contribution as it provides baseline subsequently introduced to the Black Sea data for future studies of populations in (Leppa¨koski et al. 2009). Internationally, other geographic regions and insight into B. virginica has also been reported in the invasion patterns of a widely distributed Cananeia, Guaratuba, and Babitonga species. Bays, Brazil (Bardi & Marques 2009), Argentina and Uruguay (Genzano et al. ´ 2006), Mexico (Alvarez Silva et al. 2003), Materials and Methods Portugal and Spain (Chicaro et al. 2009), the Caspian Sea, which boarders Azerbai- Collection methods.—Specimens of jan, Iran, Kazakhstan, Russia and Turk- Blackfordia virginica were collected with menistan, (Leppa¨koski et al. 2009), the plankton tows in brackish estuaries. Three Loire estuary in France, the Mira estuary disparate locations in the U.S. were in Portugal, Nova Scotia, Canada (Moore sampled: the Mattaponi River, Virginia; 1987), Lake Belona in Romania, the South Slaughter Beach, Delaware, and Lake China Sea and East China Sea in the Pontchartrain, Louisiana (Table 1). Black- Fujian Province (Zhang 1982, Lin & fordia virginica from San Francisco Bay Zhang 1990), and the Ganges and Vasishta was provided to us by Dr. Mariah Meek Godvari estuaries in India (Sastry & from the University of California, Davis. Chandramohan 1989). No genetic studies Our collections include all geographic have thus far been carried out to address regions in which B. virginica had previous- the dynamics of any population of B. ly been reported in the literature, with the virginica. exception of Hampton Roads, Virginia This study investigates the population and Coos Bay, Oregon. A new collection structure of B. virginica in the U.S. to shed location in Louisiana was included. The light on the conflicting hypotheses on its newly reported B. virginica from Louisiana provenance. It also provides the first was preserved in a formalin and seawater report of B. virginica near Lake Pontchar- solution. We examined morphological VOLUME 126, NUMBER 2 93 Table 1.—Collection information for Blackfordia virginica from 2009–2010. Date Location n Salinity (ppt) Latitude Longitude 11 Jun 10 Mattaponi River, Virginia 3 5–23 36832026.6600 N76847021.6700 W 9 Jun 09 Lake Pontchartrain, Louisiana 10 5–10 30805034.4300 N89846037.6300 W 18 Jun 10 Slaughter Beach, Delaware 3 18–20 38856006.2600 N75819026.2400 W 8 Aug 09 San Francisco Bay, California 4 Unknown – – characteristics to determine whether there composed as follows: 1x reaction buffer, was phenotypic variation in the B. virgin- 1x BSA, 0.08 mM of dNTPs, 0.032 mM of ica collected at this site compared to MgCl, 0.24 lM of each primer, 1 unit of previous descriptions of the species. These taq and 2 ll of genomic DNA. The same characters include quantifying the number cycling protocol was used for 16S and of tentacles, the absence or presence of COI: 958C for 7 minutes followed by 40 pigmentation between tentacles, the num- cycles of 958C for 30 seconds, 488C for 30 ber of statocysts between tentacles, the seconds, 728C for 1 minute, followed by an mouth, oral arms, and radial canals were extension of 728C for 5 minutes, and a quantified and photographed. Photo- 108C holding temperature. The cycling graphs of nematocysts were taken using protocol for ITS1 was the same with the SPOT v. 4.6 software (Diagnostic Instru- exception of a 608C annealing tempera- ments). Measurements were taken using a ture. We verified
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