Marine Biology

, Volume 158, Issue 9, pp 1915–1927 | Cite as

Assemblage shift following population collapse of a non-indigenous bivalve in an urban lagoon

  • Jennifer L. BurnafordEmail author
  • Scottie Y. Henderson
  • Bruno Pernet
Original Paper


The quahog Mercenaria mercenaria has been introduced repeatedly to the Pacific coast of North America, but only one population is known to have become established. In the 1970s, the population of M. mercenaria at Colorado Lagoon, in Los Angeles County, California (33o46′16″N, 118o08′05″W), was estimated at more than 300,000 individuals. To determine the current status of this non-indigenous species (NIS), in 2009, we sampled 57 intertidal and 20 shallow subtidal plots, identifying and quantifying collected bivalves. No quahogs were found among the 2,490 living bivalves in our plots, though two were found intertidally outside of our plots. The M. mercenaria population has thus collapsed since 1980, but the native community has not recovered. Six of the fourteen living bivalve species we encountered were NIS; three are new records for the location, including the clam Venerupis philippinarum, which made up 87.6% of collected individuals. Though M. mercenaria is likely on its way to extinction on the US Pacific coast, the bivalve assemblage at this location remains heavily dominated by NIS.


Bivalve Intertidal Zone Subtidal Zone Population Collapse Mercenaria Mercenaria 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.



We thank T. Parker and E. Zahn for facilitating our work at Colorado Lagoon and for answering many questions about the history and management of the lagoon. Intertidal surveys would not have been possible without the help of numerous volunteers from CSU Fullerton, CSU Long Beach, and the residential neighborhood surrounding Colorado Lagoon. We are grateful to Dr. P. Valentich-Scott for verifying identifications, Dr. J. Harding for estimating the ages of two specimens of Mercenaria mercenaria, and T. Vu and L. Cano for measuring many clams. Drs. J. Carlton, C. Lowe, D. Reish, and C. Whitcraft provided much appreciated information about the site and its inhabitants. Drs. J. Carlton, F. Bulleri, C. Whitcraft, and an anonymous reviewer made very helpful comments on an earlier draft of the paper. This work was partially supported by the Departments of Biological Science of both CSU Fullerton and CSU Long Beach.


