Benthic ecosystems and environmental change


Marine ecosystems are highly valuable to human societies, through the provision of ecological goods and services. They are, however, changing rapidly in the face of multiple concurrent stressors, such as seawater warming, ocean acidification, nutrient and pollution input, over-fishing and the spread of non-native species. Understanding how life in the sea is responding to global change is critical if we are to effectively manage and plan for further change and, ultimately, conserve precious living marine resources for future generations.


We are a young but steadily growing research group based at the Marine Biological Association of the United Kingdom. We use a combination of traditional ecological techniques and innovative experimental approaches (lab and field) to better understand how life in the sea is changing, and to predict how it is likely to change in the future.


We try to do two things. First, we collaborate. We are ‘bucket and spade’ ecologists but we collaborate with microbiologists, physiologists, oceanographers and climate scientists. We work with like-minded researchers across Europe, Australia and elsewhere. We collaborate because complex systems and problems require complex approaches to better understand them. Second, we observe. Our team and our wider network of collaborators have spent countless days working in shallow-water marine ecosystems in recent years. We do this because appreciating spatial and temporal variability in ecological pattern and process is the first step to understanding human impacts on natural systems.

Research profiles

Benthic ecosystems and environmental change

Kelp forest ecology

Kelp forests dominate shallow rocky habitats across much of the world’s temperate coastline. As foundation species, kelps support high levels of primary productivity, magnified secondary productivity, and provide habitat for highly diverse associated assemblages. 

Kelp forests also serve as habitat and nursery grounds for socioeconomically important species of fish, crustaceans and molluscs. However, the structure and extent of kelp forests is affected by environmental change factors, including ocean warming, extreme climatic events and the spread on non-native species. 

A key research focus is to address pressing knowledge gaps in our fundamental understanding of the ecological structure and functioning of these critical marine habitats, in order to better predict how kelp populations and communities will respond to current and future environmental changes. 

Ecosystem responses to ocean warming and marine heatwaves

The global ocean has warmed significantly in recent decades. As a result, the geographical ranges of many marine species have shifted, generally polewards, with unexpected consequences for the structure and functioning of marine ecosystems.

Understanding how gradual warming trends and extreme climatic events affect marine biodiversity is a major focus of the research group. To achieve this, we are examining the physical drivers of ocean warming and quantifying regional and global patterns in warming trends and the occurrence of marine heatwaves.

We are also assessing the impacts of warming trends and events on marine populations, communities and ecosystems.  This research will improve the wider understanding and appreciation of the effects of ocean warming on marine biodiversity.

Impacts of non-native species

The spread of non-native species in marine environments poses a major threat to biodiversity and ecosystem structure and function, and costs the global economy billions of pounds each year. Some marine organisms, including some seaweeds and sea squirts, which can be translocated on ship’s hulls or in ballast water, are potent global invaders that impact aquaculture, biofouling and local biodiversity.

Understanding how the ecological performance of native versus non-native species will be influenced by environmental change is critically important. We are using novel experimental techniques to examine how warming influences both native and non-native species (e.g. ascidians, bryozoans, seaweeds) attached to submerged hard surfaces (e.g. rocky reefs, pylons, jetties).

Furthermore, we are examining how non-native species, such as the Japanese Kelp Undaria pinnatifida (‘Wakame’), may spread from artificial habitats (e.g. ports and marinas) into natural habitats (e.g. rocky reefs and invade native assemblages. The impacts of Wakame invasions on native plants and animals are poorly understood, especially in Europe, and we aim to better understand its impacts in southern England through field and laboratory experiments.

Extreme Climatic Events in Marine Ecosystems

This multi-faceted project focusses on how short-term extreme climatic events, such as marine heatwaves and anomalous storm and freshening events, impact coastal marine ecosystems. The research will examine (1) trends in the frequency and magnitude of these events; (2) the impacts of...

How will ocean warming and changes in storminess and turbidity affect the structure, productivity and resilience of UK kelp forests?

This large-scale field-based project will examine the structure and functioning of UK kelp forests along existing natural gradients in temperature, wave exposure and turbidity. The project employs scientific diving as a tool to access highly productive and diverse kelp forests on shallow...

