Foundation species and invasive species strongly influence community diversity and structure, but typically in different ways. However, when widespread invasive species provide novel habitat within a community, their net effect may depend on both the environment and community composition. Fouling communities in northern and southern California harbors were surveyed to determine whether there was variation between two locations in the percent cover of an invasive bryozoan, Watersipora subtorquata (d'Orbigny, 1852), and its relationship to community diversity and composition in these two environments. Diversity significantly differed between locations and had a location-dependent association with W. subtorquata abundance. Communities were significantly dissimilar between locations, and W. subtorquata abundance had the highest percent contribution to community dissimilarity when compared to other species in the community. These results suggest that invasive species may have both facilitative and inhibitive relationships with species in a community, but that the net balance of these interactions depends on both the environmental and community context.
Angelini, C. and B.R. Silliman. 2014. Secondary foundation species as drivers of trophic and functional diversity: Evidence from a tree-epiphyte system. Ecology 95: 185–196.
Bax, N., A. Williamson, M. Aguero, E. Gonzalez and W. Geeves. 2003. Marine invasive alien species: a threat to global biodiversity. Mar. Policy 27: 313–323.
Bertness, M.D. and R. Callaway. 1994. Positive interactions in communities. Trends Ecol. Evol. 9: 187–191.
Bruno, J.F., J.J. Stachowicz and M.D. Bertness. 2003. Inclusion of facilitation into ecological theory. Trends Ecol. Evol. 18: 119–125.
Carlton, J.T. 2009. Deep invasion ecology and the assembly of communities in historical time. In: Rilov, G. and Crooks, J. A. (eds), Biological Invasions in Marine Ecosystems – Ecological, Management and Geographic Perspectives. Springer, Berlin. pp. 13–56.
Carlton, J.T. 2008. The Light and Smith Manual: Intertidal Invertebrates from Central California to Oregon. Q. Rev. Biol. 83: 130–130.
Cifuentes, M., I. Krueger, C.P. Dumont, M. Lenz and M. Thiel. 2010. Does primary colonization or community structure determine the succession of fouling communities? J. Exp. Mar. Bio. Ecol. 395: 10–20.
Colautti, R.I. and H.J. MacIsaac. 2004. A neutral terminology to define ‘invasive’ species. Divers. Distrib. 10: 135–141.
Crooks, J.A. 2002. Characterizing ecosystem-level consequences of biological invasions: the role of ecosystem engineers. Oikos 2: 153–166.
Crowl, T.A., T.O. Crist, R.R. Parmenter, G. Belovsky and A.E. Lugo. 2008. The spread of invasive species and infectious disease as drivers of ecosystem change. Front. Ecol. Environ. 6: 238–246.
Davis, K. and D.J. Marshall. 2014. Offspring size in a resident species affects community assembly. J. Anim. Ecol. 83: 322–331.
Dayton, P.K. 1972. Toward an understanding of community resilience and the potential effects of enrichment to the benthos at McMurdo Sound, Antarctica. Proc. Colloq. Conserv. Probl. Antarct. 81–96.
Dayton, P.K. and R.R. Hessler. 1972. Role of biological disturbance in maintaining diversity in the deep sea. Deep Sea Res. Oceanogr. Abstr. 19: 199–208.
Edwards, K.F. and J.J. Stachowicz. 2010. Multivariate trade-offs, succession, and phenological differentiation in a guild of colonial invertebrates. Ecology 91: 3146–3152.
Edwards, K.F. and J.J. Stachowicz. 2012. Temporally varying larval settlement, competition, and coexistence in a sessile invertebrate community. Mar. Ecol. Prog. Ser. 462: 93–102.
Floerl, O., T. Pool and G. Inglis. 2004. Positive interactions between nonindigenous species facilitate transport by human vectors. Ecol. Appl. 14: 1724–1736.
Gittenberger, A. 2009. Invasive tunicates on Zeeland and Prince Edward Island mussels, and management practices in The Netherlands. Aquat. Invasions 4: 279–281.
Gittenberger, A. 2007. Recent population expansions of non-native ascidians in The Netherlands. J. Exp. Mar. Bio. Ecol. 342: 122–126.
Graebner, R.C., R.M. Callaway and D. Montesinos. 2012. Invasive species grows faster, competes better, and shows greater evolution toward increased seed size and growth than exotic non-invasive congeners. Plant Ecol. 213: 545–553.
