Abstract
Morphological variation in the three Iberian Succisella species (S. andreae-molinae, S. carvalhoana and S. microcephala) is here examined based on novel diagnostic characters, notably the shape of the outermost involucral bracts, stolon and rosette position, and primary peduncle length. Accordingly, a new identification key and synoptic table are presented in a revised taxonomic treatment, with unprecedented photographic iconography of plants in situ. Succisella carvalhoana, here lectotypified, resurfaces as a Critically Endangered narrow endemic, confined to willow swamp forests in coastal central Portugal, requiring urgent conservation attention. Plants from Spain, and east-central Portugal, previously identified as S. carvalhoana are redetermined as S. microcephala, which is, accordingly, identified as a new country record for Portugal. The three distribution centres of S. microcephala (viz. Northern Meseta, Upper Alberche basin, and the southern foothills of the Central System) should be treated as different conservation units, requiring further taxonomic investigation. Succisella andreae-molinae and S. microcephala are preliminarily downlisted to Near Threatened, due to the improved understanding of their occurrence.
References
- Amich, F., Devesa, J.A. & Bernardos, S. (2004) Taxonomic revision of the genus Succisella (Dipsacaceae) in the Iberian Peninsula. Botanical Journal of the Linnean Society 144 (3): 351–364. https://doi.org/10.1111/j.1095-8339.2003.00243.x
- Amich, F. & Bernardos, S. (2006) Succisella carvalhoana (Mariz) Baksay. In: Bañares, Á., Blanca, G., Güemes, J., Moreno, J.C. & Ortiz, S. (Eds.) Atlas y Libro Rojo de la Flora Vascular Amenazada de España, Adenda 2006. Dirección General para la Biodiversidad-Sociedad Española de Biología de la Conservación de Plantas, Madrid, pp. 82–83.
- Amich, F. & Bernardos, S. (2009) Succisella microcephala (Mariz) Baksay. In: Bañares, Á., Blanca, G., Güemes, J., Moreno, J.C. & Ortiz, S. (Eds.) Atlas y Libro Rojo de la Flora Vascular Amenazada de España, Adenda 2008. Dirección General para la Biodiversidad-Sociedad Española de Biología de la Conservación de Plantas, Madrid, pp. 118–119.
- Anthos (2024) Sistema de información de las plantas de España. Real Jardín Botánico, CSIC-Fundación Biodiversidad. Available from: httpps://www.anthos.es. (accessed 17 November 2024)
- APG IV (2016) An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV. Botanical Journal of the Linnean Society 181 (1): 1–20. https://doi.org/10.1111/boj.12385
- Araújo, P.V. (2019) Histórias da Lista Vermelha: Succisella carvalhoana. Dias com árvores. Available from: https://dias-com-arvores.blogspot.com/2019/04/historias-da-lista-vermelha-succisella.html (accessed 25 November 2024)
- Araújo, P.V. (2020) Zonas húmidas de Covões, Cantanhede. In: Porto, M. (ed.) Sítios de Interesse Botânico de Portugal Continental. Imprensa Nacional, Lisbon, pp. 214–223.
- Araújo, P.V., Covelo, F., Clemente, A., Fura, I., Pereira, P., Portela-Pereira, E., González, P., Matos, A. et al. (2024) Succisella carvalhoana (Mariz) Baksay—mapa de distribuição. Flora-On: Flora de Portugal Interactiva, Sociedade Portuguesa de Botânica. Available from: http://www.flora-on.pt/#wSuccisella+carvalhoana (accessed 17 November 2024)
- Araújo Díaz de Terán, J., López González, G., Izuzquiza, A., Sánchez Mata, D. & Martínez Labarga, J.M. (2020) Aportaciones a la flora de la comunidad de Madrid (España). Acta Botanica Malacitana 45: 181–186. https://doi.org/10.24310/abm.v45i.7183
- Bachman, S., Moat, J., Hill, A.W., De La Torre, J. & Scott, B. (2011) Supporting Red List threat assessments with GeoCAT: Geospatial conservation assessment tool. ZooKeys 150: 117–126. https://doi.org/10.3897/zookeys.150.2109
- Baksay, L. (1955) Anatomische und systematische Untersuchungen über die Gattung Succisella. Annales Historico-Naturales Musei Nationalis Hungarici 6: 167–176.