The influence of multiple global change stressors on marine communities: a novel field approach

This project will develop new technologies (i.e. heated settlement panels) to manipulate key physical variables (e.g. temperature) in marine habitats, to better understand how environmental change may affect marine populations and communities under realistic conditions. The development of novel...

Understanding the global impacts and implications of range-shifting species in marine systems

This large, multi-national project aims to better understand the wider implications of species' range shifts in marine ecosystems. Many marine species have shifted their geographical distributions, predominantly poleward, in response to recent oceanic warming. The wider impacts of the...

Staff List

Dan Smale
MBA Research Fellow
Dan is a community ecologist with broad research interests and a particular focus on understanding how global environmental change is influencing the structure and functioning of coastal marine ecosystems.
Email: Telephone Number:
+44(0)1752 426274
Nora Salland
PhD Student
Nora is a PhD student working on climate change effects on kelp forest ecology in Dan Smale’s group. Her project is part of the INSPIRE DTP with the University of Southampton and the National Oceanography Centre.
Email: Telephone Number:
+44(0)1752 426330
Cat Wilding
Senior Research Assistant
Email: Telephone Number:
+44(0)1752 426330
Kathryn Smith
Postdoctoral Research Assistant

Katie works as a PDRA in the BEECH (Smale) Group, focussing primarily on work funded by Dan’s Future Leaders Fellowship award. Broadly, her research will explore the impacts of marine heatwaves on benthic ecosystems.

Email: Telephone Number:
+44(0)1752 426347
Sophie Corrigan
PhD Student

Sophie is a PhD student working with the University of Exeter, the MBA, and Cefas.

Larisa Lewis
PhD Student

Larisa is working in collaboration with the Cornish Seaweed Company whith her PhD focussing on developing and testing innovative approaches for UK seaweed cultivation.

Email: Telephone Number:
+44 (0)1752 426347

Peer-reviewed journal articles


Bue, M., Smale, D. A., Moore, P. J. (2020) Multiple-scale interactions structure macroinvertebrate assemblages associated with kelp understory algae. Diversity and Distributions, accepted 05/07/2020

Smale, D. A., Pessarrodona, A., King, N., Burrows, M. T., Yunnie. A., Vance, T., Moore, P. J. (2020) Environmental factors influencing primary productivity of the forest-forming kelp Laminaria hyperborea in the northeast Atlantic. Scientific Reports, accepted 05/07/2020

Holbrook, N., Sen Gupta, A., Oliver, E., Hobday, A., Benthuysen, J., Scannell, H., Smale, D. A., Wernberg, T. (2020) Keeping pace with marine heatwaves. Nature Reviews Earth and Environment, accepted 01/06/2020

King, N., McKeown, N., Smale, D. A., Bradbury, S., Jueterbock, A., Egilssdotter, H., Stamp, T., Groves, E., Moore, P., Hierarchical genetic structuring in the cool boreal kelp, Laminaria digitata: implications for conservation and management (2020). ICES Journal of Marine Science, accepted 12/03/2020

Brewin, R. J. W., Cyronak, T. Bresnahan, P. J., Andersson, A. J., Richard, J. Hammond, K., Billson, O., de Mora, L., Jackson, T., Smale, D. A., Dall'Olmo, G. (2020) Comparison of two methods for measuring sea surface temperature when surfing. Oceans 1: 6-26

Hereward, H. F. R., King, N. G., Smale, D. A. (2020) Intra-annual variability in responses of a canopy forming kelp to cumulative low tide heat stress: implications for populations at the trailing range edge. Journal of Phycology (Accepted 11/9/2019)

Smale, D. A. (2020) Impacts of ocean warming on kelp forest ecosystems. New Phytologist 225: 1447-1454


Burdett, H., Wright, H., Smale, D. A. (2019) Photophysiological responses of canopy-forming kelp species to short-term acute warming. Frontiers in Marine Science 6: a516

