Comparisons of macrofaunal communities occupying shores across the full particle-size spectrum reveals pebble beaches to be a distinct coastal habitat type

References

• Asano T, Sato S, Lui H, Takagawa T. 2010. Experimental study on effect and advantage of beach nourishment using coarse sand. Journal of Japan Society of Civil Engineers 66: 631-635.
   Google Scholar
• Apolinario M. 1999. The role of pre-recruitment processes in the maintenance of a barnacle (Chthamalus challengeri Hoek) patch on an intertidal pebble shore in Japan. Revista Brasil Biologia 59: 225-237.
   Google Scholar
• Auguie B. 2017. gridExtra: miscellaneous functions for "grid" graphics. R package version 2.3. Available at https://CRAN.R-Project.org/package=gridExtra.
   Google Scholar
• Bally R. 1981. The ecology of three sandy beaches on the west coast of South Africa. PhD thesis, University of Cape Town, South Africa.
   Google Scholar
• Bally R. 1987. The ecology of sandy beaches of the Benguela ecosystems. South African Journal of Marine Science 5: 759-770.
   Web of Science ®Google Scholar
• Bally R, McQuaid CD, Brown AC. 1984. Shores of mixed sand and rock: an unexplored ecosystem. South African Journal of Science 80: 500-503.
   Web of Science ®Google Scholar
• Bishop MJ, McSweeney SL, Altieri AH, Keith DA. 2020. MT1.4 Boulder and cobble shores. IUCN Global ecosystem typology 2.0: Descriptive profiles for biomes and ecosystem functional groups. Gland, Switzerland: International Union for the Conservation of Nature. Available at https://portals.iucn.org/library/sites/library/files/documents/2020-037-En.pdf [accessed 2022].
   Google Scholar
• Blott SJ, Pye K. 2001. Gradistat: a grain size distribution and statistics package for the analysis of unconsolidated sediments. Earth Surface Processes and Landforms 26: 1237-1248.
   Web of Science ®Google Scholar
• Branch GM, Griffiths CL, Branch ML, Beckley LE. 2010. Two oceans: a guide to the marine life of southern Africa. Cape Town, South Africa: Struik Nature.
   Google Scholar
• Brown AC. 1971. The ecology of sandy beaches of the Cape Peninsula, South Africa. Part 1: Introduction. Transactions of the Royal Society of South Africa 39:247-279.
   Web of Science ®Google Scholar
• Bujan N, Cox R , Masselink G. 2019. From fine sands to boulders: examining the relationship between beach-face slope and sediment size. Marine Geology 417: article 106012.
   Web of Science ®Google Scholar
• Bustamante R , Branch G. 1996. Large scale patterns and trophic structure of southern African rocky shores: the roles of geographic variation and wave exposure. Journal of Biogeography 23: 339-351.
   Web of Science ®Google Scholar
• Bustamante R, Branch G, Eekhout S. 1997. The influences of physical factors on the distribution and zonation patterns of South African rocky-shore communities. South African Journal of Marine Science 18: 119-136.
   Google Scholar
• Chapman MG. 2002. Early colonization of shallow subtidal boulders in two habitats. Journal of Experimental Marine Biology and Ecology 275: 95-116.
   Web of Science ®Google Scholar
• Chapman MG. 2003. The use of sandstone blocks to test hypotheses about colonization of intertidal boulders. Journal of the Marine Biological Association of the United Kingdom 83: 415-423.
   Web of Science ®Google Scholar
• Chapman MG. 2005. Molluscs and echinoderms under boulders: test of generality of patterns of occurrence. Journal of Experimental Marine Biology and Ecology 325: 65-83.
   Web of Science ®Google Scholar
• Chapman MG. 2008. Patterns of spatial and temporal variation of macrofauna under boulders in a sheltered boulder field. Austral Ecology 27: 211-228.
   Google Scholar
• Clarke KR. 1993. Non-parametric multivariate analyses of changes in community structure. Australian Journal of Ecology 18: 117-143.
   Web of Science ®Google Scholar
• Clarke KR, Gorley RN. 2006. PRIMER (Plymouth Routines in Multivariate Ecological Research) v6: user manual/tutorial 1st edition. Plymouth, UK: PRIMER-E.
   Google Scholar
• Day JH. 1959. The biology of Langebaan Lagoon: a study of the effect of shelter from wave action. Transactions of the Royal Society of South Africa 35: 475-547.
