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Moonshine worms (Diopatra aciculata: Onuphidae, Annelida) in the Knysna Estuary, South Africa; taxonomy and distribution

Published online by Cambridge University Press:  25 September 2020

H. van Rensburg Open the ORCID record for H. van Rensburg [Opens in a new window] ,
C. A. Matthee  and
C. A. Simon Open the ORCID record for C. A. Simon [Opens in a new window]
H. van Rensburg
Affiliation:
Department of Botany and Zoology, Stellenbosch University, Stellenbosch, South Africa
C. A. Matthee
Affiliation:
Department of Botany and Zoology, Stellenbosch University, Stellenbosch, South Africa
C. A. Simon*
Affiliation:
Department of Botany and Zoology, Stellenbosch University, Stellenbosch, South Africa
*
Author for correspondence: C. A. Simon, E-mail: [email protected]
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Abstract

Moonshine worms are a popular bait species used for fishing. The taxon was not detected during surveys of the macrobenthos conducted in Knysna in the 1940s and 1990s, and was first reported as a harvested bait species in the mid-2000s, suggesting that it appeared for the first time in the estuary in the last three decades. A previous molecular analysis identified the worms as Diopatra aciculata, a species first described from Australia. This study provides an updated detailed morphological description of D. aciculata in South Africa to facilitate future identifications and also investigates the species' distribution and population size in the Knysna Estuary. Specimens were examined by scanning electron, stereo- and compound microscopes. Diopatra aciculata has tubes that protrude from the sediment in sandy areas, often decorated with algae and shell fragments; a large body size, up to 600 mm long and 11.5 mm wide. It has 10–18 rings on ceratophores; 5–10 teeth on pectinate chaetae; uni- and bidentate pseudo-compound falcigers and dorsal cirri approximately as long as branchiae. Diopatra aciculata was detected up to 12 km from the mouth of the Knysna Estuary with densities measured at 18 sampled sites. Statistical analysis retrieved high and low density groups that were significantly different from one another (Kruskal-Wallis H(14, 800) = 376.55; P = 0.01), but distribution of high density sites was patchy. We estimate that the population comprises 20–24 million individuals. Given the size of individual worms and the population estimate, this species can be expected to have significant ecological impacts in the estuary.

Keywords

Baitcryptogenicdensitymorphologypolychaetepopulation
Type
Research Article
Information
Journal of the Marine Biological Association of the United Kingdom , Volume 100 , Issue 6 , September 2020 , pp. 897 - 907
Copyright
Copyright © Marine Biological Association of the United Kingdom 2020

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References

Allanson, BR and Herbert, DG (2005) A newly discovered population of the critically endangered false limpet Siphonaria compressa Allanson, 1958 (Pulmonata: Siphonariidae), with observations on its reproductive biology. South African Journal of Science 101, 95–97.Google Scholar
Allanson, BR, Maree, B and Grange, N (2000 a) An introduction to the chemistry of the water column of the Knysna Estuary with particular reference to nutrients and suspended solids. Transactions of the Royal Society of South Africa 55, 141–162.10.1080/00359190009520440CrossRefGoogle Scholar
Allanson, BR, Nettleton, J and de Villiers, CJ (2000 b) Benthic macrofauna richness and diversity in the Knysna Estuary: a 50 year comparison. Transactions of the Royal Society of South Africa 55, 177–185.10.1080/00359190009520442CrossRefGoogle Scholar