- Beck, G. (1893) Flora von Nieder-Österreich, 2. Druck und Verlag von Carl Gerold’s Sohn, Vienna, 1396 pp.
- Beentje, H. (2016) The Kew plant glossary an illustrated dictionary of plant terms, 2nd edition. The Board of Trustees of the Royal Botanic Gardens, Kew, 184 pp.
- Blanca, G. & Díaz de la Guardia, C. (1985) Avellara, género nuevo de la tribu Lactuceae (Compositae) en la Península Ibérica. Candollea 40 (2): 447–458.
- Bonsanto, D., Biscotti, N. & Wagensommer, R.P. (2023) New morphological, distribution, and ecological data on Scabiosa garganica (Caprifoliaceae), a poorly known species of the Italian flora, with evaluation of its conservation status and typification of the name. Plants 12 (9): 1915. https://doi.org/10.3390/plants12091915
- Brotero, F.A. (1804) Flora Lusitanica, 1. Typographia Regia, Lisbon, 607 pp.
- Cannon, J.F.M. (1976) Succisella G.Beck. In: Tutin, T.G., Heywood, V.H., Burges, N.A. & Valentine, D.H. (eds.) Flora Europaea, 4. p. Cambridge University Press, Cambridge, 534 pp.
- Cánovas Rubio, J.L., Sánchez Gómez, P., Jiménez, J.F., Catalán, A.E. & López Donate, J.A. (2019) Flora de interés de las Sierras de Nerpio (Albacete). Diputación de Albacete, 250 pp.
- Carapeto, A., Francisco, A., Pereira, P. & Porto, M. (2020) Lista Vermelha da Flora Vascular de Portugal Continental. Imprensa Nacional, Lisbon, 372 pp. [https://listavermelha-flora.pt/wp-content/uploads/2020/10/Lista_Vermelha_Flora_Vascular_Portugal_Continental_2020_versao_digital.pdf]
- Carlson, S.E., Mayer, V. & Donoghue, M.J. (2009) Phylogenetic relationships, taxonomy, and morphological evolution in Dipsacaceae (Dipsacales) inferred by DNA sequence data. Taxon 58 (4): 1075–1091. https://doi.org/10.1002/tax.584003
- Castroviejo, S. et al. (1986–2021) Flora iberica, vols. 1–21. Real Jardín Botánico-CSIC, Madrid.
- Coutinho, A.X.P. (1939) Flora de Portugal (plantas vasculares) disposta em chaves dicotómicas, 2nd edition. Bertrand (Irmãos), Lisbon, 938 pp.
- Devesa, J.A. (2007) Dipsacaceae. In: Devesa, J.A., Gonzalo, R. & Herrero, A. (eds.) Flora iberica, 15. Real Jardín Botánico-CSIC, Madrid, pp. 265–269.
- Devesa, J.A. & Amich, F. (2007) Succisella Beck. In: Devesa, J.A., Gonzalo, R. & Herrero, A. (eds.) Flora iberica, 15. Real Jardín Botánico-CSIC, Madrid, pp. 305–310.
- Escudero, A. & Pajarón, S. (1994) Una planta nueva de la sierra de Segura (España), Succisella andreae-molinae, sp. nov. (Dipsacaceae). Anales Jardín Botánico de Madrid 51 (2): 249–254.
- Farminhão, J. & Carapeto, A. (2024) A new combination for a neglected member of Linaria subsect. Versicolores (Plantaginaceae, Antirrhineae) endemic to the Algarve, Portugal. PhytoKeys 243: 31–45. https://doi.org/10.3897/phytokeys.243.122788
- Fernández Galiano, E. & Heywood, V.H. (1960) Catálogo de plantas de la provincia de Jaén (Mitad Oriental). Intituto Estudios Giennenses, Madrid, 205 pp.
