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Research article

https://doi.org/10.48027/hnb.48.061

First records of the invasive plant Carpobrotus edulis from the Bulgarian Black Sea Coast


Chavdar Gussev; Magdalena Valcheva https://orcid.org/0000-0003-4327-6501; Bogdan Prodanov https://orcid.org/0000-0002-8118-3034

Volume 48 (Issue 6): 145–158 · 1 June 2026



Abstract: First observations in Bulgaria of Carpobrotus edulis from both sand dunes and coastal cliffs are presented in the current study. In 2024, first signs of naturalisation of C. edulis, a species with high potential for invasiveness, were registered on two locations on the Southern Black Sea Coast: in embryonic shifting dunes, on a small beach west of Chernomorets Town, and on a coastal cliff in the town of Ravda. Images of the most significant morphological characteristics, that distinguish C. edulis from C. acinaciformis, are presented. In 2025, a field survey, monitoring of invasive plants, was carried out at 23 dune systems from the Southern Bulgarian Black Sea Coast and Carpobrotus species were not registered in the sand dune habitats of the studied dune complexes. Nevertheless, collected data shows, that unregulated ornamental planting is a major source of propagules for several invasive species in the dune habitats, and the appearance of Carpobrotus edulis as a garden escapee from gardens established around hotels, bungalows, beach bars, etc., is a matter of time, given that the species is widely used for ornamental purposes in the area. Therefore, monitoring of priority areas (coastal dunes and cliffs) and timely action by management authorities is needed to eradicate the recorded populations and raise awareness among the local community about the high invasiveness of the species and the existing ecological risk to coastal habitats – sand dunes and coastal cliffs.

Keywords: alien plant invasion, coastal cliffs, Hottentots-fig, psammo- and hasmophytic vegetation, sand dune habitats, western Pontic coast

References [66]