Oliver, E., Burrows, M., Donat, M., Sen Gupta, A., Alexander, L., Perkins-Kirkpatrick, S., Benthuysen, J., Hobday, A., Holbrook N., Moore, P., Thomsen, M., Wernberg, T., Smale, D. A. (2019) Projected marine heatwaves in the 21st century and the potential for ecological impact.  Frontiers in Marine Science 6: a734

Straub, S., Wernberg, T., Thomsen, M. S., Moore, P. J., Burrows, M. T., Harvey, B. P., Smale, D. A. (2019) Resistance, extinction, and everything in between – the diverse responses of seaweeds to marine heatwaves. Frontiers in Marine Science 6: a763

Macreadie, P. I., Anton, A., Raven, J., Beaumont, N., Connolly, R. M., Friess, D. A., Kelleway, J. J., Kennedy, H., Kuwae, T., Lavery, P. S., Lovelock, C. E., Smale, D. A., Apostolaki, E. T., Atwood, T. B., Baldock, J., Bianchi, T. S., Chmura, G. L., Eyre, B. D., Fourqurean, J. W., Hall-Spencer, J., Huxham, M., Hendriks, I. E., Krause-Jensen, D., Laffoley, D., Luisetti, T., Marbà, N., Masque, P., McGlathery, K. J., Megonigal, P. J., Murdiyarso, D., Russell, B. D., Santos, R., Serrano, O., Silliman, B. R., Watanabe, K., Carlos M. Duarte, C. M. (2019). The Future of Blue Carbon Science. Nature Communications 10: 1-13

Epstein, G., Foggo, A., Smale, D. A. (2019) Inconspicuous impacts: widespread marine invader causes subtle but significant changes in native macroalgal assemblages. Ecosphere 10: e02814

Epstein, G., Hawkins, S. J., Smale, D. A. (2019) Identifying niche and fitness dissimilarities in invaded marine macroalgal canopies within the context of contemporary coexistence theory. Scientific Reports 9: 8816

Holbrook, N. J., Scannell, H. A., Sen Gupta, Alexander, Benthuysen, J. A., Feng, M., Oliver, E. C. J., Alexander, L., Burrows, M. T., Donat, M. G., Hobday, A., Moore, P. J., Perkins-Kirkpatrick, S. E., Smale, D. A., Straub, S. C., Wernberg, T. (2019) A global assessment of marine heatwaves and their drivers. Nature Communications 10: 2624

Smale, D. A., Epstein, G., Parry, M., Attrill, M. A. (2019) Spatiotemporal variability in the structure of seagrass meadows and associated macrofaunal assemblages in southwest England (UK): using citizen science to benchmark ecological pattern. Ecology & Evolution 9: 3958-3972

Smale, D. A., Wernberg, T., Oliver, E. C. J., Thomsen, M., Harvey, B. P., Straub, S. C., Burrows, M. T., Alexander, L. V., Benthuysen, J. A., Donat, M. G., Feng, M., Hobday, A. J., Holbrook, N. J., Perkins-Kirkpatrick, S. E.,, Scannell, H. A., Sen Gupta, A., Payne, B., Moore, P. J. (2019) Marine heatwaves threaten global biodiversity and the provision of ecosystem services Nature Climate Change 9:306-312.

King, N., Wilcockson, D., Hoelters, L., Moore, P., McKeown, N., Groves, E., Smale, D. A., Stamp, T. (2019) Evidence for different thermal ecotypes in range centre and trailing edge kelp populations. Journal of Experimental Marine Biology and Ecology 515:10-17

Pessarrodona, A., Foggo, A., Smale, D. A (2019) Can ecosystem functioning be maintained despite climate-driven shifts in species composition? Insights from novel marine forests. Journal of Ecology 107: 91-104


Bates, A., Helmuth, B., Burrows, M. T., Duncan, M. I.,  Garrabou, J., Guy-Haim, T., Lima, F., Queiros, A. M., Seabra, R., Marsh, R., Belmaker, J., Bensoussan, N., Dong, Y., Mazaris, A. D., Smale, D. A., Wahl, M., Rilov, G. (2018) Biologists ignore ocean weather at their peril. Nature 560: 299-301