- Fernández‐Mazuecos, M., Jiménez‐Mejías, P., Martín‐Bravo, S., Buide, M.L., Álvarez, I. & Vargas, P. (2016) Narrow endemics on coastal plains: Miocene divergence of the critically endangered genus Avellara (Compositae). Plant Biology 18 (4): 729–738. https://doi.org/10.1111/plb.12458
- Fox, J., Weisberg, S., Price, B., Adler, D., Bates, D., Baud-Bovy, G., Bolker, B., Ellison, S., Firth, D., Friendly, M., Gorjanc, G., Graves, S., Heiberger, R., Krivitsky, P., Laboissiere, R., Maechler, M., Monette, G., Murdoch, D., Nilsson, H., Ogle, D., Ripley, B., Short, T., Venables, W., Walker, S., Winsemius, D., Zeileis, A. & R-Core. (2024) car: Companion to Applied Regression, version 3.1-3. Available from: https://cran.r-project.org/web/packages/car/index.html (accessed 27 February 2025)
- Franco, J.A. (1984) Nova Flora de Portugal (Continental e Açores), 2. Escolar Editora, Lisbon, 660 pp.
- Fura, I. (2024) Um modelo para a conservação: riachos e lagoas de Febres e Covões. Unpublished MSc thesis, University of Coimbra, 162 pp.
- GBIF: The Global Biodiversity Information Facility (2024) What is GBIF? Available from: https://www.gbif.org/what-is-gbif (accessed 25 November 2024)
- García Cardo, Ó. & García Cardo, J.M. (2023) Succisella andreae-molinae Pajarón & Escudero, otro endemismo Bético descubierto en la Serranía de Cuenca. Flora Montiberica 87: 125–127.
- Giráldez, X. (1986) Contribución al conocimiento de la flora zamorana. Studia Botanica (Universidad de Salamanca) 5: 175–178.
- Gomes Machado, C.M. (1868) Catalogo methodico das plantas observadas em Portugal. Jornal de sciencias mathematicas, physicas e naturaes 1: 26–36
- Gutiérrez, D. (1908) Apuntes para la flora del Partido Judicial de Olmedo e indicaciones de los usos medicinales que algunas plantas reciben. Tip. Benito Manuel, Ávila, 136 pp.
- Haller, A. (1768) Historia Stirpium Indigenarum Helvetiae Inchoata, 2. Sumptibus Societatis typographicae, Bern, 323 pp.
- Heleno, R., Costa, J.M., Covelo, F., Santos, J., Lopes, P., Gouveia, A.C., Matos, A., Salgado, A., da Cruz, M.T.G., Farminhão, J., Horta, M., Barreto, G., Marques, A.V., Craveiro, L., Pinto, P., Santos, M., Nunes, B., Barreiro, M., Dias, A., Rodrigues, G., Esteves, L., Wanderley, M., Santos, I., Artiaga, J.P., Veríssimo, J., Vilhena, I., Moniz, L., Leão, A., Couras, M., Mendes, S.B., Nereu, M., da Silva, A.M.D., Sales, F., Gonçalves, M.T., Coutinho, A., Freitas, H., Silva, J.S., Ramos, J., Marchante, E. & Timóteo, S. (2024) O armário: Fruiting phenology data for 4,462 plant taxa in Portugal (1926–2013). Scientific Data 11 (1): 669. https://doi.org/10.1038/s41597-024-03520-9
- Henriques, J. (1876) O Jardim Botânico da Universidade de Coimbra. Imprensa da Universidade, Coimbra, 54 pp.
- Hervé, M. (2023) Testing and plotting procedures for biostatistics—Package ‘RVAideMemoire, version 0.9-83-7. Available from: https://cran.r-project.org/web/packages/RVAideMemoire/RVAideMemoire.pdf (accessed 27 February 2025)
- IUCN Standards and Petitions Committee (2024) Guidelines for Using the IUCN Red List Categories and Criteria. Version 16. Prepared by the Standards and Petitions Committee. Available from: https://www.iucnredlist.org/resources/redlistguidelines (accessed 27 February 2025)
- Jiménez‐Mejías, P., Fernández‐Mazuecos, M., Amat, M.E. & Vargas, P. (2015) Narrow endemics in European mountains: high genetic diversity within the monospecific genus Pseudomisopates (Plantaginaceae) despite isolation since the late Pleistocene. Journal of Biogeography 42 (8): 1455–1468.