  1. Allen LG (1976) Additions to the list of fish species known from Alamitos Bay, California, based on studies in Colorado Lagoon. Calif Fish and Game 62:310–314Google Scholar
  2. Allen LG, Horn MH (1975) Abundance, diversity and seasonality of fishes in Colorado Lagoon, Alamitos Bay, California. Estuar Coast Mar Sci 3:371–380CrossRefGoogle Scholar
  3. Byers JE (2005) Marine reserves enhance abundance but not competitive impacts of a harvested nonindigenous species. Ecology 86:487–500CrossRefGoogle Scholar
  4. Carlton JT (1979) History, biogeography, and ecology of the introduced marine and estuarine invertebrates of the Pacific coast of North America. Ph.D. dissertation, University of California, DavisGoogle Scholar
  5. Carlton JT (1992) Introduced marine and estuarine mollusks of North America: an end-of-the-20th-century perspective. J Shellf Res 11:489–505Google Scholar
  6. Carlton JT (1999) Molluscan invasions in marine and estuarine communities. Malacologia 41:439–454Google Scholar
  7. Chambers Group (2004) Habitat assessment for the Colorado Lagoon restoration feasibility study for the City of Long Beach. Prepared for Moffatt and Nichol. p 25. Downloaded from
  8. Chew KK (2001) Introduction of the hard clam (Mercenaria mercenaria) to the Pacific Coast of North America with notes on its introduction to Puerto Rico, England, and France. In: Kraueter JN, Castagna M (eds) Biology of the Hard Clam. Elsevier Science, New York, pp 701–709CrossRefGoogle Scholar
  9. Coan EV, Valentich-Scott P (2007) Bivalvia. In: Carlton JT (ed) The Light and Smith Manual, 4th edn. UC Press, Berkeley, pp 807–859Google Scholar
  10. Coan EV, Valentich-Scott P, Bernard FR (2000) Bivalve Seashells of Western North America. Santa Barbara Museum of Natural History, Santa BarbaraGoogle Scholar
  11. Cohen AN, Harris LH, Bingham BL, Carlton JT, Chapman JW, Lambert CC, Lambert G, Ljubenkov JC, Murray SN, Rao LC, Reardon K, Schwindt E (2002) Project report for the Southern California exotics expedition 2000, a rapid assessment survey of exotic species in sheltered coastal waters. Report prepared for CA department of fish and game, state water resources control board, and natl fish and wildlife foundation, p 23Google Scholar
  12. Crane JM, Allen LG, Eisemann C (1975) Growth rate, distribution, and population density of the northern quahog Mercenaria mercenaria in Long Beach, California. Calif Fish and Game 61:68–81 (errata 61:261)Google Scholar
  13. Davis MA (2009) Invasion biology. Oxford University Press Inc., New York, p 244Google Scholar
  14. Doering PH (1976) A burrowing response of Mercenaria mercenaria (Linnaeus, 1758) elicited by Asterias forbesi (Desor, 1848). The Veliger 19:167–175Google Scholar
  15. FAO (2010). Fishery and Aquaculture Statistics Capture Production 2008 Yearbook. 590 pp. Downloaded from
  16. FAO (2011) Cultured aquatic species information programme. Mercenaria mercenaria. Cultured aquatic species information programme. Rome. FAO© 2004–2011. Downloaded 24 March 2011 from
  17. Forsyth R (1997) An introduction to introduced marine molluscs in the Pacific Northwest. Of Sea and Shore 19:188–190, 202Google Scholar
  18. Forsyth RG, Forsyth TJ (2001) A note on Mercenaria mercenaria in British Columbia, Canada. Festivus 33:85Google Scholar
  19. Geller JB (1999) Decline of a native mussel masked by sibling species invasion. Conserv Biol 13:661–664CrossRefGoogle Scholar
  20. Gillespie GE (2007) Distribution of non-indigenous intertidal species on the Pacific Coast of Canada. Nippon Suisan Gakkaishi 73:1133–1137CrossRefGoogle Scholar
  21. Gillespie GE, Parker M, Merilees W (1999) Distribution, abundance, biology and fisheries potential of the exotic varnish clam (Nuttallia obscurata) in British Columbia. Canadian Stock Assessment Secretariat Research Document 99/193Google Scholar
  22. Grizzle RE, Bricelj VM, Shumway SE (2001) Physiological ecology of Mercenaria mercenaria. In: Kraueter JN, Castagna M (eds) Biology of the hard clam. Elsevier Science, New York, pp 305–382CrossRefGoogle Scholar
  23. Hertz J, Hertz CM (1992) Unusual finds at Mission Bay, San Diego. Festivus 24:61–62Google Scholar
  24. Le Page SD (1996) Spatial distribution and temporal fluctuations of bivalve assemblages in Newport Bay, California. M.S. thesis, California State University, FullertonGoogle Scholar
  25. Levitan DR (1995) The ecology of fertilization in free-spawning invertebrates. In: McEdward LR (ed) Ecology of marine invertebrate larvae. CRC Press, Boca Raton, pp 123–156Google Scholar