Epstein, G. & Smale, D. A. (2018) Between-habitat variability in the population dynamics of a global marine invader may drive management uncertainty. Marine Pollution Bulletin 137: 488-500

Hereward, H., Foggo, A., Hinckley, S., Greenwood, J. Smale, D. A. (2018) Seasonal variability in the population structure of a habitat-forming kelp and a conspicuous gastropod grazer: Do blue-rayed limpets (Patella pellucida) exert top-down pressure on Laminaria digitata populations? Journal of Experimental Marine Biology and Ecology 506: 171-181

Epstein, G. Hawkins, S. J., Smale, D. A. (2018) Removal treatments alter the recruitment dynamics of a global marine invader - Implications for management feasibility. Marine Environmental Research 140: 322-331

Teagle, H. Moore, P. J., Jenkins, H., Smale, D. A. (2018) Spatial variability in the diversity and structure of faunal assemblages associated with kelp holdfasts (Laminaria hyperborea) in the northeast Atlantic. PLoS ONE 13 (7) e0200411

Brewin, R. J. W., Smale, D. A., Moore, P. J., Dall’Olmo, G., Miller, P. I., Taylor, B. H., Smyth, T. J., Fishwick, J. R., Yang, M. (2018) Evaluating operational AVHRR Sea Surface Temperature data at the coastline using benthic temperature loggers. Remote Sensing 10 (6): 1-23

Pessarrodona, A., Moore, P. J., Sayer, M., Smale, D. A. (2018) Carbon assimilation and transfer through kelp forests in the NE Atlantic is diminished under a warmer ocean climate. Global Change Biology 24: 4386-4398

Hobday, A. J., Oliver, E. C. J., Sen Gupta, A., Benthuysen, J. A., Burrows, M. T. Donat, M. G., Holbrook, N., Moore, P. J., Thomsen, M. S., Wernberg, T., Smale, D. A. (2018) Categorizing and naming marine heatwaves. Oceanography 31: 162-173

Teagle, H. & Smale, D. A. (2018) Climate-driven substitution of habitat-forming species leads to reduced biodiversity within a temperate marine community. Diversity and Distributions 24: 1367-1380

Oliver E., Donat M.G., Burrows M.T., Moore P.J., Smale D.A., Alexander L.V., Benthuysen J.A., Feng M., Sen Gupta A., Hobday A.J., et al. (2018) Ocean warming brings longer and more frequent marine heatwaves. Nature Communications 9:1324.

Smale, D. A., Moore, P. J., Queirós, A. M., Higgs, N. D., Burrows, M. T. (2018) Appreciating interconnectivity between habitats is key to Blue Carbon management. Frontiers in Ecology and the Environment 16 (2): 71-73


Epstein, G. & Smale, D. A. (2017) Environmental and ecological factors influencing the spillover of the non-native kelp, Undaria pinnatifida, from marinas into natural rocky reef communities. Biological Invasions 20 (4): 1049-1072

King, N., McKeown, N. J., Smale, D. A., Moore, P. J. (2017) The importance of phenotypic plasticity and local adaptation in driving intraspecific variability in thermal niches of marine macrophytes. Ecography 41: 1469-1484

King, N., Wilcockson, D., Smale, D. A., Webster, R., Hoelters, L., Moore, P. J. (2017) Cumulative stress restricts niche filling potential of habitat-forming kelps in a future climate. Functional Ecology 32 (2): 288-299

Epstein, G. & Smale, D. A. (2017) Undaria pinnatifida: a case study to highlight challenges in marine invasion ecology and management. Ecology and Evolution 7: 8624–8642

Smale, D. A., Taylor, J. D., Coombs, S. H., Moore, G., Cunliffe, M. (2017) Community responses to seawater warming are conserved across diverse biological groupings and taxonomic resolutions. Proceedings of the Royal Society B: Biological Sciences 284: 20170534.