Acosta A., Carranza M.L., Di Martino L., Frattaroli A., Izzi C.F., Stanisci A. 2008 Patterns of native and alien plant species occurrence on coastal dunes in Central Italy. In: Tokarska-Guzik B., Brock J.H., Brundu G., Child L., Daehler C.C., Pišek P. (eds) Plant Invasions: Human Perception, Ecological Impacts and Management. Backhuys, Leiden, The Netherlands, 235–248.
Alados C.L., Pueyo Y., Barrantes O., Escós J., Giner L., Robles A.B. 2004 Variations in landscape patterns and vegetation cover between 1957 and 1994 in a semiarid Mediterranean ecosystem. Landscape Ecology 19: 543–559. https://doi.org/10.1023/B:LAND.0000036149.96664.9a
Andreu J., Manzano-Piedras E., Bartomeus I., Dana E.D., Vilà, M. 2010 Vegetation Response after Removal of the Invasive Carpobrotus Hybrid Complex in Andalucía, Spain. Ecological Restoration 28 (4): 440–448.
Asensi A., Díez-Garretas B., Pereña J. 2016 Alien plants of coastal dune habitats in southern Spain. Plant Biosystems 150: 477–483. https://doi.org/10.1080/11263504.2014.973463
Cabello J. 2009 Aizoaceae. In: Blanca G., Cabezudo B., Cueto M., Fernández López C., Morales Torres C. (eds) Flora Vascular de Andalucía Oriental, II vols. Sevilla, Consejería de Medio ambiente, Junta de Andalucía, 180–185.
Campos J.A., Herrera M., Biurrum I., Loidi J. 2004 The role of alien plants in the natural coastal vegetation in central-northern Spain. Biodiversity and Conservation 13: 2275–2293. https://doi.org/10.1023/B:BIOC.0000047902.27442.92
Campoy J.G., Acosta A.T.R., Affre L., Barreiro R., Brundu G., Buisson E., González L., Lema M., Novoa A., Retuerto R., Roiloa S.R., Fagundez J. 2018 Monographs of invasive plants in Europe: Carpobrotus. Botany Letters 165: 440–475. https://doi.org/10.1080/23818107.2018.1487884
Carboni M., Santoro R., Acosta A. 2010 Are some communities of the coastal dune zonation more susceptible to alien plant invasion? Journal of Plant Ecology 3: 139–147. https://doi.org/10.1093/jpe/rtp037
Carranza M.L., Carboni M., Feola S., Acosta A.T.R. 2010 Landscape-scale patterns of alien plant species on coastal dunes: the case of iceplant in central Italy. Applied Vegetation Science 13: 135–145. https://doi.org/10.1111/j.1654-109X.2009.01065.x
Conser C., Connor E.F. 2009 Assessing the residual effects of Carpobrotus edulis invasion, implications for restoration. Biological Invasions 11: 349–358. https://doi.org/10.1007/s10530-008-9252-z
Cullen J. 2011 Carpobrotus. In: Cullen J., Kness S.G., Cubey H.S. (eds) The European Garden Flora: Flowering Plants, 71, III vols. Cambridge University Press, Cambridge, 640.
Davey F.H. 1909 Flora of Cornwall. Chegwidden, Penryn.
Defeo O., McLachlan A., Schoeman D.S., Schlacher T.A., Dugan J., Jones A., Lastra M., Scapini F. 2009 Threats on sandy beach ecosystems: A review. Estuarine, Coastal and Shelf Science 81: 1–12. https://doi.org/10.1016/j.ecss.2008.09.022
Delbaere B.C.W. 1998 Facts and Figures on European Biodiversity; State and Trends 1998–1999. European Centre for Nature Conservation: Tilburg, The Netherlands, 115.
Doody J.P. 2013 Sand Dune Conservation, Management and Restoration. Springer, Dordrecht, The Netherlands, 303.
Drius M., Bongiorni L., Depellegrin D., Menegon S., Pugnetti A., Stifter S. 2019 Tackling challenges for Mediterranean sustainable coastal tourism: An ecosystem service perspective. Science of the Total Environment 652: 1302–1317. https://doi.org/10.1016/j.scitotenv.2018.10.121
Drius M., Jones L., Marzialetti F., de Francesco M.C., Stanisci A., Carranza M.L. 2019 Not just a sandy beach. The multi-service value of Mediterranean coastal dunes. Science of the Total Environment 668: 1139–1155. https://doi.org/10.1016/j.scitotenv.2019b.02.364
Gaertner M., Den Breeyen A., Hui C., Richardson D.M. 2009 Impacts of alien plant invasions on species richness in Mediterranean-type ecosystems: A meta-analysis. Progress in Physical Geography 33: 319–338. https://doi.org/10.1177/0309133309341607
