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Department of Zoology



After his BSc Hons in Zoology at University College London (1965) and PhD at the University of Queensland, Brisbane (1968), Richard spent short periods at the University of Bristol and at the CERL Fawley Marine Laboratory before becoming a UTO in the Cambridge Zoology Department from 1972 to 2011.  His fieldwork is now carried out mainly from Rhodes (South Africa) and Queensland (Australia), via his honorary positions on the research staff of those universities, being then based at the Knysna Field Laboratory and Moreton Bay Research Station respectively.  He is also an Emeritus Fellow of St Catharine's College Cambridge. 

At various times in the recent past he has been Chairman of the European Union for Coastal Conservation (U.K.) and Secretary of the Estuarine & Coastal Sciences Association (of which he was one of the founders). He is also the author of several books, including An Introduction to Marine Ecology (with Roger Hughes of Bangor) [Blackwell Science 1982, 1988, 1999], The Invertebrates (with Peter Calow of Sheffield and Peter Olive of Newcastle) [Blackwell Science 1988, 1993, 2001], Estuarine Biology [Arnold 1974, 1984], Coastal Lagoons [CUP 1980] and The Brackish-Water Fauna of Northwestern Europe [CUP 1994]. He is a recipient of the Linnean Society of London's bicentenary medal. In total his publications have received >5,500 citations, with a Google Scholar h-index of 33 & an i10 of 101.


Richard is a marine and brackish-water ecologist interested in the benthic invertebrate communities of soft coastal sediments, especially in the spatial patterns of variation displayed by their biodiversity and related phenomena, from latitudinal scales at one extreme to sub-1 m ones at the other. He studies the macrofaunal assemblages that dominate intertidal seagrass beds, mangrove swamps, and bare expanses of sand and mud in a variety of geographical regions, especially in the Garden Route National Park (Western Cape, RSA) and the Moreton Bay Marine Park (Australia), but also in central Indonesia (Taman Nasional Wakatobi), Seychelles (Curieuse Marine National Park) and north-western Europe (including the north Norfolk coast and the Anse Lostrouc'h). 


Key publications: 
  • Barnes, R.S.K., Seath, J.L. 2024. The occurrence of a rich subtidal macrobenthic fauna in through-flow residential marina canals, and the potential of such systems to be managed as biodiversity assets. Ocean and Coastal Management254, 107191.
  • Barnes, R.S.K., Hamylton, S.M., Borburgh, L. 2024. Microscale dispersion of intertidal seagrass macrofauna. Marine Environmental Research, 196, 106385.
  • Barnes, R.S.K., Seath, J.L., Arendse, C.J. 2024. Differential sampling in the assessment of conservation and biodiversity merit: A comparison of the seagrass macrofauna in three nearby South African estuaries. Biodiversity and Conservation33, 509-532. 
  • Barnes, R.S.K. 2021. What does measuring species diversity in estuarine seagrass systems actually assess? Marine Environmental Research172, 105500.
  • Barnes, R.S.K. 2021. Within-species relationship of patchiness to both abundance and occupancy, as exemplified by seagrass macrobenthos.  Oecologia, 196, 1107-1117.
  • Barnes, R.S.K. 2020. Interspecific relationship of patchiness to occupancy and abundance, as exemplified by seagrass macrobenthos. Ecological Indicators121, 107083.
  • Barnes, R.S.K. & Claassens, L. 2020. Do beds of subtidal estuarine seagrass constitute a refuge for macrobenthic biodiversity threatened intertidally? Biodiversity and Conservation, 29, 3227-3244.
  • Barnes, R.S.K. 2020. Do different sympatric seagrass species support macrobenthic faunas of differing composition, abundance, biodiversity or patchiness? Marine Environmerntal Research160, 104983.
  • Barnes, R.S.K. 2019. Abundance/occupancy/patchiness relations in estuarine seagrass macrobenthos. Estuarine, Coastal and Shelf Science, 228, 106360.
  •  Barnes, R.S.K. 2019. Local patchiness of macrobenthic faunal abundance displays homogeneity across the disparate seagrass systems of an estuarine bay. Marine Environmental Research, 148, 99-107.