De Leij, R., Epstein, G., Brown, M., Smale, D. A. (2017) The influence of native macroalgal canopies on the distribution and abundance of the non-native kelp Undaria pinnatifida in natural reef habitats. Marine Biology 164: 156-163

Simpson, T. J. S., Smale, D. A., Justin I. McDonald, J. I., Wernberg, T. (2017) Large scale variability in the structure of sessile invertebrate assemblages in artificial habitats reveals the importance of local-scale processes. Journal of Experimental Marine Biology and Ecology 494:10-19

Smale, D. A., Wernberg, T., Vanderklift, M. (2017) Regional-scale variability in the response of benthic macroinvertebrate assemblages to a marine heatwave. Marine Ecology Progress Series 568: 17-30

Joint, I. & Smale, D. A. (2017) Marine heatwaves and optimal temperatures for microbial assemblage activity. FEMS Microbiology Ecology 93 (2): fiw243

Hargrave, M., Foggo, A., Pessarrodona, A., Smale, D. A. (2017) The effects of warming on the ecophysiology of two co-existing kelp species with contrasting distributions. Oecologia: 183: 531-543

Smale, D. A. & Moore, P. J. (2017) Variability in kelp forest structure along a latitudinal gradient in ocean temperature. Journal of Experimental Marine Biology and Ecology 486: 255–264

Teagle, H., Hawkins, S. J., Moore, P. J., Smale, D. A. (2017) The role of kelp species as biogenic habitat formers in coastal marine ecosystems. Journal of Experimental Marine Biology and Ecology 492: 81-98


Wernberg, T.Bennett, S., Babcock, R.C., de Bettignies, T., Cure, K., Depczynski, M., Dufois, F., Fromont, J., Fulton, C.J., Hovey, R.K., Harvey, E.S., Holmes, T.H., Kendrick, G.A., Radford, B., Santana-Garcon, J., Saunders, B.J., Smale, D. A., Thomsen, M.S., Tuckett, C.A., Tuya, F., Vanderklift, M.A., Wilson, S.K. (2016) Climate driven regime shift of a temperate marine ecosystem. Science 353 (6295): 169-172.

Hobday, A. J., Alexander, L. V., Perkins, S. E., Smale, D. A., Straub, S. C., Oliver, E. C., Benthuysen, J., Burrows, M. T., Donat, M. G., Feng, M., Holbrook, N. J., Moore, P. J., Scannell, H. A., Sen Gupta, A., Wernberg, T. (2016) A hierarchical approach to defining marine heatwaves. Progress in Oceanography 141: 227-238

Smale, D. A, Burrows, M. T., Evans, A. J., King, N., Sayer, M. D. J., Yunnie, A. L. E., Moore, P. J. (2016) Linking environmental variables with regional-scale variability in ecological structure and standing stock of carbon within UK kelp forests. Marine Ecology Progress Series 542:79-95

Arnold, M., Teagle, H., Brown, M., Smale, D. A. (2016) The structure of biogenic habitat and epibiotic assemblages associated with the global invasive kelp Undaria pinnatifida in comparison to native macroalgae. Biological Invasions 18: 661-676


Smale, D. A. and Vance, T. (2015) Climate-driven shifts in species’ distributions may exacerbate the impacts of storm disturbances on northeast Atlantic kelp forests. Marine and Freshwater Research 67: 65-74.

Sunday J.M., Pecl G.T., Frusher S., Hobday A.J., Hill N.A., Holbrook N.J., Edgar G.J., Stuart-Smith R.D., Barrett N.S., Wernberg T., Watson R.A., Smale D.A., Fulton E.A., Slawinski D., Feng M., Radford B.T. and Bates A.E. (2015) Species traits and climate velocity explain geographic range shifts in an ocean warming hotspot. Ecology Letters 18: 944-953

Smale, D. A., Vance, T., Yunnie, A. L. E., Widdicombe, S. (2015). Disentangling the impacts of heat wave magnitude, duration and timing on the structure and diversity of sessile marine assemblages. PeerJ, 3 e863

Marzinelli E.M., Williams S.B., Babcock R.C., Barrett N.S., Johnson C.R., Jordan A., Kendrick G.A., Pizarro O.R., Smale D.A. and Steinberg P.D. (2015) Large-scale geographic variation in distribution and abundance of Australian deep-water kelp forests. PLoS ONE, 10, e0118390.