   Google Scholar
• Defeo O, Mclachlan A. 2013. Global patterns in sandy beach macrofauna: species richness, abundance, biomass and body size. Geomorphology 199: 106-114.
   Web of Science ®Google Scholar
• de la Huz R, Lastra M, Lopez J. 2002. The influence of sediment grain size on burrowing, growth and metabolism of Donax trunculus L. (Bivalvia: Donacidae). Journal of Sea Research 47: 85-95.
   Web of Science ®Google Scholar
• Dexter OM. 1992. Sandy beach community structure: the role of exposure and latitude. Journal of Biogeography 19: 59-66.
   Web of Science ®Google Scholar
• Dinno A. 2017. dunn.test: Dunn's test of multiple comparisons using rank sums. R package version 1.3.5. Available at https://CRAN.R-projeel.org/package=dunn.test.
   Google Scholar
• Fanini L, Coleman CO, Lowry JK. 2019. Insights into the ecology of Cryptorchestia garbinii on the shores of the urban Lake Tegel (Berlin, Germany). Life and Environment 69: 187-191.
   Google Scholar
• Field JG, Griffiths CL, Griffiths RJ, Jarman N, Zoutendyk P, Velimirov B, Bowes A. 1980. Variation in structure and biomass of kelp communities along the south-west Cape coast. Transactions of the Royal Society of South Africa 44: 145-203.
   Web of Science ®Google Scholar
• Fiora SM, Carcedo MC. 2015. Influence of grain size on burrowing and alongshore distribution of the yellow clam (Amarilladesma mactroides). Journal of Shellfish Research 34: 785-789.
   Web of Science ®Google Scholar
• Forgie SA, StJohn MG, Wiser SK. 2012. Invertebrate communities and drivers of their composition on gravel beaches in New Zealand. New Zealand Journal of Ecology 37: 95-104.
   Web of Science ®Google Scholar
• Furota T, Ito T. 1999. Life cycle and environmentally induced semelparity in the shore isopod Ligia cinerascens (Ligidae) on a cobble shore along Tokyo Bay, central Japan. Journal of Crustacean Biology 19: 752-761.
   Web of Science ®Google Scholar
• Garner CJ. 2013. Characterisation and biotic classification of Eastern Cape mixed substrate shores. PhD thesis, Nelson Mandela Metropolitan University, South Africa.
   Google Scholar
• Gauci MJ, Deidun A, Schembri PJ. 2005. Faunistic diversity of Maltese pocket sandy and shingle beaches: are these of conservation value? Oceanologia 47:219-241.
   Web of Science ®Google Scholar
• Govender N. 2009. Aspects of the ecology of sandy beaches along Durban's urbanised coastline. MSc thesis, University of KwaZulu-Natal, South Africa.
   Google Scholar
• Griffiths CL, Robinson TB, Lange L, Mead A. 2010. Marine biodiversity in South Africa: an evaluation of current states of knowledge. PLoS ONE 8: e12008.
   Google Scholar
• Harris LR. 2008. The ecological implications of sea-level rise and storms for sandy beaches in KwaZulu-Natal. MSc thesis, University of KwaZulu-Natal, South Africa.
   Google Scholar
• Harris LR. 2012. An ecosystem-based spatial conservation plan for the South African sandy beaches. PhD thesis, Nelson Mandela Metropolitan University, South Africa.
   Google Scholar
• Harris LR, Nel R, Schoeman D. 2011. Mapping beach morphodynamics remotely: a novel application tested on South African sandy shores. Estuarine, Coastal and Shelf Science 92: 78-89.
   Web of Science ®Google Scholar
• Harris LR, Campbell E, Nel R, Shoeman D. 2014. Rich diversity, strong endemism, but poor protection: addressing the neglect of sandy beach ecosystems in coastal conservation planning. Diversity and Distribution 10: 1120-1135.
   Google Scholar
• Harris LR, Sink KJ, Skowno AL, van Niekerk L. 2019a. South African National Biodiversity Assessment 2018. Technical Report, vol. 5: coast. Pretoria, South Africa: South African National Biodiversity Institute. Available at http://hdl.handle.net/20.500.12143/6374 [accessed 2022].
   Google Scholar
• Harris LR, Bessinger M, Dayaram A, Holness S, Kirkman S, Livingstone TC et al. 2019b. Advancing land-sea integration for ecologically meaningful coastal conservation and management. Biological Conservation 237: 81-89.