Allanson, BR, Human, LRD and Claassens, L (2016) Observations on the distribution and abundance of a green tide along an intertidal shore, Knysna Estuary. South African Journal of Botany 107, 49–54.10.1016/j.sajb.2016.02.197CrossRefGoogle Scholar
Arias, A, Paxton, H (2014) First record of the polychaetous annelid Diopatra micrura Pires et al., 2010 in the Mediterranean Sea. Mediterranean Marine Science 15, 5–8.10.12681/mms.377CrossRefGoogle Scholar
Arias, A and Paxton, H (2015) The cryptogenic bait worm Diopatra biscayensis Fauchald et al., 2012 (Annelida: Onuphidae) – revisiting its history, biology and ecology. Estuarine, Coastal and Shelf Science 163, 22–36.10.1016/j.ecss.2015.05.033CrossRefGoogle Scholar
Arias, A, Anadón, N and Paxton, H (2010) New records of Diopatra marocensis (Annelida: Onuphidae) from northern Spain. Zootaxa 68, 67–68.10.11646/zootaxa.2691.1.5CrossRefGoogle Scholar
Arias, A, Paxton, H and Budaeva, N (2016) Redescription and biology of Diopatra neapolitana (Annelida: Onuphidae), a protandric hermaphrodite with external spermaducal papillae. Estuarine, Coastal and Shelf Science 174, 1–17.10.1016/j.ecss.2016.03.002CrossRefGoogle Scholar
Bailey-Brock, JH (1984) Ecology of the tube-building polychaete Diopatra leuckarti Kinberg, 1865 (Onuphidae) in Hawaii: community structure, and sediment stabilizing properties. Zoological Journal of the Linnean Society 80, 191–199.10.1111/j.1096-3642.1984.tb01972.xCrossRefGoogle Scholar
Baird, D, Marais, JFK and Wooldridge, TH (1981) The influence of a marina canal system on the ecology of the Kromme estuary, St Francis Bay. South African Journal of Zoology 16, 21–34.10.1080/02541858.1981.11447729CrossRefGoogle Scholar
Bergamo, G, Carrerette, O and de Matos Nogueira, JM (2019) Continuous and non-seasonal reproductive cycle of the alien species Diopatra neapolitana (Onuphidae, Annelida) in a tropical bay of SW Atlantic. Estuarine, Coastal and Shelf Science 231.10.1016/j.ecss.2019.106479CrossRefGoogle Scholar
Berke, SK, Mahon, AR, Lima, FP, Halanych, KM, Wethey, DS and Woodin, SA (2010) Range shifts and species diversity in marine ecosystem engineers: patterns and predictions for European sedimentary habitats. Global Ecology and Biogeography 19, 223–232.10.1111/j.1466-8238.2009.00509.xCrossRefGoogle Scholar
Branch, GM and Pringle, A (1987) The impact of the sand prawn Callianassa kraussi Stebbing on sediment turnover and on bacteria, meiofauna, and benthic microflora. Journal of Experimental Marine Biology and Ecology 107, 219–235.Google Scholar
Branch, GM, Griffiths, CL, Branch, ML and Beckley, LE (2016) In Bowles, E (ed.), Two Oceans: A Guide to the Marine Life of Southern Africa. Cape Town: Penguin Random House South Africa.Google Scholar
Britz, PJ, Sauer, WHH, Mather, D, Oellerman, LK, Cowley, PD, ter Morshuizen, L and Bacela, N (2001) Baseline Study of the Utilisation of Living Marine Resources in the Eastern Cape Province. Report Prepared for the Department of Economic Affairs, Environment and Tourism, Eastern Cape Province.Google Scholar
Conti, G and Massa, F (1998) Experienze de allavamento del polichete Diopatra neapolitana Delle Chiaje, 1841 nella Laguna di S. Gilla (Sardegna Meridionale). Biologia Marina Meditteranea 5, 1473–1480.Google Scholar
Cowley, PD, Childs, AR and Bennett, HR (2013) The trouble with estuarine fisheries in temperate South Africa, illustrated by a case study on the Sundays Estuary. African Journal of Marine Science 35, 117–128.10.2989/1814232X.2013.789079CrossRefGoogle Scholar
Cunha, T, Hall, A and Queiroga, H (2005) Estimation of the Diopatra neapolitana annual harvest resulting from digging activity in Canal de Mira, Ria de Aveiro. Fisheries Research 76, 56–66.10.1016/j.fishres.2005.05.008CrossRefGoogle Scholar