Bates, A. E., Bird, T. J., Stuart-Smith, R. D., Wernberg, T., Sunday, J. M., Barrett, N. S., Edgar, G. J., Frusher, S., Hobday, A. J., Pecl, G. T., Smale, D. A., McCarthy, M.  (2015) Distinguishing geographical range shifts from artifacts of detectability and sampling effort. Diversity and Distributions 21: 13-22

Smale, D. A., Vance, T., Yunnie, A. L. E., Wernberg, T. (2015) The rise of Laminaria ochroleuca in the Western English Channel (UK) and comparisons with its competitor and assemblage dominant Laminaria hyperborea. Marine Ecology 36: 1033-1044


Verges, A. et al incl. Smale, D. A. (2014) The tropicalisation of temperate marine ecosystems: climate-mediated changes in herbivory and community phase shifts. Proceedings of the Royal Society B. 281: 1789

Brodie J., Williamson C.J., Smale D.A., Kamenos N.A., Mieszkowska N., Santos R., Cunliffe M., Steinke M., Yesson C., Anderson K.M., Asnaghi V., Brownlee C., Burdett H.L., Burrows M.T., Collins S., Donohue P.J.C., Harvey B., Foggo A., Noisette F., Nunes J., Ragazzola F., Raven J.A., Schmidt D.N., Suggett D., Teichberg M. & Hall-Spencer J.M. (2014). The future of the northeast Atlantic benthic flora in a high CO2 world. Ecology and Evolution, 4: 2787-2798

Bates, A. E., Pecl, G. T., Frusher S., Hobday, A. J., Wernberg, T. Smale, D. A., Sunday, J. M., Hill, N., Dulvy, N. K., Colwell, R. K., Holbrook, N., Fulton, E. A., Dirk Slawinski, D., Feng, M., Edgar, G. J., Radford, B. T., Thompson, P. A., Watson, R. A. (2014) Defining and observing climate-mediated range shifts in marine systems. Global Environmental Change, 26: 27-38

Foster, S., Smale, D. A., How, J. de Lestang, S., Brearley, A., Kendrick, G. A. (2014) Regional-scale patterns of mobile invertebrate assemblage structure on artificial habitats off Western Australia. Journal of Experimental Marine Biology and Ecology, 453: 43-53

Smale D.A. and Wernberg T. (2014) Population structure of the purple sea urchin Heliocidaris erythrogramma along a latitudinal gradient in southwest Australia. Journal of the Marine Biological Association of the United Kingdom, 94: 1033-1040

Azzarello, J., Smale, D. A., Langlois, T., Hansson, E. (2014) Linking habitat characteristics to abundance patterns of canopy-forming macroalgae and sea urchins in southwest Australia. Marine Biology Research 10: 682-693


Smale, D. A., Burrows, M., Moore, P., O’Connor, N. and Hawkins, S. (2013) Threats and knowledge gaps for ecosystem services provided by kelp forests: a northeast Atlantic perspective. Ecology and Evolution 3: 4016-4038

Smale, D. A. & Wernberg, T. (2013) Extreme climatic event drives range contraction of a habitat-forming species. Proceedings of the Royal Society B  280: 20122829

Smale, D. A. (2013) Multi-scale patterns of spatial variability in sessile assemblage structure do not alter predictably with development time. Marine Ecology Progress Series 482: 29-41.