- JSTOR (2000–2024) Global Plants. Available from: https://doi.org/10.2307/4108552 (accessed 25 November 2024)
- Kassambara, A. (2023) rstatix: Pipe-Friendly Framework for Basic Statistical Tests, version 0.7.2. Available from: https://cran.r-project.org/web/packages/rstatix/index.html (accessed 27 February 2025)
- Lange, J. (1861) Pugillus plantarum imprimis hispanicarum, quas in itinere 1851–52 legit. Joh. Lange II. Videnskabelige meddelelser fra den Naturhistoriske forening i Kjöbenhavn 1861: 33–116. https://doi.org/10.5962/bhl.title.10857
- López Vélez, G. (1994) Aportación a la flora de las Sierras del Sur de Albacete (Calar del Mundo y Sierras adyacentes). Acta Botanica Malacitana 19: 214–216. https://doi.org/10.24310/abm.v19i.8903
- Maguilla, E. & Martín-Díaz, A. (2022) Succisella carvalhoana. The IUCN Red List of Threatened Species 2022: e.T163379779A163385219. https://doi.org/10.2305/IUCN.UK.2022-2.RLTS.T163379779A163385219.en
- Mariz, J. (1890) Flora lusitanica exciccata: Centuriae IX et X. Boletim da Sociedade Broteriana 8: 141–158.
- Mariz, J. (1891) Especies distribuidas: 1890. Boletim da Sociedade Broteriana 9: 45–56.
- Mayer, V. (2016) Dipsacaceae (inclusive Triplostegia). In: Kadereit, J. & Bittrich, V. (eds.) Flowering Plants. Eudicots. The Families and Genera of Vascular Plants, 14. Springer, Cham, pp. 145–163. https://doi.org/10.1007/978-3-319-28534-4_11
- Mayer, V. & Ehrendorfer, F. (2013) The phylogenetic position of Pterocephalidium and the new African genus Pterothamnus within an improved classification of Dipsacaceae. Taxon 62 (1): 112–126. https://doi.org/10.1002/tax.621010
- Médail, F. & Diadema, K. (2009) Glacial refugia influence plant diversity patterns in the Mediterranean Basin. Journal of Biogeography 36 (7): 1333–1345. https://doi.org/10.1111/j.1365-2699.2008.02051.x
- Meusel, H. & Jäger, J. (1992) Vergleichende Chorologie der zentraleuropäischen Flora, 3 (2). Gustav Fischer Verlag, Jena, 266 pp.
- Moench, C. (1794) Methodus plantas horti botanici et agri Marburgensis: a staminum situ describendi. Marburgi Cattorum, in officina nova libraria academiae, Marburg, 368 pp. https://doi.org/10.5962/bhl.title.304
- Pereira, A.J., Francisco, A. & Porto, M. (2016) Flora-On: Occurrence data of the vascular flora of mainland Portugal. PhytoKeys 69: 105–119. https://doi.org/10.3897/phytokeys.69.9432
- Porto, M. (2021) Nave de Haver. In: Farminhão, J. (ed.) Sítios de Interesse Botânico de Portugal Continental, 2. Imprensa Nacional, Lisbon, pp. 140–151.
- QGIS Development Team (2024) QGIS Geographic Information System. QGIS Association. Available from: http://qgis.osgeo.org (accessed 27 February 2025)
- R Core Team (2024) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna. Available from: http://www.R-project.org/ (accessed 27 February 2025)
- Rico, E. (1984) Aportaciones y comentarios sobre la flora del centro-oeste español. Anales del Jardín Botánico de Madrid 41 (2): 407–423.