Giulio S., Cao Pinna L., Carboni M., Marzialetti F., Acosta A.T.R. 2021 Invasion success on European coastal dunes. Plant Sociology 58 (1): 29–39. https://doi.org/10.3897/pls2021581/02
Gonçalves M.L. 1990 Carpobrotus N. E. Br. In: Castroviejo S., Laínz M., López González G., Montserrat P., Muñoz Garmendia F., Paiva J., Villar L. (eds) Flora Iberica, II vols. C.S.I.C, Madrid, 82–85.
Haller I., Stybel N., Schumacher S., Mossbauer M. 2011 Will beaches be enough? Future challenges on coastal tourism at the German Baltic Sea. Journal of Coastal Research 61: 70–80.
Hartmann H.E. (ed.) 2002 Illustrated handbook of succulent plants: Aizoaceae FZ (Vol. 2). Springer Science and Business Media.
Hartmann H.E.K. 1993 Aizoaceae. In: Kubitzki K., Rohwer J.G., Bittrich V. (eds) The Families and Genera of Vascular Plants. Flowering Plants – Dicotyledons. Magnoliid, Hamamelid and Caryophyllid Families. Springer-Verlag, Berlin, 37–69.
Hesp P.A., Martínez M.L. 2007 Disturbance processes and dynamics in coastal dunes. In: Johnson E.A., Miyanishi K. (eds) Plant Disturbance Ecology: The Process and the Response. Elsevier/Academic Press, Cambridge, MA, USA, 215–247.
Innangi M., Marzialetti F., Di Febbraro M., Acosta A.T.R., De Simone W., Frate L., Finizio M., Villalobos Perna P., Carranza M.L. 2023 Coastal Dune Invaders: Integrative Mapping of Carpobrotus sp. pl. (Aizoaceae) Using UAVs. Remote Sensing 15 (2): 503. https://doi.org/10.3390/rs15020503
Interpretation Manual of European Union Habitats – EUR28 2013 Interpretation Manual of European Union Habitats – EUR28. European Commission, DG Environment, Nature ENV B.3.
Janssen J., Rodwell J., García Criado M., Gubbay S., Haynes T., Nieto A., Sanders N., Landucci F., Loidi J., Ssymank A., Tahvanainen T., Valderrabano M., Acosta A.T.R., Aronsson M., Arts G., Attorre F., Bergmeier E., Bijlsma R.-J., Bioret F., Biţă-Nicolae C., Biurrun I., Calix M., Capelo J., Čarni A., Chytrý M., Dengler J., Dimopoulos P., Essl F., Gardfjell H., Gigante D, Giusso del Galdo G., Hájek M., Jansen F., Jansen J., Kapfer J., Mickolajczak A., Molina J.A., Molnár Z., Paternoster D., Piernik A., Poulin B., Renaux B., Schaminée J., Šumberová K., Toivonen H., Tonteri T., Tsiripidis I., Tzonev R., Valachovič M. 2016 European Red List of Habitats – Part 2 Terrestrial and freshwater habitats. European Union. https://doi.org/10.2779/091372
Jucker T., Carboni M., Acosta A.T.R. 2013 Going beyond taxonomic diversity: deconstructing biodiversity patterns reveals the true cost of iceplant invasion. Diversity and Distributions 19: 1566–1577. https://doi.org/10.1111/ddi.12124
Kavrakova V., Dimova D., Dimitrov M., Tsonev R., Belev T., Rakovska K. 2009 Rakovodstvo za opredelyane na mestoobitania ot evropeyska znachimost v Bulgaria. World Wide Fund for Nature, Danube-Carpathian Program and federation “Zeleni Balkani”, Sofia, 131. (In Bulgarian)
Lousley J.E. 1971 Flora of the Isles of Scilly. David and Charles, Newton Abbot.
Mack R., Simberloff D., Lonsdale W.M., Evans H., Clout M., Bazzaz F.A. 2000 Biotic invasions: Causes, epidemiology, global consequences and control. Ecological Applications 10: 689–710.
Malavasi M., Santoro R., Cutini M., Acosta A.T.R, Carranza L. 2014 The impact of human pressure on landscape patterns and plant species richness in Mediterranean coastal dunes. Plant Biosystems 150 (1): 73–82. https://doi.org/10.1080/11263504.2014.913730
Malavasi M., Santoro R., Cutini M., Acosta A.T.R., Carranza M.L. 2013 What has happened to coastal dunes in the last half century? A multitemporal coastal landscape analysis in Central Italy. Landscape and Urban Planning 119: 54–63. https://doi.org/10.1016/j.landurbplan.2013.06.012
Manning J. 2018 Field guide to fynbos. Penguin Random House, South Africa.
Marloth R. 1913 The Flora of South Africa, I vols. Darter Bros, Capetown.