Wernberg, T., Smale, D. A., Tuya, F. et al (2013) An extreme climatic event alters marine ecosystem structure in a global biodiversity hotspot. Nature Climate Change 3: 78-82


Smale, D. A. (2012) Extreme spatial variability in sessile assemblage development in subtidal habitats off southwest Australia (southeast Indian Ocean). Journal of Experimental Marine Biology and Ecology 438: 76-83

Smale, D.A., Kendrick, G.A., Harvey, E.S. et al (2012) Regional-scale benthic monitoring for Ecosystem-Based Fisheries Management (EBFM) using an Autonomous Underwater Vehicle (AUV). ICES Journal of Marine Science  69: 1108-1118

Rose, T., Smale, D. A. and Botting, G. (2012) The 2011 marine heat wave in Cockburn Sound, temperate Western Australia. Ocean Science 8: 545-550

Smale, D.A. and Wernberg, T (2012) Short-term in situ warming influences early development of sessile assemblages. Marine Ecology Progress Series 453: 129-136

Wernberg, T., Smale, D. A. and Thomsen, M. (2012) A decade of climate change experiments on marine organisms: procedures, patterns and problems. Global Change Biology 18, 1491–1498

Smale, D. A  and Childs, S. (2012) The occurrence of a widespread marine invader, Didemnum perlucidum (Tunicata, Ascidiacea) in Western Australia. Biological Invasions 14: 1325-1330

Smale, D.A. and Wernberg, T (2012) Ecological observations associated with an anomalous warming event at the Houtman Abrolhos Islands, Western Australia. Coral Reefs 31: 441

Jamieson, R.E., Rogers, A.D., Billett, D.S.M., Smale, D.A. and Pearce, D.A. (2012) Patterns of marine bacterioplankton biodiversity in the surface waters of the Scotia Arc, Southern Ocean. FEMS Microbiology Ecology 80:452-468

Smale D. A., Barnes, D. K. A., Barnes R. S. K., Smith D. J., Suggett, D. J. (2012) Spatial variability in the structure of intertidal crab and gastropod assemblages within the Seychelles Archipelago (Indian Ocean). Journal of Sea Research 69: 8-15


Smale, D. A., Wernberg, T., Vance, T. (2011) Community development on temperate subtidal reefs: the influences of wave energy and the stochastic recruitment of a dominant kelp. Marine Biology. 158:1757–1766

Smale, D. A., Wernberg, T., Peck, L. S. and Barnes, D. K. A. (2011) Turning on the heat: ecological response to simulated warming in the sea. PLoS One 6: e16050 1-4

Smale, D. A., Kendrick, G. A., and Wernberg, T. (2011) Subtidal macroalgal richness, diversity and turnover, at multiple spatial scales, along the southwestern Australian coastline. Estuarine, Coastal and Shelf Science 91: 224-231

Wernberg, T., Russell, B. D., Moore, P. J., Ling, S. D., Smale, D. A., Campbell, A., Coleman, M., Steinberg, P. D., Kendrick, G. A., Connell, S. D. (2011) Impacts of climate change in a global hotspot for temperate marine biodiversity and ocean warming. Journal of Experimental Marine Biology and Ecology 400: 7-16

Smale, D. A., Langlois, T., Kendrick, G. A., Meeuwig, J. Harvey, E. S. (2011) From fronds to fish: the use of indicators for ecological monitoring in marine benthic ecosystems, with case studies from temperate Western Australia. Reviews in Fish Biology and Fisheries 21: 311-337


Smale, D. A. (2010) Monitoring marine macroalgae: the influence of spatial scale on the usefulness of biodiversity surrogates. Diversity and Distributions 16: 985-995

Smale, D. A., Kendrick, G. A., and Wernberg, T. (2010) Assemblage turnover and taxonomic sufficiency of subtidal macroalgae at multiple spatial scales. Journal of Experimental Marine Biology and Ecology 384: 76-86.

Smale, D. A., Kendrick, G. A., Waddington, K. I. Van Niel, K. P., Meeuwig, J. J. and Harvey, E. S. (2010) Benthic assemblage composition on subtidal reefs along a latitudinal gradient in Western Australia. Estuarine, Coastal and Shelf Science 86: 83-92.


Smale D. A. and Wernberg T. (2009) Satellite-derived SST data as a proxy for water temperature in nearshore benthic ecology. Marine Ecology- Progress Series 387: 27-37.


Smale, D. A., Brown, K. M., Barnes, D. K. A., Fraser, K. P. P., Clarke, A. (2008) Ice scour disturbance in Antarctic shallow waters. Science 321: 371.