  • Barnes, R.S.K. & Hamylton, S.M. 2019. Isometric scaling of faunal patchiness: Seagrass macrobenthic abundance across small spatial scales. Marine Environmental Research146, 89-100.
  • Barnes, R.S.K. & Laurie, H. 2018. Seagrass macrofaunal abundance shows both multifractality and scale-invariant patchiness. Marine Environmental Research, 138, 84-95.
  • Barnes, R.S.K. 2017. Are seaward pneumatophore fringes transitional between mangrove and lower-shore system compartments? Marine Environmental Research, 125, 99-109.
  • Barnes, R.S.K. 2016. Spatial homogeneity of benthic microfaunal biodiversity across small spatial scales. Marine Environmental Research, 122, 148-157.
  • Barnes, R.S.K. & Hamylton, S. 2016. On the very edge:  faunal and functional responses to the interface between benthic seagrass and unvegetated-sand assemblages. Marine Ecology Progress Series, 553, 33-48.
  • Barnes, R.S.K. & Hamylton, S. 2015. Uniform functional structure across spatial scales in an intertidal benthic assemblage. Marine Environmental Research, 106, 82-91.
  • Barnes, R.S.K. & Barnes, M.K.S. 2014. Spatial uniformity of biodiversity is inevitable if the available species are distributed independently of each other. Marine Ecology Progress Series, 516, 263-266.
  • Barnes, R.S.K. & Barnes, M.K.S. 2014. Biodiversity differentials between the numerically-dominant macrobenthos of seagrass and adjacent unvegetated sediment in the absence of sand flat bioturbation. Marine Environmental Research, 99, 34-43.
  • Barnes, R.S.K. & Barnes, M.K.S.  2012.  Shore height and differentials between macrobenthic assemblages in vegetated and unvegetated areas of an intertidal sandflat. Estuarine, Coastal & Shelf Science, 106, 112-120.
  • Barnes, R.S.K.  2010.  A remarkable case of fiddler crab (Uca spp,) alpha diversity in Wallacea.  Hydrobiologia, 637, 249-253.
Other publications: 
  • Barnes, R.S.K. 2023. Seagrass macrobenthic biodiversity does not vary in conformity with a leaky-lagoonal confinement gradient. Marine Environmental Research, 185, 105897.
  • Barnes, R.S.K. 2022. Interspecific abundance-occupancy relations along estuarine gradients. Marine Environmental Research181, 105755.
  • Barnes, R.S.K., Claassens, L. & Seath, J. 2022. Where ecologically 'tis better to go brown than green: Enhanced seagrass macrobenthic biodiversity within the canals of a brownfield coastal marina. Biodiversity and Conservation31, 2981-2997.
  • Barnes, R.S.K. 2022. Biodiversity differentials between seagrass and adjacent bare sediment change along an estuarine gradient. Estuarine, Coastal & Shelf Science274, 107951. 
  • Barnes, R.S.K. 2022. Intraspecific abundance - occupancy - patchiness relations in the intertidal benthic macrofauna of a cool-temperate North Sea mudflat. Estuaries and Coasts, 45, 827-838. 
  • Barnes, R.S.K. 2020. Do species display characteristic intraspecific levels of patchiness in a given habitat type? The case of intertidal seagrass macrobenthos. Marine Biology167, 177.
  • Barnes, R.S.K. 2019. Spatial structure of a multi-species guild: the dominant biofilm-grazing microgastropods of seagrass. Hydrobiologia, 827, 293-307.
  • Barnes, R.S.K. 2018. Context dependency in the effect of Ulva-induced loss of seagrass cover on estuarine macrobenthic abundance and biodiversity. Aquatic Conservation, 29, 163-174.
  • Hamylton, S.M. Barnes, R.S.K. 2018. The effect of sampling effort on spatial autocorrelation in macrobenthic intertidal invertebrates. Hydrobiologia811, 239-250.
  • Barnes, R.S.K.  & Hendy, I.W. 2015. Seagrass-associated macrobenthic functional diversity and functional structure along an estuarine gradient. Estuarine, Coastal and Shelf Science, 164, 233-243.
  • Barnes, R.S.K. & Hendy, I.W. 2015. Functional uniformity underlies the common spatial structure of macrofaunal assemblages in intertidal seagrass beds. Biological Journal of the Linnean Society115, 114-126.
