Biogenic habitat transitions influence facilitation in a marine soft-sediment ecosystem.

Habitats are often defined by the presence of key species and biogenic features. However, the ecological consequences of interactions among distinct habitat-forming species in transition zones where their habitats overlap remain poorly understood. We investigated transition zone interactions by conducting experiments at three locations in Mahurangi Harbour, New Zealand, where the abundance of two habitat-forming marine species naturally varied. The two key species differed in form and function: One was a sessile suspension-feeding bivalve that protruded from the sediment (Atrina zelandica; Pinnidae); the other was a mobile infaunal urchin that bioturbated sediment (Echinocardium cordatum; Spatangoida). The experimental treatments established at each site reflected the natural densities of the species across sites (Atrina only, Echinocardium only, Atrina and Echinocardium together, and plots with neither species present). We identified the individual and combined effects of the two key species on sediment characteristics and co-occurring macrofauna. After five months, we documented significant treatment effects, including the highest abundance of co-occurring macrofauna in the Atrina-only treatments. However, the facilitation of macrofauna by Atrina (relative to removal treatments) was entirely negated in the presence of Echinocardium at densities >10 individuals/m2. The transitional areas in Mahurangi Harbour composed of co-occurring Atrina and Echinocardium are currently widespread and are probably more common now than monospecific patches of either individual species, due to the thinning of dense Atrina patches into sparser mixed zones during the last 10-15 years. Thus, although some ecologists avoid ecotones and habitat edges when designing experiments, suspecting that it will skew the extrapolation of results, this study increased our understanding of benthic community dynamics across larger proportions of the seascape and provided insights into temporal changes in community structure associated with patch dynamics. Particularly in situations where non-abrupt habitat transitions are commonplace, documentation of community dynamics in individual biogenic habitats and in mixed transition zones is required in order to scale-up and generalize results.

[1]  R. Rosenberg,et al.  Scale- and intensity-dependent disturbance determines the magnitude of opportunistic response , 2006 .

[2]  R. Zajac Macrobenthic biodiversity and sea floor landscape structure , 2008 .

[3]  K. Weathers,et al.  An Interdisciplinary and Synthetic Approach to Ecological Boundaries , 2003 .

[4]  Martin Solan,et al.  Extinction and Ecosystem Function in the Marine Benthos , 2004, Science.

[5]  S. Thrush,et al.  Benthic‐pelagic coupling and suspension‐feeding bivalves: Linking site‐specific sediment flux and biodeposition to benthic community structure , 2001 .

[6]  G. Coco,et al.  Feedbacks between bivalve density, flow, and suspended sediment concentration on patch stable states. , 2006, Ecology.

[7]  Sandra Díaz,et al.  Scaling environmental change through the community‐level: a trait‐based response‐and‐effect framework for plants , 2008 .

[8]  J. Jackson,et al.  Interphyletic Competition Among Marine Benthos , 1979 .

[9]  Matthew M. Yarrow,et al.  Toward Conceptual Cohesiveness: a Historical Analysis of the Theory and Utility of Ecological Boundaries and Transition Zones , 2007, Ecosystems.

[10]  G. Brenchley Mechanisms of spatial competition in marine soft-bottom communities , 1982 .

[11]  C. Heip,et al.  Biodiversity links above and below the marine sediment–water interface that may influence community stability , 2004, Biodiversity & Conservation.

[12]  S. Thrush,et al.  Disturbance to Marine Benthic Habitats by Trawling and Dredging: Implications for Marine Biodiversity , 2002 .

[13]  A. Hector,et al.  Biodiversity and ecosystem multifunctionality , 2007, Nature.

[14]  S. Pickmere,et al.  Benthic nutrient fluxes along an estuarine gradient : influence of the pinnid bivalve Atrina zelandica in summer , 2005 .

[15]  Yasuo Nakamura Autoecology of the heart urchin, Echinocardium cordatum, in the muddy sediment of the Seto Inland Sea, Japan , 2001, Journal of the Marine Biological Association of the United Kingdom.

[16]  K. Weathers,et al.  A Framework for a Theory of Ecological Boundaries , 2003 .

[17]  D. Twichell,et al.  Responses of infaunal populations to benthoscape structure and the potential importance of transition zones , 2003 .

[18]  Pierre Legendre,et al.  Integrating heterogeneity across spatial scales: interactions between Atrina zelandica and benthic macrofauna , 2002 .

[19]  S. Thrush,et al.  Indirect effects of Atrina zelandica on water column nitrogen and oxygen fluxes: The role of benthic macrofauna and microphytes , 2006 .

[20]  S. Thrush,et al.  Rapid reworking of subtidal sediments by burrowing spatangoid urchins , 2005 .

[21]  J. Ellis,et al.  Muddy waters: elevating sediment input to coastal and estuarine habitats , 2004 .

[22]  G. Brenchley Disturbance and community structure an experimental study of bioturbation in marine soft bottom environments , 1981 .

[23]  J. Ellis,et al.  Broad-scale disturbance of intertidal and shallow sublittoral soft-sediment habitats; effects on the benthic macrofauna , 2000 .

[24]  S. Thrush,et al.  Biogenic disturbance determines invasion success in a subtidal soft-sediment system. , 2008, Ecology.

[25]  J. Hewitt,et al.  Conditional outcomes of facilitation by a habitat-modifying subtidal bivalve. , 2006, Ecology.

[26]  P. Snelgrove Getting to the Bottom of Marine Biodiversity: Sedimentary Habitats: Ocean bottoms are the most widespread habitat on Earth and support high biodiversity and key ecosystem services , 1999 .

[27]  M. Posey Influence of relative mobilities on the composition of benthic communities , 1987 .

[28]  D. Rhoads,et al.  Animal-sediment relations in Cape Cod Bay, Massachusetts II. Reworking by Molpadia oolitica (Holothuroidea) , 1971 .

[29]  WILLIAM F. FAGAN,et al.  Integrating Edge Detection and Dynamic Modeling in Quantitative Analyses of Ecological Boundaries , 2003 .

[30]  W. Lauenroth,et al.  Dominant species, rather than diversity, regulates temporal stability of plant communities , 2011, Oecologia.

[31]  Simon F. Thrush,et al.  Variable effect of a large suspension-feeding bivalve on infauna: experimenting in a complex system , 2001 .

[32]  R. Rosenberg,et al.  Macrobenthic succession in relation to organic enrichment and pollution of the marine environment , 1978 .

[33]  K. Reise,et al.  Sediment destabilizing and stabilizing bio-engineers on tidal flats: cascading effects of experimental exclusion , 2009, Helgoland Marine Research.

[34]  S. Thrush,et al.  Bioturbators enhance ecosystem function through complex biogeochemical interactions , 2004, Nature.

[35]  DAVID L. STRAYER,et al.  A Classification of Ecological Boundaries , 2003 .

[36]  S. Thrush,et al.  Seabed drag coefficient over natural beds of horse mussels (Atrina zelandica) , 1998 .

[37]  R. Rosenberg,et al.  Importance of functional biodiversity and species-specific traits of benthic fauna for ecosystem functions in marine sediment , 2007 .

[38]  Pierre Legendre,et al.  Untangling Multiple Factors in Spatial Distributions: Lilies, Gophers, and Rocks , 1996 .

[39]  P. Jumars,et al.  FACILITATION OF SOFT-BOTTOM BENTHIC SUCCESSION BY TUBE BUILDERS' , 1983 .