Graham, M.H. 2004. Effects of local deforestation on the diversity and structure of Southern California giant kelp forest food webs. Ecosystems 7: 341–357.
Holbrook, S.J., A.J. Brooks, R.J. Schmitt and H.L. Stewart. 2008. Effects of sheltering fish on growth of their host corals. Mar. Biol. 155: 521–530.
Hughes, B.B. 2010. Variable effects of a kelp foundation species on rocky intertidal diversity and species interactions in central California. J. Exp. Mar. Bio. Ecol. 393: 90–99.
Knight, N.S., C. Prentice, M. Tseng and M.I. O'Connor. 2015. A comparison of epifaunal invertebrate communities in native eelgrass Zostera marina and non-native Zostera japonica at Tsawwassen, BC. Mar. Biol. Res. 11: 564–571.
Kohler, K.E. and S.M. Gill. 2006. Coral Point Count with Excel extensions (CPCe):A Visual Basic program for the determination of coral and substrate coverage using random point count methodology. Comput. Geosci. 32: 1259–1269.
Legendre, P., D. Borcard and P.R. Peres-Neto. 2005. Analyzing beta diversity: Partitioning the spatial variation of community composition data. Ecol. Monogr. 75: 435–450.
Levine, J.M., P.B. Adler and S.G. Yelenik. 2004. A meta-analysis of biotic resistance to exotic plant invasions. Ecol. Lett. 7: 975–989.
Lockwood, J.L., M.F. Hoopes, M.P. Marchetti. 2013. Invasion Ecology. Blackwell Publishing, Malden, USA.
Lord, J., R. Whitlatch and E.D. Grosholz. 2015. Predicting competitive shifts and responses to climate change based on latitudinal distributions of species assemblages. Ecology 96: 1264–1274.
Lord, J.P. 2016. Temperature, space availability, and species assemblages impact competition in global fouling communities. Biol. Invasions.
Mackie, J.A., M.J. Keough and L. Christidis. 2006. Invasion patterns inferred from cytochrome oxidase I sequences in three bryozoans, Bugula neritina, Watersipora subtorquata, and Watersipora arcuata. Mar. Biol. 149: 285–295.
Mackie, J.A, J.A. Darling and J.B. Geller. 2012. Ecology of cryptic invasions: latitudinal segregation among Watersipora (Bryozoa) species. Sci. Rep. 2: 871.
Marasinghe, M.M.K.I., R.R.M.K.P. Ranatunga and A.C. Anil. 2018. Settlement of non-native Watersipora subtorquata (d'Orbigny, 1852) in artificial collectors deployed in Colombo Port, Sri Lanka. BioInvasions Rec. 7 (1): 7–14.
McCuller, M.I. and J.T. Carlton. 2018. Transoceanic rafting of bryozoa (Cyclostomata, cheilostomata, and ctenostomata) across the north pacific ocean on Japanese tsunami marine debris. Aquat. Invasions 13 (1): 137–162.
McKenzie, L.A, R. Brooks and E.L. Johnston. 2011. Heritable pollution tolerance in a marine invader. Environ. Res. 111: 926–32.
Mckenzie, L.A., R.C. Brooks and E.L. Johnston. 2012. A widespread contaminant enhances invasion success of a marine invader. J. Appl. Ecol. 49: 767–773.
Miranda, A.A., A.C.S. Almeida and L.M. Vieira. 2018. Non-native marine bryozoans (Bryozoa: Gymnolaemata) in Brazilian waters: Assessment, dispersal and impacts. Mar. Pollut. Bull. 130: 184–191.
Oksanen, J., F.G. Blanchet, R. Kindt, P. Legendre, R.B. O'Hara, G.L. Simpson, P. Solymos, M.H.H. Stevens and H. Wagner. 2015. vegan: Community Ecology Package. R package version 2.5-1. https://CRAN.R-project.org/package=vegan
Palardy, J.E. and J.D. Witman. 2014. Flow, recruitment limitation, and the maintenance of diversity in marine benthic communities. Ecology 95: 286–297.
Parker, I., D. Simberloff and W. Lonsdale. 1999. Impact: toward a framework for understanding the ecological effects of invaders. Biol. Invasions 1: 3–19.
Posey, M.H. 1988. Community changes associated with the spread of an introduced seagrass, Zostera japonica. Ecology 69 (4): 974–983.
Power, M.E., D. Tilman, J.A. Estes, B.A. Menge, W.J. Bond, L.S. Mills, G. Daily, J.C. Castilla, J. Lubchenco and R.T. Paine. 1996. Challenges in the quest for keystones. Bioscience 46: 609–620.