  • Barnes, R.S.K. 2014.  The nature and location of spatial change in species assemblages: a new approach illustrated by the seagrass macrofauna of the Knysna estuarine bay, South Africa. Transactions of the Royal Society of South Africa, 69, 75-80.
  • Barnes, R.S.K. 2014. Is spatial uniformity of soft-sediment biodiversity widespread and, if so, over what scales? Marine Ecology Progress Series, 504, 147-158.
  • Barnes, R.S.K. 2013. Spatial stability of macrobenthic seagrass biodiversity. Marine Ecology Progress Series, 493, 127-139.
  • Barnes, R.S.K. 2013. Distribution patterns of macrobenthic biodiversity in the intertidal seagrass beds of an estuarine system, and their conservation significance. Biodiversity and Conservation22, 357-372.
  • Barnes, R.S.K. & Hamylton, S. 2013. Abrupt transitions between macrobenthic faunal assemblages across seagrass bed margins. Estuarine, Coastal & Shelf Science131, 213-223.
  • Barnes, R.S.K. & Ellwood, M.D.F.  2012. Spatial variation in the macrobenthic assemblages of intertidal seagrass along the long axis of an estuary. Estuarine, Coastal & Shelf Science112, 173-182.
  • Barnes, R.S.K. & Ellwood, M.D.F. 2012.  The critical scale of small-scale spatial variation in ecological patterns and processes in intertidal macrobenthic seagrass assemblages.  Estuarine, Coastal & Shelf Science,98, 119-125.
  • 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 Research69, 8-15.
  • Clifton, J., Etienne, M., Barnes, D.K.A., Barnes, R.S.K., Suggett, D.J. & Smith, D.J. 2012. Marine conservation policy in Seychelles: Current constraints and prospects for improvement.  Marine Policy36, 823-831.
  • Vieira, S., Coelho, H., Nolasco, R., Serodio, J., Barnes, R.S.K. & Queiroga, H.  2012.  Repeated cycles of immersion and emersion amplify the crawling rhythm of the intertidal gastropod Hydrobia ulvae (Pennant, 1777). Journal of the Marine Biological Association of the U.K., 92, 565-570.
  • Barnes, R.S.K. & Barnes, M.K.S.  2011. Hierarchical scales of spatial variation in the smaller surface and near-surface macrobenthos of a subtropical intertidal seagrass system in Moreton Bay, Queensland.  Hydrobiologia673, 169-178.
  • Barnes, R.S.K. & Ellwood, M.D.F.  2011.  The significance of shore height in intertidal macrobenthic seagrass ecology and conservation. Aquatic Conservation21, 614-624.
  • Barnes, R.S.K. & Ellwood, M.D.F.  2011.  Macrobenthic assemblage structure in a cool-temperate intertidal dwarf-eelgrass bed in comparison with those in lower latitudes. Biological Journal of the Linnean Society104, 527-540.
  • Barnes, R.S.K.  2010. Spatial variation in abundance and diversity of the smaller surface and near-surface eelgrass-associated intertidal macrobenthos within a warm-temperate estuarine bay in the Garden Route National Park, RSA.  Aquatic Conservation20, 762-772.
  • Vieira, S., Coelho, H., Nolasco, R., Serodio, J., Barnes, R.S.K. & Queiroga, H.  2010.  The circatidal rhythm of the estuarine gastropod Hydrobia ulvae (Gastropoda, Hydrobiidae).  Biological Journal of the Linnean Society100, 439-450.
  • Barnes, R.S.K.  2010.  A review of the sentinel and allied crabs (Crustacea: Brachyura: Macrophthalmidae), with particular reference to the genus Macrophthalmus.  Raffles Bulletin of Zoology,  58, 31-49.
  • Barnes, R.S.K.  2010.  Regional and latitudinal variation in the diversity, dominance and abundance of microphagous microgastropods and other benthos in intertidal beds of dwarf eelgrass, Nanozostera spp. Marine Biodiversity, 40, 95- 106.
  • Barnes, D.K.A., Barnes, R.S.K., Smith, D.J. & Rothery, P. 2009. Littoral biodiversity across scales in the Seychelles, Indian Ocean. Marine Biodiversity, 39, 109-119.