Daǧli, E, Ergen, Z and Çinar, ME (2005) One-year observation on the population structure of Diopatra neapolitana Delle Chiaje (Polychaeta: Onuphidae) in Izmir Bay (Aegean sea, eastern Mediterranean). Marine Ecology 26, 265–272.10.1111/j.1439-0485.2005.00055.xCrossRefGoogle Scholar
Day, JH (1967) A monograph on the Polychaeta of Southern Africa. Part 1: Errantia. Annals of the South African Museum 45, 375–548.Google Scholar
Day, JH, Millard, NAH and Harrison, AD (1951) The ecology of South African estuaries. Part III: Knysna: a clear open estuary. Transactions of the Royal Society of South Africa 33, 367–413.10.1080/00359195109519891CrossRefGoogle Scholar
de Carvalho, AN, Vaz, ASL, Sérgio, TIB and Dos Santos, PJT (2013) Sustainability of bait fishing harvesting in estuarine ecosystems – case study in the Local Natural Reserve of Douro Estuary, Portugal. Revista de Gestão Costeira Integrada 13, 157–168.10.5894/rgci393CrossRefGoogle Scholar
Elgetany, AH, van Rensburg, H, Hektoen, M, Matthee, CA, Budaeva, N, Simon, CA and Struck, TH (2020) Species delineation in the speciation grey zone – the case of Diopatra (Annelida, Onuphidae). Zoologica Scripta zsc.12421.Google Scholar
Fauchald, K and Jumars, PA (1979) The diet of worms: a study of polychaete feeding guilds. Oceanography and Marine Biology: An Annual Review 17, 193–284.Google Scholar
Fauchald, K, Berke, SK and Woodin, SA (2012) Diopatra (Onuphidae: Polychaeta) from intertidal sediments in southwestern Europe. Zootaxa 3395, 47–58.10.11646/zootaxa.3395.1.5CrossRefGoogle Scholar
Freitas, R, Pires, A, Velez, C, Almeida, Â, Wrona, FJ, Soares, AMVM and Figueira, E (2015) The effects of salinity changes on the Polychaete Diopatra neapolitana: impacts on regenerative capacity and biochemical markers. Aquatic Toxicology 163, 167–176.10.1016/j.aquatox.2015.04.006CrossRefGoogle ScholarPubMed
Gaigher, CM (1979) Aspects of the Population Dynamics and Ecology of the Bloodworm (Arenicola loveni Kinberg). Unpublished thesis, University of Cape Town.Google Scholar
Hakkim, VMA (1975) Salinity tolerance of Diopatra neapolitana Dille Chiaje: Annelida–Polychaeta. Indian Journal of Marine Science 4, 99–101.Google Scholar
Harwell, MC and Orth, RJ (2001) Influence of a tube-dwelling polychaete on the dispersal of fragmented reproductive shoots of eelgrass. Aquatic Botany 70, 1–7.10.1016/S0304-3770(00)00147-9CrossRefGoogle Scholar
Henninger, TO and Froneman, PW (2013) Role of the sandprawn Callichirus kraussi as an ecosystem engineer in a South African temporarily open/closed estuary. African Journal of Aquatic Science 38, 101–107.10.2989/16085914.2012.750593CrossRefGoogle Scholar
Hodgson, AN, Allanson, BR and Cretchley, R (2000) The exploitation of Upogebia africana (Crustacea: Thalassinidae) for bait in the Knysna Estuary. Transactions of the Royal Society of South Africa 55, 197–204.10.1080/00359190009520444CrossRefGoogle Scholar
Huttel, M (1990) Influence of the lugworm Arenicola marina on porewater nutrient profiles of sand flat sediments. Marine Ecology Progress Series 62, 241–248.10.3354/meps062241CrossRefGoogle Scholar
Jumars, PA, Dorgan, KM and Lindsay, SM (2015) Diet of worms emended: an update of polychaete feeding guilds. Annual Review of Marine Science 7, 497–520.10.1146/annurev-marine-010814-020007CrossRefGoogle ScholarPubMed
Kara, J, Macdonald, AHH and Simon, CA (2018) Integrative taxonomic methods reveal an incorrect synonymisation of the South African Pseudonereis podocirra (Schmarda) as the widespread Pseudonereis variegata (Grube) from Chile. Invertebrate Systematics 32, 1282–1297.10.1071/IS18016CrossRefGoogle Scholar
Knox, GA and Cameron, DB (1971) Port Phillip Bay Survey 1957–1963, Victoria, Australia. Part 2(4) Polychaeta. Memoirs of the National Museum of Victoria 32, 21–42.10.24199/j.mmv.1971.32.04CrossRefGoogle Scholar