   Web of Science ®Google Scholar
• Henninger TO, Hodgson AN. 2000. Foraging activity of Helcion pruinosus (Patellogastropoda) on a South African boulder shore. Journal of Molluscan Studies 67: 59-68.
   Web of Science ®Google Scholar
• Henninger TO, Hodgson AN. 2001. The reproductive cycle of Helcion pruinosus (Patellogastropoda) on two South African boulder shores. Journal of Molluscan Studies 67: 385-394.
   Web of Science ®Google Scholar
• Jackson LF, Lipschitz S. 1984. Coastal sensitivity atlas of Southern Africa. Cape Town, South Africa: Department of Transport.
   Google Scholar
• Jazdzewski K, De Broyer C, Pudlarz M, Dauby P. 2000. Amphipods of a stony beach in the maritime Antarctic. Polskie Archiwum Hydrobiologii 47: 569-577.
   Google Scholar
• Jażdżewski K, De Broyer C, Pudlarz M, Zielinski D. 2001. Seasonal fluctuations of vagile benthos in the uppermost sublittoral of a maritime Antarctic fjord. Polar Biology 24: 910-917.
   Web of Science ®Google Scholar
• Kurihara T. 2001. Spatial and temporal fluctuations in densities of gastropods and bivalves on subtropical cobbled shores. Bulletin of Marine Science 68: 409-426.
   Web of Science ®Google Scholar
• Kurihara T. 2002. Spatial and temporal fluctuation in the density of the intertidal limpet, Patelloida striata Quoy & Gaimard, on subtropical cobbled shores. Journal of Molluscan Studies 68: 79-86.
   Web of Science ®Google Scholar
• Kurihara T. 2007. Life-history traits of a gastropod, Nerita squamulata Le Guillou 1841, on a subtropical cobbled shore disturbed by sand. Plankton and Benthos Research 2: 213-218.
   Google Scholar
• LeHir M, Hily C. 2005. Macrofaunal diversity and habitat structure in intertidal boulder fields. Biodiversity and Conservation 14: 233-250.
   Web of Science ®Google Scholar
• Lieberman M. 1979. Ecology of subtidal algae on seasonally devastated cobble substrates off Ghana. Ecology 60: 1151-1161.
   Web of Science ®Google Scholar
• Mathers KL, Rice SP, Wood PJ. 2019. Predator, prey, and substrate interactions: the role of faunal activity and substrate characteristics. Ecosphere 10: e02545.
   Web of Science ®Google Scholar
• Matsumoto H, Young AP, Guza RT. 2019. Observations of surface cobbles at two southern California beaches. Marine Geology 419: 1-10.
   Web of Science ®Google Scholar
• McGuinness KA. 1984. Species-area relations of communities on intertidal boulders: testing the null hypothesis. Journal of Biogeography 11: 439-456.
   Web of Science ®Google Scholar
• McGuinness KA. 1987a. Disturbance and organisms on boulders. I. Patterns in the environment and the community. Oecologia 71: 409-419.
   PubMed Web of Science ®Google Scholar
• McGuinness KA. 1987b. Disturbance and organisms on boulders. II. Causes in patterns of diversity and abundance. Oecologia 71: 420-430.
   PubMed Web of Science ®Google Scholar
• McGuinness KA. 1988. Short-term effects of sessile organisms on colonization of intertidal boulders. Journal of Experimental Marine Biology and Ecology 106: 159-175.
   Google Scholar
• Mclachlan A. 1996. Physical factors in benthic ecology: effects of changing sand particle size on beach fauna. Marine Ecology Progress Series 131: 205-217.
   Web of Science ®Google Scholar
• Mclachlan A. 2001. Coastal beach ecosystems. In: Lewin R (ed.), Encyclopedia of biodiversity. New York: Academic Press. pp 741-751.
   Google Scholar
• Mclachlan A, Defeo O. 2017. The ecology of sandy shores (3rd edn). Amsterdam, The Netherlands: Academic Press.
   Google Scholar
• Mclachlan A, Jaramillo E, Donn TE, Wessels F. 1993. Sandy beach macrofauna communities and their control by the physical environments: a geographic comparison. Journal of Coastal Research 15: 27-38.