  • Barnes, R.S.K., Smith, D.J., Barnes, D.K.A. & Gerlach, J. 2008. Variation in the distribution of supralittoral vegetation around an atoll cay: Desroches (Amirante Islands, Seychelles). Atoll Research Bulletin, 565, 1-6.
  • Barnes, R.S.K. 2006. Variation in feeding rate of the intertidal mudsnail Hydrobia ulvae in relation to the tidal cycle. Marine Ecology, 27, 154-159.
  • Schuwerack, P-M.M., Barnes, R.S.K., Underwood, G.J.C. & Jones, P.W.  2006.  Gender and species differences in sentinel crab (Macrophthalmus) feeding on an Indonesian mudflat. Journal of Crustacean Biology, 26, 119-123.
  • Barnes, R.S.K. 2005. Interspecific competition and rarity in mudsnails: feeding interactions between and within Hydrobia acuta neglecta and sympatric Hydrobia species. Aquatic Conservation, 15, 485-493.
  • Barnes, R.S.K. 2003. Feeding rates of continually submerged Hydrobia ulvae vary during the daylight hours. Journal of the Marine Biological Association of the U.K.83, 1273-1275.
  • Barnes, R.S.K. 2003. Interactions between benthic molluscs in a Sulawesi mangal, Indonesia: the cerithiid mud-creeper Cerithium coralium and potamidid mudwhelks, Terebralia spp.  Journal of the Marine Biological Association of the U.K., 83, 483-487.
  • Barnes, R.S.K. 2002. The occurrence and ecology of a marine hydrobiid mudsnail in the southern hemisphere:  the Knysna Estuary, South Africa. African Journal of Ecology, 40, 289-294.
  • Barnes, R.S.K. 2001. Interference competition in the intertidal mudsnail Hydrobia ulvae: egestion rates revisited. Journal of the Marine Biological Association of the U.K., 81, 491-495.
  • Barnes, R.S.K. & de Villiers, C.J. 2000. Animal abundance and food availability in coastal lagoon and intertidal marine sediments. Journal of the Marine Biological Association of the U.K., 80, 193-202. 
  • Barnes, R.S.K.  2000.  The geomorphology and ecology of the shingle-impounded coastal lagoon systems of Britain. In:  Packham, J.R., Randall, R.E., Barnes, R.S.K. & Neal, A.  (Ed.), Ecology and Geomorphology of Coastal Shingle, pp 320-335. Westbury, Otley.
  • Barnes, R.S.K. 1999. The conservation of brackish-water systems:  priorities for the 21st Century. Aquatic Conservation, 9, 523-527.
  • Barnes, R.S.K. 1999. What determines the distribution of coastal hydrobiid mudsnails within north-western Europe? Marine Ecology20,  97-110.
  • Barnes, R.S.K. 1998. The effects of movement on population density estimates of mudflat epifauna. Journal of the Marine Biological Association of the U.K., 78, 377-385.
  • Barnes, R.S.K. (Ed.) 1998. The Diversity of Living Organisms.  Blackwell Science, Oxford
  • Barnes, R.S.K. & Gandolfi, S.M. 1998. Is the lagoonal mudsnail Hydrobia neglecta rare because of competitively-induced reproductive depression and, if so, what are the implications for its conservation? Aquatic Conservation, 8, 737-744.
  • Barnes, R.S.K., Bradley, P., Calado, M., Demirayak, F., Doody, P., Granja, H., Hecker, N., Randall, R.E., Smit, C.J., Teixeira, A., Walmsley, J., Huggett, D. & Norris, K. 1997. Coastal habitats.  In:  Tucker, G.M. & Evans, M.I.,  Habitats for Birds in Europe. A Conservation Strategy for the Wider Environment, pp 93-123. BirdLife International, Cambridge.
  • Barnes, R.S.K. 1996. Breeding, recruitment and survival in a mixed intertidal population of the mudsnails Hydrobia ulvae and H. neglecta. Journal of the Marine Biological Association of the U.K., 76, 1003-1012.
  • Barnes, R.S.K. 1996. European coastal lagoons: values and threats. In: Tekke, R.M.H. (Ed.), Management of Coastal Lagoons in Albania, pp 141-146. EUCC, Leiden.