Largier, JL, Attwood, CG and Harcourt-Baldwin, JL (2000) The hydrographic character of the Knysna Estuary. Transactions of the Royal Society of South Africa 55, 107–122.10.1080/00359190009520437CrossRefGoogle Scholar
Lewis, C and Karageorgopoulos, P (2008) A new species of Marphysa (Eunicidae) from the western Cape of South Africa. Journal of the Marine Biological Association of the United Kingdom 88, 277–287.10.1017/S002531540800009XCrossRefGoogle Scholar
Luckenbach, MW (1986) Sediment stability around animal tubes : the roles of hydrodynamic processes and biotic activity. Limnology and Oceanography 31, 779–787.10.4319/lo.1986.31.4.0779CrossRefGoogle Scholar
MacNae, W (1956) Aspects of life on muddy shores in South Africa. South African Journal of Science 53, 40–44.Google Scholar
MacNae, W (1957) The ecology of plants and animals in the intertidal regions of the Zwartkops Estuary, near Port Elizabeth, South Africa. Journal of Ecology 45, 361–387.10.2307/2256924CrossRefGoogle Scholar
MacNae, W and Kalk, M (eds) (1958) A Natural History of Inhaca Island, Mozambique. Johannesburg: Witwatersrand University Press.Google Scholar
Mangum, CP, Santos, SL and Rhodes, WRJ (1968) Distribution and feeding in the onuphid polychaete, Diopatra cuprea (Bosc). Marine Biology 2, 33–34.10.1007/BF00351635CrossRefGoogle Scholar
Maree, B (2010) Structure and status of the intertidal wetlands of the Knysna Estuary. Transactions of the Royal Society of South Africa 55, 163–176.10.1080/00359190009520441CrossRefGoogle Scholar
McLachlan, A, Cockcroft, AC and Malan, DE (1984) Benthic faunal response to a high energy gradient. Marine Ecology Progress Series 16, 51–63.10.3354/meps016051CrossRefGoogle Scholar
Mead, A, Griffiths, CL, Branch, GM, Mcquaid, CD, Blamey, LK, Bolton, JJ, Anderson, RJ, Dufois, F, Rouault, M, Froneman, PW, Whitfield, AK, Harris, LR, Nel, R, Pillay, D and Adams, JB (2013) Human-mediated drivers of change – impacts on coastal ecosystems and marine biota of South Africa. African Journal of Marine Science 35, 403–425.10.2989/1814232X.2013.830147CrossRefGoogle Scholar
Napier, VR, Turpie, JK and Clark, BM (2009) Value and management of the subsistence fishery at Knysna Estuary, South Africa. African Journal of Marine Science 31, 297–310.10.2989/AJMS.2009.31.3.3.991CrossRefGoogle Scholar
Parameswaran, VS (1973) Distribution of Diopatra neapolitana Delle Chiaje (Polychaeta) in the South-West Coast of India. Indian Journal of Marine Sciences 2, 62–63.Google Scholar
Paxton, H (1993) Diopatra Audouin and Milne Edwards (Polychaeta: Onuphidae) from Australia, with a discussion of developmental patterns in the genus. The Beagle, Records of the Northern Territory Museum of Arts and Sciences 10, 115–154.Google Scholar
Paxton, H (1998) The Diopatra chiliensis confusion – redescription of D. chiliensis (Polychaeta, Onuphidae) and implicated species. Zoologica Scripta 27, 31–48.10.1111/j.1463-6409.1998.tb00427.xCrossRefGoogle Scholar
Paxton, H (2016) A new species of Diopatra (Annelida: Onuphidae) from Namibia, southwestern Africa. Marine Biodiversity 46, 889–895.10.1007/s12526-016-0451-zCrossRefGoogle Scholar
Paxton, H and Bailey-Brock, JH (1986) Diopatra dexiognatha, a new species of Onuphidae (Polychaeta) from Oahu, Hawaiian Islands. Pacific Science 40, 1–6.Google Scholar
Perez-Hurtado, A, Goss-Custard, JD and Garcia, F (1997) The diet of wintering waders in Cádiz Bay, southwest Spain. Bird Study 44, 45–52.10.1080/00063659709461037CrossRefGoogle Scholar
Philippart, CJM (1994) Interactions between Arenicola marina and Zostera noltii on a tidal flat in the Wadden Sea. Marine Ecology Progress Series 111, 251–257.10.3354/meps111251CrossRefGoogle Scholar