   Google Scholar
• Menge BA, Branch GM. 2001. Rocky intertidal communities. In: Bertness MD, Gaines S, Hay ME (eds), Marine community ecology. Sutherland, Massachusetts: Sinauer Associates. pp 221-257.
   Google Scholar
• Miroshnikova YM, Neradovsky YN. 2019. Pebble beaches of Murmansk coast unique formations of the Kola Peninsula [Paper]. Proceedings of the 4th International Scientific Conference "Arctic: History and Modernity", 17-18 April, Saint Petersburg, Russian Federation. lOP Conference Series: Earth and Environmental Science, vol. 302.
   Google Scholar
• Nakaoka Y, Wada K. 2017. Habitat preference of Cyclograpsus pumilio (Decapoda, Brachyura, Varunidae) inhabiting the upper intertidal limit of pebble shores. Japanese Journal of Benthology 72: 12-15.
   Google Scholar
• Nel P. 2000. Physical and biological factors structuring sandy beach macrofauna communities. PhD thesis, University of Cape Town, South Africa.
   Google Scholar
• Nhon DH, Nguyen TML, Lai TBT, Nguyen NN, Nguyen HH, Bui MO. 2019. Mineral compositions and grain sizes of sediments in intertidal zone in northern Vietnam. Vietnam Journal of Marine Science and Technology 19: 63-75 [in Vietnamese, with English abstract].
   Google Scholar
• Packman JR, Randall RE, Barnes RSK, Neal A (eds). 2001. Ecology and geomorphology of coastal shingle. Otley, UK: Westbury.
   Google Scholar
• Pubill E, Abello P, Ramon M, Baeta M. 2011. Faunistic assemblages of a sublittoral coarse sand habitat of the northwestern Mediterranean. Scientia Marina 75: 189-196.
   Web of Science ®Google Scholar
• QGIS Development Team. 2019. Quantum GIS 3.10 (A Coruna). QGIS Geographic Information System, Open Source Geospatial Foundation Project. Available at http://qgis.osgeo.org.
   Google Scholar
• R Core Team. 2013. R: a language and environment for statistical computing. Vienna, Austria: R Foundation for Statistical Computing.
   Google Scholar
• Raffaelli D, Hawkins S. 1996. Intertidal ecology. London: Chapman and Hall.
   Google Scholar
• Riccardi A, Bourget E. 1998. Weight-to-weight conversion factors for marine benthic macro-invertebrates. Marine Ecology Progress Series 163: 245-251.
   Web of Science ®Google Scholar
• Ruessink G, Brinkkemper JA, Kleinhans MG. 2015. Geometry of wave-formed orbital ripples in coarse sand. Journal of Marine Science and Engineering 3: 1568-1594.
   Google Scholar
• Russell RJ. 2006. Where most grains of very coarse sand and fine gravel are deposited. Sedimentology 11: 31-38.
   Google Scholar
• Short A. 1996. The role of wave height, period, slope, tide range and embaymentisation in beach classifications: a review. Revista Chilena de Historia Natural 69: 589-604.
   Web of Science ®Google Scholar
• Short A, Wright LD. 1983. Physical variability of sandy beaches. In: Mclachlan A, Erasmus T (eds), Sandy beaches as ecosystems. Developments in hydrobiology, vol. 19. Dordrecht, The Netherlands: Springer. pp 133-144.
   Google Scholar
• Sink K, Holness S, Harris L, Majiedt P, Atkinson L, Robinson T, Kirkman S et al. 2012. South African National Biodiversity Assessment 2011. Technical Report, vol. 4: marine and coastal component. Pretoria, South Africa: South African National Biodiversity Institute.
   Google Scholar
• Soares AG. 2003. Sandy beach morphodynamics and macrobenthic communities in temperate subtropical and tropical regions- a macro ecological approach. PhD thesis, University of Cape Town, South Africa.
   Google Scholar
• Sousa WP. 1979a. Disturbance in marine intertidal boulder fields: the nonequilibrium maintenance of species diversity. Ecology 60: 1225-1239.
   Web of Science ®Google Scholar
• Sousa WP. 1979b. Experimental investigations of disturbance and ecological succession in a rocky intertidal algal community. Ecological Monographs 49: 227-254.
   Web of Science ®Google Scholar
• Sousa WP. 1980. The responses of a community to disturbance: the importance of successional age and species life histories. Oecologia 45: 72-81.