  • Barnes, R.S.K. 1995. European coastal lagoons: macrotidal versus microtidal contrasts. Biologia Marina Mediterranea, 2(2), 3-7.
  • Barnes, R.S.K. 1994.  A critical appraisal of the application of Guélorget and Perthuisot's concepts of the 'paralic ecosystem' and 'confinement' to macrotidal Europe. Estuarine, Coastal & Shelf Science, 38, 41-48.
  • Barnes, R.S.K. 1994. Macrofaunal community structure and life histories in coastal lagoons. Elsevier Oceanography Series, 60, 311-362 [Kjerfve, B. (Ed.) Coastal Lagoon Processes].
  • Barnes, R.S.K. 1994. Investment in eggs in lagoonal Hydrobia ventrosa  and a comparison of life-history strategy in north-west European Hydrobia species. Journal of the Marine Biological Association of the U.K., 74, 637-650.
  • Barnes, R.S.K. 1993. Reproductive strategies in contrasting populations of the coastal gastropod Hydrobia.  III.  Lagoonal versus intertidal-marine H. neglecta. Vie et Milieu, 43, 73-83.
  • Barnes, R.S.K. 1991. Dilemmas in the theory and practice of biological conservation as exemplified by British coastal lagoons. Biological Conservation, 55, 315-328.
  • Barnes, R.S.K. & Mann, K.H. (Ed.) 1991. Fundamentals of Aquatic Ecology.  Blackwell Science, Oxford   
  • Barnes, R.S.K. 1991. On the distribution of the northwest European species of the gastropod Hydrobia, with particular reference to H. neglecta.  Journal of Conchology34, 59-62.
  • Barnes, R.S.K. 1991. European estuaries and lagoons:  A personal overview of problems and possibilities for conservation and management.  Aquatic Conservation, 1, 79-87.
  • Barnes, R.S.K. 1990. Reproductive strategies in contrasting populations of the coastal gastropod Hydrobia ulvae.  II. Longevity and life-time egg production. Journal of Experimental Marine Biology & Ecology, 138, 183-200.
  • Barnes, R.S.K. 1989. What, if anything, is a brackish-water fauna?  Transactions of the Royal Society of Edinburgh, Earth Sciences, 80, 235-240.
  • Barnes, R.S.K. 1989. The coastal lagoons of Britain:  An overview and conservation appraisal. Biological Conservation, 49, 295-313.
  • Barnes, R.S.K. 1988. The faunas of land-locked lagoons: chance differences and the problems of dispersal. Estuarine, Coastal & Shelf Science, 26, 309-318.
  • Barnes, R.S.K. 1988. On reproductive strategies in adjacent lagoonal and intertidal-marine populations of the gastropod Hydrobia ulvaeJournal of the Marine Biological Association of the U.K., 68, 365-375.
  • Barnes, R.S.K. 1988. Influence of the presence of congeneric species on the behavioural preferences of Hydrobia  species.  NATO Advanced Study Institutes, A: Life Sciences, 151, 255-263.
  • Barnes, R.S.K. 1986. Daily activity rhythms in the intertidal gastropod Hydrobia ulvae (Pennant). Estuarine, Coastal & Shelf Science, 22, 325-334.
  • Barnes, R.S.K. (Ed.) 1984. A Synoptic Classification of Living Organisms.  Blackwell Science, Oxford 
  • Barnes, R.S.K. 1981. Factors effecting climbing in the coastal gastropod Hydrobia ulvae (Pennant). Journal of the Marine Biological Association of the U.K., 61, 301-306.
  • Barnes, R.S.K. 1981. An experimental study of the pattern and significance of the climbing behaviour of Hydrobia ulvae  (Pennant). Journal of the Marine Biological Association of the U.K., 61, 285-299.
  • Barnes, R.S.K. 1981. Behavioural activities and ecological strategies in the intertidal gastropod Hydrobia ulvae. In: Jones, N.V. & Wolff, W.J. (Ed.) Feeding and Survival Strategies of Estuarine Organisms, pp. 79-90. Plenum, New York.
  • Barnes, R.S.K. & Mann, K.H. (Ed.) 1980. Fundamentals of Aquatic Ecosystems.  Blackwell Science, Oxford
  • Barnes, R.S.K. 1979. Intrapopulation variation in Hydrobia sediment preferences. Estuarine & Coastal Marine Science, 9, 231-234.