- Ríos, S., Alcaraz Ariza, F. & Robledo, A. (1995) Contribución al conocimiento de la flora ripícola del SE Ibérico, II. Anales de Biología (Universidad de Murcia) 20: 63–74. [https://revistas.um.es/analesbio/article/view/32211]
- Ríos, S., Alcaraz Ariza, F. & Valdés Franzi, A. (2003) Vegetación de sotos y riberas de la provincia de Albacete (España). Instituto de Estudios Albacetenses “Don Juan Manuel” de la Excma. Diputación de Albacete, Albacete, 365 pp.
- Salazar Mendías, C., Cano Carmona, E. & Valle Tendero, F. (1996) Aportaciones a la flora vascular de las provincias de Granada y Jaén (S. España). Acta Botanica Malacitana 21: 314–318. https://doi.org/10.24310/abm.v21i0.8693
- Sampaio, G. (1946) Flora Portuguesa, 2nd edition. Imprensa Moderna, Porto, 768 pp.
- Sánchez Gómez, P., Carrión, M.Á. & Vera Pérez, J.B. (2004) Succisella andreae-molinae Escudero & Pajarón. In: Bañares, Á., Blanca, G., Güemes, J., Moreno, J.C. & Ortiz, S. (eds.) Atlas y Libro Rojo de la Flora Vascular Amenazada de España: Táxones Prioritários. Ministerio de Medio Ambiente, Organismo Autónomo Parques Nacionales, Madrid, pp. 860–861.
- Sánchez Gómez, P., Carrión Vilches, M.Á. & Vera, J.B. (2006) Succisella andreae-molinae. The IUCN Red List of Threatened Species 2006: e.T61646A12532260. https://doi.org/10.2305/IUCN.UK.2006.RLTS.T61646A12532260.en
- Sánchez Gómez, P., Jiménez Martínez, J.F., Cánovas, J.L., Catalán, A.E. & López Donate, J.A. (2016) Flora Protegida y de Interés del Parque Natural de los Calares del Mundo y de la Sima. Consejería de Agricultura, Medio Ambiente y Desarrollo Rural de Castilla-La Mancha, Toledo, 510 pp.
- Sánchez Sánchez, J. (1980) Notas florísticas para la provincia de Salamanca (España). Anales del Jardín Botánico de Madrid 36: 265–273.
- Schneider, C.A., Rasband, W.S. & Eliceiri, K.W. (2012) NIH Image to ImageJ: 25 years of image analysis. Nature Methods 9 (7): 671–675. https://doi.org/10.1038/nmeth.2089
- Signorell, A., Aho, K., Alfons, A., Anderegg, N., Aragon, T., Arachchige, C., Arppe, A., Baddeley, A., Barton, K., Bolker, B., Borchers, H.W., Caeiro, F., Champely, S., Chessel, D., Chhay, L., Cooper, N., Cummins, C., Dewey, M., Doran, H.C., Dray, S., Dupont, C., Eddelbuettel, D., Ekstrom, C., Elff, M., Enos, J., Farebrother, R.W., Fox, J., Francois, R., Friendly, M., Galili, T., Gamer, M., Gastwirth, J.L., Gegzna, V., Gel, Y.R., Graber, S., Gross, J., Grothendieck, G., Harrell Jr, F.E., Heiberger, R., Hoehle, M., Hoffmann, C.W., Hojsgaard, S., Hothorn, T., Huerzeler, M., Hui, W.W., Hurd, P., Hyndman, R.J., Jackson, C., Kohl, M., Korpela, M., Kuhn, M., Labes, D., Leisch, F., Lemon, J., Li, D., Maechler, M., Magnusson, A., Mainwaring, B., Malter, D., Marsaglia, G., Marsaglia, J., Matei, A., Meyer, D., Miao, W., Millo, G., Min, Y., Mitchell, D., Moser, C.F., Mueller, F., Naepflin, M., Navarro, D., Nilsson, H., Nordhausen, K., Ogle, D., Ooi, H., Parsons, N., Pavoine, S., Plate, T., Prendergast, L., Rapold, R., Revelle, W., Rinker, T., Ripley, B.D., Rodriguez, C., Russell, N., Sabbe, N., Scherer, R., Seshan, V.E., Smithson, M., Snow, G., Soetaert, K., Stahel, W.A., Stephenson, A., Stevenson, M., Stubner, R., Templ, M., Temple Lang, D., Therneau, T., Tille, Y., Torgo, L., Trapletti, A., Ulrich, J., Ushey, K., VanDerWal, J., Venables, B., Verzani, J., Villacorta Iglesias, P.J., Warnes, G.R., Wellek, S., Wickham, H., Wilcox, R.R., Wolf, P., Wollschlaeger, D., Wood, J., Wu, Y., Yee, T. & Zeileis, A. (2024) DescTools: Tools for Descriptive Statistics, version 0.99.58. Available from: https://cran.r-project.org/web/packages/DescTools/index.html (accessed 27 February 2025)
- Simões de Carvalho, J.A. (1872) Memoria historica da faculdade de philosophia. Imprensa da Universidade, Coimbra, 335 pp.