Pillay, D, Branch, GM and Forbes, AT (2007) The influence of bioturbation by the sandprawn Callianassa kraussi on feeding and survival of the bivalve Eumarcia paupercula and the gastropod Nassarius kraussianus. Journal of Experimental Marine Biology and Ecology 344, 1–9.10.1016/j.jembe.2006.10.044CrossRefGoogle Scholar
Pillay, D, Branch, GM, Griffiths, CL, Williams, C and Prinsloo, A (2010) Ecosystem change in a South African marine reserve (1960–2009): role of seagrass loss and anthropogenic disturbance. Marine Ecology Progress Series 415, 35–48.10.3354/meps08733CrossRefGoogle Scholar
Pillay, D, Williams, C and Whitfield, AK (2012) Indirect effects of bioturbation by the burrowing sandprawn Callichirus kraussi on a benthic foraging fish, Liza richardsonii. Marine Ecology Progress Series 453, 151–158.10.3354/meps09642CrossRefGoogle Scholar
Pires, A, Paxton, H, Quintino, V and Rodrigues, AM (2010) Diopatra (Annelida: Onuphidae) diversity in European waters with the description of Diopatra micrura, new species. Zootaxa 2395, 17–33.10.11646/zootaxa.2395.1.2CrossRefGoogle Scholar
Pires, A, Freitas, R, Quintino, V and Rodrigues, AM (2012 a) Can Diopatra neapolitana (Annelida: Onuphidae) regenerate body damage caused by bait digging or predation? Estuarine, Coastal and Shelf Science 110, 36–42.10.1016/j.ecss.2011.12.039CrossRefGoogle Scholar
Pires, A, Quintino, V, Gentil, F, Freitas, R, Rodrigues, AM (2012 b) Reproductive biology of a brooding Diopatra species: Diopatra marocensis Paxton et al., 1995. Estuarine, Coastal and Shelf Science 110, 85–92.Google Scholar
Pires, A, Figueira, E, Moreira, A, Soares, AMVM and Freitas, R (2015) The effects of water acidification, temperature and salinity on the regenerative capacity of the polychaete Diopatra neapolitana. Marine Environmental Research 106, 30–41.10.1016/j.marenvres.2015.03.002CrossRefGoogle ScholarPubMed
Reichardt, W (1988) Impact of bioturbation by Arenicola marina on microbiological parameters in intertidal sediments. Marine Ecology Progress Series 44, 149–158.10.3354/meps044149CrossRefGoogle Scholar
Robinson, TB, Alexander, ME, Simon, CA, Griffiths, CL, Peters, K, Sibanda, S, Miza, S, Groenewald, B, Majiedt, P and Sink, KJ (2016) Lost in translation? Standardising the terminology used in marine invasion biology and updating South African alien species lists. African Journal of Marine Science 38, 129–140.10.2989/1814232X.2016.1163292CrossRefGoogle Scholar
Rodrigues, AM, Pires, A, Mendo, S and Quintino, V (2009) Diopatra neapolitana and Diopatra marocensis from the Portuguese coast: morphological and genetic comparison. Estuarine, Coastal and Shelf Science 85, 609–617.10.1016/j.ecss.2009.10.004CrossRefGoogle Scholar
Siebert, T and Branch, GM (2005) Interactions between Zostera capensis, Callianassa kraussi and Upogebia africana: deductions from field surveys in Langebaan Lagoon, South Africa. African Journal of Marine Science 27, 345–356.10.2989/18142320509504094CrossRefGoogle Scholar
Siebert, T and Branch, GM (2006) Ecosystem engineers: interactions between eelgrass Zostera capensis and the sandprawn Callianassa kraussi and their indirect effects on the mudprawn Upogebia africana. Journal of Experimental Marine Biology and Ecology 338, 253–270.10.1016/j.jembe.2006.06.024CrossRefGoogle Scholar
Siebert, T and Branch, GM (2007) Influences of biological interactions on community structure within seagrass beds and sandprawn-dominated sandflats. Journal of Experimental Marine Biology and Ecology 340, 11–24.10.1016/j.jembe.2006.08.007CrossRefGoogle Scholar
