Reproduction and Population Structure of Corbicula Fluminea in an Oligotrophic Subalpine Lake Author(S) :Marianne E
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Reproduction and Population Structure of Corbicula fluminea in an Oligotrophic Subalpine Lake Author(s) :Marianne E. Denton, Sudeep Chandra, Marion E. Wittmann, John Reuter and Jeffrey G. Baguley Source: Journal of Shellfish Research, 31(1):145-152. 2012. Published By: National Shellfisheries Association DOI: http://dx.doi.org/10.2983/035.031.0118 URL: http://www.bioone.org/doi/full/10.2983/035.031.0118 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. Journal of Shellfish Research, Vol. 31, No. 1, 145–152, 2012. REPRODUCTION AND POPULATION STRUCTURE OF CORBICULA FLUMINEA IN AN OLIGOTROPHIC SUBALPINE LAKE MARIANNE E. DENTON,1* SUDEEP CHANDRA,1 MARION E. WITTMANN,3† JOHN REUTER3 AND JEFFREY G. BAGULEY2 1Aquatic Ecosystems Analysis Laboratory, Department of Natural Resources and Environmental Science, University of Nevada, 1664 N. Virginia Street, MS 186, Reno, NV, 89512; 2Department of Biology, University of Nevada, 1664 N. Virginia Street, MS 314, Reno, NV 89557; 3Tahoe Environmental Research Center, University of California, One Shields Avenue, Davis, CA, 95616 ABSTRACT Reproductive effort and population structure of the nonnative clam Corbicula fluminea were studied in an oligotrophic subalpine lake. Three shallow sites (5 m) and one deeper site (20 m) were studied between May 11, 2010, and November 5, 2010, to determine spatial variation and the influence of environmental conditions (e.g., temperature and food availability as determined by total organic carbon (TOC) and sediment particulate organic matter (SPOM) on reproductive effort. The clam C. fluminea exhibited a univoltine spawn cued by increases in temperature. Reproductive effort calculated for adult clams (13.67 ± 0.03 mm (SE), n ¼ 1,875) across sites was not influenced by TOC and SPOM concentrations, and overall reproductive effort was less than more productive ecosystems, which may be a result of Lake Tahoe’s ultraoligotrophy. All 3 shallow sites had similar levels of reproductive effort. Once veligers were observed, of the 603 clams then dissected, there were 10 ± 2 veligers per clam (±SE), 25 clams had $100 veligers per clam (286 ± 28 veligers per clam), 78 clams contained less than 100 veligers (20 ± 2 veligers per clam), and 498 clams had no veligers present, indicating the population exhibits a highly variable reproductive effort. There was, at a minimum, a 4-wk delay from the point that temperatures reached a threshold for fertilization and veliger release until they were observed in dissected clams. At 20 m, C. fluminea were high in abundance compared with shallow sites, but contained few fully developed juveniles, indicating a potential population sink. Overall population structure was dominated by adult clams ($13 mm), with a minimal presence of juveniles (#4mm). KEY WORDS: clam, Corbicula fluminea, reproduction, fecundity, population structure INTRODUCTION larval incubation periods as short as 6 days, normally upward to 2 wk or as lengthy as 60 days in a wide range of environmental Nonnative aquatic species that are predisposed to reach conditions (King et al. 1986, Kraemer & Galloway 1986, nuisance levels are tolerant to a wide range of environmental McMahon 2000, Rajagopal et al. 2000). Eggs from C. fluminea conditions, able to use food and space efficiently, reach early are held in the inner demibranches of the ctenidia (gills) after sexual maturity, and/or exhibit high fecundity (e.g., Kolar & release from the gonads, then fertilized, and embryos are brooded Lodge 2001, Kulhanek et al. 2011, Karatayev et al. 2009). Within in the same structure. This may result in an annual fecundity rate a newly established area, molluscs with the greatest fecundity, of as many as 68,000 juveniles per individual (Aldridge & resulting from life history strategies such as the type of sexual McMahon 1978, McMahon 2002). Temperature initiates mul- expression, duration of brooding, and life span, are the most tiple stages of reproduction, and C. fluminea generally has a likely to become nuisance organisms (Keller et al. 2007). bivoltine reproductive cycle in response to temperature regimes The freshwater clam Corbicula fluminea, native to south- in rivers, lakes, and reservoirs (Aldridge & McMahon 1978, eastern Asia, has been introduced