   PubMed Web of Science ®Google Scholar
• Takada Y. 1992. Tide level variation of morph frequency and size structure in Monodonta labia (Gastropoda: Trochidae) at several boulder shores in Amakusa. Japanese Journal of Malacology 51: 187-195.
   Google Scholar
• Takada Y. 1995a. Inorganic content of faeces in molluscan grazers observed on a boulder shore at Amakusa. Japanese Journal of Malacology 54: 195-201.
   Google Scholar
• Takada Y. 1995b. Seasonal migration promoting assortative mating in Littorina brevicula on a boulder shore in Japan. Hydrobiologia 309: 151-159.
   Web of Science ®Google Scholar
• Takada Y. 1995c. Variation of growth rate with tidal level in the gastropod Monodonta labia on a boulder shore. Marine Ecology Progress Series 117: 103-110.
   Web of Science ®Google Scholar
• Takada Y. 1996. Vertical migration during the life history of the intertidal gastropod Monodonta labia on a boulder shore. Marine Ecology Progress Series 130: 117-123.
   Web of Science ®Google Scholar
• Takada Y. 2001. Activity patterns of the herbivorous gastropod Monodonta labia on a boulder shore at Amakusa, Japan. Venus 60: 15-25.
   Google Scholar
• Takada Y. 2003. Dimorphic migration, growth, and fecundity in a seasonally split population of Littorina brevicula (Mollusca: Gastropoda) on a boulder shore. Population Ecology 45: 141-148.
   Web of Science ®Google Scholar
• Takada Y. 2008. Contrasting characteristics in increasing and decreasing phases of the Nerita japonica (Mollusca: Gastropoda) population on a boulder shore. Population Ecology 50: 391-403.
   Web of Science ®Google Scholar
• Tonisson H, Orviku K, Kont A, Suursaar O, Jaagus J, Rivis R. 2007. Gravel-pebble shores on Saaremaa Island, Estonia and the relationship with formation conditions. Journal of Coastal Research 50: 810-815.
   Google Scholar
• Tucker L, Griffiths CL, Schroeter F, Vetter HD. 2017. Boulder shores in South Africa- a distinct but poorly documented coastal habitat type. African Journal of Marine Science 39: 193-202.
   Web of Science ®Google Scholar
• van Rensburg C, Robbins AE, Griffiths CL. 2021. Temperature cycles beneath , and adjacent to, intertidal boulders and associated differences in biotic composition. African Journal of Marine Science 43: 435-441.
   Web of Science ®Google Scholar
• Warnes GR , Bolker B, Bonebakker L, Gentleman R, Huber W, Liaw A et al. 2020. gplots: various R programming tools for plotting data. R package version 3.1.1. Available at https://CRAN.R-projeel.org/package=gplots.
   Google Scholar
• Wentworth CK. 1922. A scale of grade and class terms for clastic sediments. The Journal of Geology 30: 377-392.
   Google Scholar
• Wickham H. 2016. Ggplot2: elegant graphics for data analysis. New York: Springer-Verlag.
   Google Scholar
• Wickham H. 2020. forcats: tools for working with categorical variables (factors). R package version 0.5.0. Available at https://CRAN.R-project.org/package=forcats.
   Google Scholar
• Wickham H, Henry L, Pedersen TL, Luciani M, Decorde M, Lise V. 2020. svglite: an 'SVG' graphics device. R package version 1.2.3.2. Available at https://CRAN.R-project.org/package=svglite.
   Google Scholar
• Wilke CO. 2020. ggtext: improved text rendering support for 'ggplot2'. R package version 0.1.0. Available at https://CRAN.R-projeel.org/package=ggtext.
   Google Scholar
• Williams JJ, Bell PS, Betteridge K, Thorne P. 2002. Transport of coarse sand by tidal currents and waves. In: Smith JM (ed.), Coastal engineering 2002 - solving coastal conundrums. Proceedings of the 28th International Conference on Coastal Engineering, 7-12 July, Cardiff, Wales. Singapore: World Scientific.
   Google Scholar
• Williams JJ, Bell PS, Thorne P. 2003. Field measurements of flow fields and sediment transport above mobile bed forms. Journal of Geophysical Research 108: article 3109.
   PubMed Web of Science ®Google Scholar
• WoRMS Editorial Board. 2020. World Register of Marine Species. Available at http://www.marinespecies.org at VLIZ [accessed 2020].
   Google Scholar