  26. MacDonald KB (1969) Quantitative studies of salt marsh mollusc faunas from the North American Pacific coast. Ecol Mon 39:33–60CrossRefGoogle Scholar
  27. Mann R, Harding JM, Southworth MJ, Wesson JA (2005) Northern quahog (hard clam) Mercenaria mercenaria abundance and habitat use in Chesapeake Bay. J Shellf Res 24:509–516CrossRefGoogle Scholar
  28. Mikkelsen PM, Bieler R (2008) Seashells of southern Florida: living marine mollusks of the Florida Keys and adjacent regions: bivalves. Princeton University Press, Princeton, p 503Google Scholar
  29. Moffatt and Nichol (2004) Tidal and flood hydraulics study. Prepared for the City of Long Beach, p 25. Downloaded from
  30. Moffatt and Nichol (2010) Alternatives analysis report, Phase 2 study, Colorado Lagoon Restoration Project. Prepared in association with WRA and Blue Line Consulting for the City of Long Beach and Port of Long Beach, p 88. Downloaded from
  31. Murphy RC (1983) The introduced bivalve, Mercenaria mercenaria, in a shallow coastal ecosystem: 1) factors affecting its distribution, and 2) contribution to benthic community metabolism. Ph.D. dissertation, University of Southern CaliforniaGoogle Scholar
  32. Murphy RC (1985) Factors affecting the distribution of the introduced bivalve, Mercenaria mercenaria, in a California lagoon—the importance of bioturbation. J Mar Res 43:673–692CrossRefGoogle Scholar
  33. Murphy RC, Kremer JN (1985) Bivalve contribution to benthic metabolism in a California lagoon. Estuaries 8:330–341CrossRefGoogle Scholar
  34. Peterson CH (2002) Recruitment overfishing in a bivalve mollusc fishery: hard clams (Mercenaria mercenaria) in North Carolina. Can J Fish Aquat Sci 59:96–104CrossRefGoogle Scholar
  35. Ranasinghe JA, Mikel TK, Velarde RG, Weisberg SB, Montagne DE, Cadien DB, Dalkey A (2005) The prevalence of non-indigenous species in southern California embayments and their effects on benthic macroinvertebrate communities. Biol Inv 7:679–686CrossRefGoogle Scholar
  36. Roberts D, Rittschof D, Gerhart DJ, Schmidt AR, Hill LG (1989) Vertical migration of the clam Mercenaria mercenaria (L.) (Mollusca:Bivalvia): environmental correlates and ecological significance. J Exp Mar Biol Ecol 126:271–280CrossRefGoogle Scholar
  37. Sakai AK, Allendorf FW, Holt JS, Lodge DM, Molofsky J, With KA, Baughman S, Cabin RJ, Cohen JE, Ellstrand NC, McCauley DE, O’Neil P, Parker IM, Thompson JN, Weller SG (2001) The population biology of invasive species. Ann Rev Ecol Syst 32:305–332CrossRefGoogle Scholar
  38. Salchak A, Haas J (1971) Occurrence of the northern quahog, Mercenaria mercenaria, in Colorado Lagoon, Long Beach, California. Calif Fish and Game 57:126–128Google Scholar
  39. Simberloff D, Gibbons L (2004) Now you see them, now you don’t!—population crashes of established introduced species. Biol Inv 6:161–172CrossRefGoogle Scholar
  40. Turgeon DD, Quinn JF, Bogan AE, Coan EV, Hochberg FG, Lyons WG, Mikkelsen PM, Neves RJ, Roper CFE, Rosenberg G, Roth B, Scheltema A, Thompson FG, Vecchione M, Williams JD (1998) Common and scientific names of aquatic invertebrates from the United States and Canada: Mollusks. Second edition. American Fisheries Society special publication 26. American Malacological Union, Bethesda, p 526Google Scholar
  41. Tyler RM (2007) Effects of coverage by benthic seaweed mats on (northern quahog = hard clam) Mercenaria mercenaria in a eutrophic estuary. J Shellf Res 26:1021–1028CrossRefGoogle Scholar
  42. Whiteley J, Bendell-Young L (2007) Ecological implications of intertidal mariculture: observed differences in bivalve community structure between farm and reference sites. J Appl Ecol 44:495–505CrossRefGoogle Scholar
  43. Williams SL, Grosholz ED (2008) The invasive species challenge in estuarine and coastal environments: marrying management and science. Estuaries Coasts: J CERF 31:3–20CrossRefGoogle Scholar
  44. Zar JH (2010) Biostatistical analysis, 5th edn. Pearson Prentice Hall, New Jersey, p 944Google Scholar

Copyright information

© Springer-Verlag 2011

Authors and Affiliations

  • Jennifer L. Burnaford
    • 1
    Email author
  • Scottie Y. Henderson
    • 1
    • 3
  • Bruno Pernet
    • 2
  1. 1.Department of Biological ScienceCalifornia State University FullertonFullertonUSA
  2. 2.Department of Biological SciencesCalifornia State University Long BeachLong BeachUSA
  3. 3.Department of BiologyCerritos CollegeNorwalkUSA

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