- Sinkevičienė, Z. (2013) Succisella inflexa—A neglected species in meadows of Lithuania. Botanica Lithuanica 19 (1): 67–71. https://doi.org/10.2478/botlit-2013-0008
- Systematics Association Committee for Descriptive Biological Terminology (1962) II. Terminology of simple symmetrical plane shapes (chart 1). Taxon 11: 145−156. https://doi.org/10.2307/1216718
- Thiers, B. (2024) Index herbariorum: a global directory of public herbaria and associated staff. New York Botanical Garden’s Virtual Herbarium, New York. Available from: http://sweetgum.nybg.org/ih/ (accessed 25 November 2024)
- Turland, N.J., Wiersema, J.H., Barrie, F.R., Greuter, W., Hawksworth, D.L., Herendeen, P.S., Knapp, S., Kusber, W.H., Li, D.Z., Marhold, K., May, T.W., McNeill, J., Monro, A.M., Prado, J., Price, M.J. & Smith, G.F. (eds.) (2018) Chapter II Section 2. In: International Code of Nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017. Koeltz Botanical Books, Glashütten. https://doi.org/10.12705/Code.2018
- Vázquez, F.M., Ramos, S. Fernández, A. & López, J.M. (2004) Especies protegidas de Extremadura: flora. Junta de Extremadura, Mérida, 414 pp.
- Vázquez, F.M., Gutiérrez Esteban, M., Blanco, J., García Alonso, D., Guerra Barrena, M.J., Márquez-García, F., Cabeza de Vaca Molina, M.A., López Chaparro, J.L., Sánchez García, Á., Palacios González, M.J. & Mateos, J.A. (2010) Catálogo Regional de Especies Amenazadas de Extremadura. Junta de Extremadura. Consejería de Industria, Energía y Medio Ambiente, Mérida, 447 pp.
- Vázquez, F.M. & Márquez García, F. (2021) Herbario HSS Finca La Orden-Valdesequera (CICYTEX). Junta de Extremadura, La Orden Estate, Dep. Forest Production and Grasslands, Agriculture and Environment Office, Regional Government of Extremadura. Occurrence dataset: https://doi.org/10.15468/siye1z (accessed 23 November 2024)
- Verlaque, R. (1986) Étude biosystématique et phylogénétique des Dipsacaceae. IV. Tribus des Scabioseae (phylum, 1, 2, 3). Revue de Cytologie et de Biologie Végétales—Le Botaniste 9 (1): 5–72.
- Wickham, H., Francois, R., Henry, L. & Müller, K. (2023) dplyr: A Grammar of Data Manipulation, R Package Version 1.1.4. Available from: https://cran.r-project.org/web/packages/dplyr/index.html (accessed 27 February 2025)
- Willkomm, M. (1851) Sertum Florae Hispanicae. Flora 34 (47): 739–750.
- Willkomm, M. (1852) Sertum Florae Hispanicae. Commision bei Joh. Friedr. Hartknoch, Leipzig, 173 pp.