globally and is generally Kennedy & Van Huekelem 1985, Rajagopal et al. 2000, considered to be an aquatic invasive species of nuisance status. Mouthon & Parghentanian 2004). An initial spawn commonly After establishing in the Pacific Northwest of the United States occurs during the spring after threshold temperatures have been during the 1930s, C. fluminea spread throughout North America reached (at least 16–18°C for at least 10 degree-days); however, (McMahon 1982). Its establishment has resulted in negative once temperatures exceed 27–28°C, reproductive output is ecological and economic impacts, including colonization in restricted (McMahon 2000, Mouthon 2001, Mouthon 2001b). water intake systems of power generating systems (McMahon A subsequent, weaker spawn may occur after a return to lower tem- 2002). It can dominate the benthic biomass of aquatic ecosystems peratures (Aldridge & McMahon 1978, Kennedy & Van Huekelem (Karatayev et al. 2003) and lead to ecological changes, including 1985, Rajagopal et al. 2000, Mouthon & Parghentanian 2004). disruption of food webs because of the size-selective filtration Although temperature is the primary cue for initiation of of seston (Cohen et al. 1984, Phelps 1994, McMahon 2002), reproduction, food availability is also important for embryo suppression of native mollusc populations (Strayer 1999), and development and successful brooding (Doherty et al. 1987, alteration of nutrient cycling dynamics (Hakenkamp & Palmer Mouthon 2001b). Overall food availability has been found to 1999, Vaughn & Hakenkamp 2001). enhance gonad development and fecundity, and increases both Reproduction of C. fluminea can be prolific as a result of the brood size and individual size of developing embryos hermaphroditism, rapid reproductive maturity, and variable (Beekey & Karlson, 2003). To support growth and reproduc- *Corresponding author. E-mail: [email protected] tion, two feeding strategies are used: suspension feeding from †Current address: Department of Biological Sciences, University of the water column and deposit feeding in the substrate. Notre Dame. Galvin Life Sciences Building, Notre Dame, IN 46556 Suspension feeding rates of C. fluminea are variable but can DOI: 10.2983/035.031.0118 be high, between 300–2,500 L/h (McMahon & Bogan 2001). In 145 146 DENTON ET AL. the absence of suspended food, such as that seen in oligotro- Marla Bay is approximately 1.5 km wide with a maximum phic ecosystems, C. fluminea can ingest sediment particulate depth of 5 m before a steep drop toward profundal depths at organic matter (SPOM) through deposit feeding (Reid et al. the edge of the bay, approximately 0.50 km from the shoreline. 1992), consuming upward of 50 mg/day and doubling growth At Nevada Beach the bottom extends approximately 110 m rates (McMahon & Bogan 2001). from the shoreline to a depth of 5 m, followed by a slope to The objective of this study was to investigate the factors that greater depths. The substrate is dominated by medium sand influence the reproductive efforts (timing and overall fecundity) (0.50–0.30 mm) at both Marla Bay (>50%) and Nevada Beach of a recently established population of C. fluminea in oligotro- (>75%), with the remaining particle sizes ranging from very phic Lake Tahoe (California to Nevada). To our knowledge, fine gravel (4.00 mm) to very fine sand. Lake Tahoe is the highest elevation and deepest lake where this species has established. First observed in 2002, C. fluminea was Field Collection found to be widely established in the southeastern littoral zone of Lake Tahoe by 2008 (Hackley et al. 2008). A recent survey We collected C. fluminea using a Petite Ponar grab (area, found C. fluminea distributed in deeper waters (20–80 m). We 225 cm2) biweekly from May through August (late spring to believe clams living in deeper waters may contribute to the summer) and monthly from September through November recruitment of nearshore populations. Utilizing a combination (fall) 2010. Lake water was collected near the water–substrate of field experiments, dissections of clams, and information interface using a Van Dorn sampler and measured for in situ gathered from a literature review, we tested the following temperature using a hobbyist digital thermometer (Coralife hypotheses: (1) temperature would have the greatest influence ESU Digital Thermometer). In situ point measurements for on the timing of reproductive initiation;