  • Crawford, R.M., Dorey, A.E., Little, C. & Barnes, R.S.K. 1979. An ecological study of the Swanpool, Falmouth   V.  Phytoplankton and nutrients. Estuarine and Coastal Marine Science9, 135-160.
  • Barnes, R.S.K., Williams, A., Little, C. & Dorey, A.E. 1979. An ecological study of the Swanpool, Falmouth IV. Population fluctuations of some dominant macrofauna. In: Jefferies, R.L. & Davy, A.J. (Ed.) Ecological Processes in Coastal Environments, pp 177-197. Blackwell, Oxford.
  • Barnes, R.S.K. 1978. Variation in paragnath number of Nereis diversicolor in relation to sediment type and salinity regime. Estuarine & Coastal Marine Science, 6, 275-283.
  • Barnes, R.S.K. & Greenwood, J.G. 1978. The response of the intertidal gastropod Hydrobia ulvae to sediments of differing particle size. Journal of Experimental Marine Biology & Ecology, 31, 43-54.
  • Barnes, R.S.K. & Head, S.M. 1977. Variation in paragnath number in some British populations of the estuarine polychaete Nereis diversicolor. Estuarine and Coastal Marine Science, 5, 771-781.
  • Barnes, R.S.K.  (Ed.) 1977. The Coastline.  Wiley, London
  • Barnes, R.S.K., Sattelle, D.B., Everton, I.J., Nicholas, W. & Scott, D.H. 1976. Intertidal sands and interstitial fauna associated with different stages of salt-marsh development. Estuarine & Coastal Marine Science, 4, 497-511.
  • Barnes, R.S.K., Coughlan, J. & Holmes, N.J. 1973. A preliminary survey of the macroscopic bottom fauna of the Solent, with particular reference to Crepidula fornicata and Ostrea edulis. Proceedings of the Malacological Society, 40, 253-275.
  • Little, C., Barnes, R.S.K. & Dorey, A.E. 1973. An ecological study of the Swanpool, Falmouth III.  Origin and history. Studhyansow Kernewek, 1, 33-48.
  • Dorey, A.E., Little, C. & Barnes, R.S.K. 1973. An ecological study of the Swanpool, Falmouth II.  Hydrography and its relation to animal distributions. Estuarine and Coastal Marine Science, 1,153-176.
  • Barnes, R.S.K. & Green, J. (Ed.) 1972. The Estuarine Environment. Applied Science, London
  • Barnes, R.S.K., Dorey, A.E. & Little, C. 1971. An ecological study of a pool subject to varying salinity (Swanpool, Falmouth).  An introductory account of the topography, fauna and flora.  Journal of Animal Ecology, 40, 709-734.
  • Barnes, R.S.K. 1968. On the evolution of elongate ocular peduncles by the Brachyura.  Systematic Zoology, 17, 182-187.
  • Barnes, R.S.K. 1968. On the affinities of three fossil ocypodid crabs and their relevance to the time and place of origin of the genus Macrophthalmus (Brachyura: Ocypodidae). Journal of Zoology, London, 154, 333-339.
  • Barnes, R.S.K. 1968. Gradual acclimation effects in Macrophthalmus setosus  M.Edw. (Brachyura: Ocypodidae). Life Sciences, 7, 171-173.
  • Barnes, R.S.K. 1968. Individual variation in osmotic pressure of an ocypodid crab. Comparative Biochemistry & Physiology, 27, 447-450.
  • Barnes, R.S.K. 1967. The osmotic behaviour of a number of grapsoid crabs with respect to their differential penetration of an estuarine system. Journal of Experimental Biology, 47, 535-551.
  • Barnes, R.S.K. 1967. The Macrophthalminae of Australasia; with a review of the evolution and morphological diversity of the type genus Macrophthalmus  (Crustacea: Brachyura). Transactions of the Zoological Society of London, 31, 195-262.



Research Fellow
Hon Research Professor, Rhodes University
Hon Associate Professor, and Member of the Centre for Marine Science, University of Queensland
Hon Professor, Institute for Coastal and Marine Research, Nelson Mandela University

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