Preskitt, L.B., P.S. Vroom and C.M. Smith. 2004. A Rapid Ecological Assessment (REA) quantitative survey method for benthic algae using photoquadrats with scuba. Pacific Sci. 58: 201–209.
R Core Team (2016). R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. URL https://www.R-project.org/.
Rodriguez, L.F. 2006. Can invasive species facilitate native species? Evidence of how, when, and why these impacts occur. Biol. Invasions 8: 927–939.
Rohr, J.R., C.G. Mahan and K.C. Kim. 2009. Response of arthropod biodiversity to foundation species declines: The case of the eastern hemlock. For. Ecol. Manage. 258: 1503–1510.
Ryland, J.S. and P.J. Hayward. 1991. Marine flora and fauna of the northeastern United States: Erect Bryozoa. NOAA Tech. Rep. NMFS. 99: 1–48.
Sellheim, K., J.J. Stachowicz and R.C. Coates. 2009. Effects of a nonnative habitat-forming species on mobile and sessile epifaunal communities. Mar. Ecol. Prog. Ser. 398: 69–80.
Simberloff, D. 1995. Why do introduced species appear to devastate islands more than mainland areas? Pacific Sci. 49: 87–97.
Sorte, C.J.B., S.L. Williams and R.A. Zerebecki. 2010. Ocean warming increases threat of invasive species in a marine fouling community. Ecology 91: 2198–2204.
Stachowicz, J.J. 2001. Mutualism, facilitation, and the structure of ecological communities. Bioscience 51: 235–246.
Stachowicz, J.J., J.F. Bruno and J.E. Duffy. 2007. Understanding the effects of marine biodiversity on communities and ecosystems. Annu. Rev. Ecol. Evol. Syst. 38: 739–66.
Stachowicz, J.J. and J.E. Byrnes. 2006. Species diversity, invasion success, and ecosystem functioning: Disentangling the influence of resource competition, facilitation, and extrinsic factors. Mar. Ecol. Prog. Ser. 311: 251–262.
Stachowicz, J.J., J.R. Terwin, R.B. Whitlatch and R.W. Osman. 2002. Linking climate change and biological invasions: Ocean warming facilitates nonindigenous species invasions. Proc. Natl. Acad. Sci. U. S. A. 99: 15497–500.
Thomsen, M.S., P.A. Stæhr, L. Nejrup and D.R. Schiel. 2013. Effects of the invasive macroalgae Gracilaria vermiculophylla on two co-occurring foundation species and associated invertebrates. Aquat. Invasions 8: 133–145.
Thomsen, M.S., T. Wernberg, P.M. South and D.R. Schiel. 2016. Non-native seaweeds drive changes in marine coastal communities around the world. In: Hu, Z.M. and Fraser, C. (eds), Seaweed Phylogeography. Springer, Dordrecht. pp. 147–186.
Van Kleunen, M., E. Weber and M. Fischer. 2010. A meta-analysis of trait differences between invasive and non-invasive plant species. Ecol. Lett. 13 (2): 235–45.
Vieira, L., M. Jones and P. Taylor. 2014. The identity of the invasive fouling bryozoan Watersipora subtorquata (d'Orbigny) and some other congeneric species. Zootaxa 3857: 151–182.
Willette, D.A. and R.F. Ambrose. 2012. Effects of the invasive sea-grass Halophila stipulacea on the native seagrass, Syringodium filiforme, and associated fish and epibiota communities in the Eastern Caribbean. Aquat. Bot. 103: 74–82.
Willig, M.R. and S.J. Presley. 2018. Biodiversity and disturbance. Encyclopedia of the Anthropocene 3: 45–51.
Wyatt, A.S.J., C.L. Hewitt, D.I. Walker and T.J. Ward. 2005. Marine introductions in the Shark Bay World Heritage Property, Western Australia: A preliminary assessment. Divers. Distrib. 11: 33–44.
York, P., P. Evangelista, S. Kumar, J. Graham, C. Flather and T. Stohlgren. 2011. A habitat overlap analysis derived from maxent for tamarisk and the south-western willow flycatcher. Front. Earth Sci. 5: 120–129.
Zerebecki, R.A. and C.J.B. Sorte. 2011. Temperature tolerance and stress proteins as mechanisms of invasive species success. PLoS One 6: e14806.