Smale, D. A. (2008) Ecological traits of benthic assemblages in shallow Antarctic waters: does ice scour disturbance select for small, mobile, scavengers with high dispersal potential? Polar Biology 31: 1225-1231.

Smale, D. A. and Barnes, D. K. A. (2008) Likely responses of the Antarctic benthos to climate-related changes in physical disturbance during the 21st Century, based primarily on evidence from the West Antarctic Peninsula region. Ecography 31: 289-305.

Smale, D.A., Barnes, D.K.A., Fraser, K.P.P. and Peck, L.S. (2008) Benthic community response to iceberg scouring at an intensely disturbed shallow water site at Adelaide Island, Antarctica. Marine Ecology- Progress Series 355: 85-94.

Smale, D. A. (2008) Spatial variability in the distribution of dominant shallow-water benthos at Adelaide Island, Antarctica. Journal of Experimental Marine Biology and Ecology 347: 140-148.

Smale, D.A. (2008) Continuous benthic community change along a depth gradient in Antarctic shallows: evidence of patchiness but not zonation. Polar Biology 31: 189-198

Barnes, D.K.A., Linse, K. Enderlein P. Smale, D. A. Fraser, K.P.P. and Brown, M. P. (2008) Marine richness and gradients at Deception Island, Antarctica. Antarctic Science 20: 271-280.


Smale, D. A. (2007) Ice disturbance intensity structures benthic communities in nearshore Antarctic waters. Marine Ecology- Progress Series 349: 89-102.

Smale, D.A., Barnes, D.K.A. and Fraser, K.P.P. (2007). The influence of depth, site exposure and season on the intensity of iceberg scouring in nearshore Antarctic waters. Polar Biology 30: 769-779.

Smale, D.A, Barnes, D.K.A., Fraser, K.P.P, Mann, P.J. and Brown M.P. (2007) Scavenging in Antarctica: intense variation between sites and seasons in shallow benthic necrophagy. Journal of Experimental Marine Biology and Ecology 349: 405-417.


Smale, D.A., Barnes, D.K.A. and Fraser, K.P.P. (2007) The influence of ice scour on benthic communities at three contrasting sites at Adelaide Island, Antarctica. Austral Ecology 32: 878-888

Contributions to compiled volumes

Moore, P. J. & Smale, D. A (2020) Impacts of climate change on shallow and shelf subtidal habitats, relevant to the coastal and marine environment around the UK. MCCIP Science Review 2020, 272–292

Wernberg, T., Campbell, A., Coleman, M.A., Connell, S.D., Kendrick, G.A., Moore, P.J., Russell, B.D.,Smale, D.A. & Steinberg, P.D. (2009). Macroalgae and Temperate Rocky Reefs. In: A Marine Climate Change Impacts and Adaptation Report Card for Australia 2009. Eds. Poloczanska, E.S., Hobday, A.J. & Richardson, A.J. NCCARF publication 05/09, ISBN 978-1-921609-03-9.

Wernberg, T. Smale, D.A., Verges, A., Campbell, A. H. Russell, B. D., Coleman, M. A., Ling, S. D., Steinberg, P. D., Johnson, C. R., Kendrick, G. A. & Connell, S. D. (2012) Macroalgae and Temperate Rocky Reefs. In: A Marine Climate Change Impacts and Adaptation Report Card for Australia 2012. Eds. Poloczanska, E.S., Hobday, A.J. & Richardson. ISBN: 978-0-643-10928-5

Barnes, D., Bergstrom, D. Bindschadler, R. and 36 others including Smale, D. A. (2009) The Next 100 Years. In Turner, J., Bindschadler, R., Convey, P., Di Prisco, G., Fahrbach, E., Gutt, J., Hodgson, D., Mayewski, P. and Summerhayes, C. (Eds.) Antarctic climate change and the environment (pp. 299-389). Scientific Committee on Antarctic Research, Cambridge UK.

Dr Dan Smale

Benthic Ecosytems and Environmental Change

Marine Biological Association of the UK

Citadel Hill, Plymouth, PL1 2PB

+44(0)1752 426489