McClintock D. 1975 The Wild Flowers of Guernsey: With Notes of the Frequencies of All Species Recorded for the Channel Islands. Collins, London.
Mifsud S. 2021 Morphology of the invasive Carpobrotus (Aizoaceae) in Europe: Malta as a case study. Mediterranean Botany 42: e71195. https://doi.org/10.5209/mbot.71195
Mucina L., Adams J., Knevel I., Rutherford M., Powrie L., Bolton J., van der Merwe J., Anderson R., Bornman T., le Roux A., Janssen J. 2006 Coastal Vegetation of South Africa. Strelitzia 19: 658–583.
Novoa A., González L. 2014 Impacts of Carpobrotus edulis (L.) NE Br. on the germination, establishment and survival of native plants: a clue for assessing its competitive strength. PLoS One 9 (9): e107557. https://doi.org/10.1371/journal.pone.0107557
Novoa A., González L., Moravcová L., Pyšek P. 2013 Constraints to Native Plant Species Establishment in Coastal Dune Communities Invaded by Carpobrotus edulis: Implications for Restoration. Biological Conservation 164: 1–9. https://doi.org/10.1016/j.biocon.2013.04.008
Novoa A., Rodríguez R., Richardson D., González L. 2014 Soil quality: a key factor in understanding plant invasion? The case of Carpobrotus edulis (L.) NE. Br. Biological Invasions 16: 429–443. https://doi.org/10.1007/s10530-013-0531-y
Ortiz D.G., Lumbreras E.L., Rosselló J.A. 2008 Flora alóctona suculenta valenciana: Aizoaceae y Portulacaceae. Monografías De Bouteloa 7: 1–68.
Petrosillo I., Zurlini G., Corlianò M.E., Zaccarelli N., Dadamo M. 2007 Tourist perception of recreational environment and management in a marine protected area. Landscape and Urban Planning 79: 29–37. https://doi.org/10.1016/j.landurbplan.2006.02.017
Preston C.D., Sell P.D. 1988 The Aizoaceae Naturalized in the British Isles. Watsonia 17 (3): 217–245.
Prodanov B., Gussev C., Sopotlieva D., Valcheva M., Bekova R., Baltakova A., Tzonev R., Popov J. 2025 A standard procedure for dune mapping along the Bulgarian Black Sea coast: an integrated approach combining UAS photogrammetry, geomorphological and phytocoenological surveys. Frontiers in Mareen Science 12:1579724. https://doi.org/10.3389/fmars.2025.1579724
Roiloa S.R. 2019 Clonal traits and plant invasiveness: the case of Carpobrotus NE Br.(Aizoaceae). Perspectives in Plant Ecology, Evolution and Systematics 40: 125479. https://doi.org/10.1016/j.ppees.2019.125479
Roiloa S.R., Rodríguez-Echeverría S., de la Pena E., Freitas H. 2010 Physiological integration increases the survival and growth of the clonal invader Carpobrotus edulis. Biological Invasions 12 (6): 1815-1823. https://doi.org/10.1007/s10530-009-9592-3
Roiloa S.R., Abalde S., Xu C.Y., López L. 2017 The effect of stolon fragmentation on the colonization of clonal invasive Carpobrotus edulis in a coastal dune system: a field test. Plant Species Biology 32: 460–465. https://doi.org/10.1111/1442-1984.12157
Santoro R., Jucker T., Carboni M., Acosta A.T.R. 2012 Patterns of plant community assembly in invaded and non-invaded communities along a natural environmental gradient. Journal of Vegetation Science 23: 483–494. https://doi.org/10.1111/j.1654-1103.2011.01372.x
Sarmati S., Conti L., Acosta A.T. 2019 Carpobrotus acinaciformis vs Carpobrotus edulis: Are there any differences in their impact on coastal dune plant biodiversity? Flora 257: 151422. https://doi.org/10.1016/j.flora.2019.151422
Schlacher T., Dugan J., Schoeman D.S., Lastra M., Jones A., Scapini F., McLachlan A., Defeo O. 2007 Sandy beaches at the brink. Diversity and Distributions 13: 556–560. https://doi.org/10.1111/j.1472-4642.2007.00363.x
Šilc U., Stešević D., Luković M., Caković D. 2020 Changes of a sand dune system and vegetation between 1950 and 2015 on Velika plaža (Montenegro, E Mediterranean). Regional Studies in Marine Science 35: 101139. https://doi.org/10.1016/j.rsma.2020.101139