Simon, CA, du Toit, AN, Smith, MKS, Claassens, L, Smith, F and Smith, P (2019) Bait collecting by subsistence and recreational fishers in Knysna Estuary may impact management and conservation. African Zoology 54, 91–103.10.1080/15627020.2019.1608862CrossRefGoogle Scholar
Simon, CA, Kara, J, Naidoo, C and Matthee, CA (2020) Genetic structure of bloodworm, Arenicola loveni (Annelida; Arenicolidae) suggests risk of local extinction in the face of overexploitation is lower than expected. African Zoology 55, 175–183. https://doi.org/10.1080/15627020.2020.1723440.CrossRefGoogle Scholar
Teske, PR, Lockyear, JF, Hecht, T and Kaiser, H (2007) Does the endangered Knysna seahorse, Hippocampus capensis, have a preference for aquatic vegetation type, cover or height? African Zoology 42, 23–30.10.1080/15627020.2007.11407373CrossRefGoogle Scholar
Thomsen, MS and McGlathery, KJ (2005) Facilitation of macroalgae by the sedimentary tube forming polychaete Diopatra cuprea. Estuarine, Coastal and Shelf Science 62, 63–73.10.1016/j.ecss.2004.08.007CrossRefGoogle Scholar
Thomsen, MS, Muth, MF and McGlathery, KJ (2011) Tube-forming polychaetes enhance invertebrate diversity and abundance in sandy sediments of Mozambique, Africa. African Journal of Marine Science 33, 327–332.10.2989/1814232X.2011.600433CrossRefGoogle Scholar
Tue, NT, Hamaoka, H, Sogabe, A, Quy, TD, Nhuan, MT and Omori, K (2012) Food sources of macro-invertebrates in an important mangrove ecosystem of Vietnam determined by dual stable isotope signatures. Journal of Sea Research 72, 14–21.10.1016/j.seares.2012.05.006CrossRefGoogle Scholar
Turpie, JK and Clark, B (2007) Development of a Conservation Plan for Temperate South African Estuaries on the Basis of Biodiversity Importance, Ecosystem Health and Economic Costs and Benefits. CAPE Regional Estuarine Management Programme (Final report).Google Scholar
van Der Westhuizen, HC and Marais, JFK (1977) Stomach content analyses of Pomadasys commersonni from the Swartkops Estuary (Pisces: Pomadasyidae). Zoologica Africana 12, 500–504.10.1080/00445096.1977.11447596CrossRefGoogle Scholar
van Herwerden, L (1989) Collection of mussel worms Pseudonereis variegata for bait – a legislative anachronism. South African Journal of Marine Science 8, 363–366.Google Scholar
van Rensburg, H (2019) Taxonomy and Distribution of Moonshine Worms (Diopatra sp.) in Knysna Estuary. Stellenbosch: Stellenbosch University.Google Scholar
Wheater, CP, Bell, JR and Cook, PA (2011) Practical Field Ecology: A Project Guide. Oxford: Wiley-Blackwell.Google Scholar
Whitfield, AK, Beckley, LE, Bennett, BA, Branch, GM, Kok, HM, Potter, IC and van der Elst, RP (1989) Composition, species richness and similarity of ichthyofaunas in eelgrass Zostera capensis beds of southern Africa. South African Journal of Marine Science 8, 251–259.10.2989/02577618909504565CrossRefGoogle Scholar
Woodin, SA (1981) Disturbance and community structure in a shallow water sand flat. Ecology 62, 1052–1066.Google Scholar
Wynberg, RP and Branch, GM (1991) An assessment of bait-collecting for Callianassa kraussi Stebbing in Langebaan Lagoon, Western Cape, and of associated avian predation. South African Journal of Marine Science 11, 141–152.10.2989/025776191784287592CrossRefGoogle Scholar
Wynberg, RP and Branch, GM (1994) Disturbance associated with bait-collection for sandprawns (Callianassa kraussi) and mudprawns (Upogebia africana): long-term effects on the biota of intertidal sandflats. Journal of Marine Research 52, 523–558.10.1357/0022240943077019CrossRefGoogle Scholar
Wynberg, RP and Branch, GM (1997) Trampling associated with bait-collection for sandprawns Callianassa kraussi Stebbing: effects on the biota of an intertidal sandflat. Environmental Conservation 24, 139–148.10.1017/S0376892997000209CrossRefGoogle Scholar

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