Simberloff D., Martin J.L., Genovesi P., Maris V., Wardle D.A., Aronson J., Courchamp F., Galil B., García-Berthou E., Pascal M., Pyšek P., Sousa R., Tabacchi E., Vilà M. 2013 Impacts of biological invasions: what’s what and the way forward. Trends in Ecology and Evolution 28: 58–66. https://doi.org/10.1016/j.tree.2012.07.013
Strid A., Tan K. 1997 Carpobrotus. In: Strid A., Tan K. (eds) Flora Hellenica I. Koeltz Scientific Books, Königsten.
Suehs C., Affre L., Médail F. 2004 Invasion dynamics of two alien Carpobrotus (Aizoaceae) taxa on a Mediterranean island: I. Genetic diversity and introgression. Heredity 92: 31–40. https://doi.org/10.1038/sj.hdy.6800374
Suehs C.M., Affre L., Médail F. 2004 Invasion dynamics of two alien Carpobrotus (Aizoaceae) taxa on a Mediterranean island: II. Reproductive strategies. Heredity 92 (6): 550–556. https://doi.org/10.1038/sj.hdy.6800454
Traveset A., Moragues E., Valladares F. 2008 Spreading of the invasive Carpobrotus aff. acinaciformis in Mediterranean ecosystems: the advantage of performing in different light environments. Applied Vegetation Science 11: 45–55. https://doi.org/10.3170/2007-7-18303
Tzonev R. 2015 Black Sea embryonic dunes. Red Data Book of the Republic of Bulgaria. Natural Habitats. BAS and MoEW, Sofia, Bulgaria.
Tzonev R. 2015 Vegetation on the Black Sea sand beaches. Red Data Book of the Republic of Bulgaria. Natural Habitats. BAS and MoEW, Sofia, Bulgaria.
Valcheva M., Sopotlieva D., Apostolova I., Tsvetkova N. 2021 Vegetation Characteristics and Recent Successional Trends of Sand Dune Habitats at the Bulgarian Black Sea Coast. Coasts 1: 1–24. https://doi.org/10.3390/coasts1010001
Van Dijk H.W.J. 1989 Ecological Impact of Drinking-water Production in Dutch Coastal Dunes. In: van der Meulen F., Jungerius P.D., Visser J., (eds) Perspectives in Coastal Dune Management: Proceedings of the European Symposium Leiden. SPB Academic Publishing, Hague, The Netherlands, 163–182.
Varone L., Catoni R., Bonito A., Gini E., Gratani L. 2017 Photochemical performance of Carpobrotus edulis in response to various substrate salt concentrations. South African Journal of Botany 111: 258–266. https://doi.org/10.1016/j.sajb.2017.03.035
Velev S. 2002 Climatic regioning. In: Kopralev I. (ed.) Geography of Bulgaria. Publishing House ForCom, Sofia, Bulgaria, 155–156.
Vilà M., Tessier M., Suehs C.M., Brundu G., Carta L., Galanidis A., Lambdon P., Manca M., Médail F., Moragues E., Traveset A., Troumbis A.Y., Hulme P.E. 2006 Local and regional assessments of the impacts of plant invaders on vegetation structure and soil properties of Mediterranean islands. Journal of Biogeography 33: 853–861. https://doi.org/10.1111/j.1365-2699.2005.01430.x
Weber E., D’Antonio C.M. 1999 Germination and growth responses of hybridizing Carpobrotus species (Aizoaceae) from coastal California to soil salinity. American Journal of Botany 86 (9): 1257–1263.
Wisura W., Glen H.F. 1993 The South African species of Carpobrotus (Mesembryanthema-Aizoaceae). Contributions to the Bolus Herbarium, vol. 15: 76–107.

Additional information

Received: 6 April 2026 · Accepted: 8 May 2026

Editor: Stanislav Abadjiev

Conflicts of interest:
No potential conflict of interest has been reported by the authors, reviewers, or subject editor.

Authors’ contributions (using CrediT ↗):
Chavdar Gussev: conceptualisation, data curation, formal analysis, investigation, methodology, validation, writing—original draft, writing—review and editing; Magdalena Valcheva: conceptualisation, data curation, formal analysis, investigation, methodology, visualisation, writing—original draft, writing—review and editing; Bogdan Prodanov: data curation, investigation, validation, visualisation, writing—review and editing.

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Gussev Ch., Valcheva M., Prodanov B. 2026 First records of the invasive plant Carpobrotus edulis from the Bulgarian Black Sea Coast. Historia naturalis bulgarica 48: 145–158.

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