(Campanulaceae) in the Western Mediterranean Region
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2274 campanula:Maquetación 1 13/06/2011 12:09 Página 15 Anales del Jardín Botánico de Madrid Vol. 68(1): 15-47 enero-junio 2011 ISSN: 0211-1322 doi: 10.3989/ajbm.2274 A taxonomic revision of the Campanula lusitanica complex (Campanulaceae) in the Western Mediterranean region by Jara Cano-Maqueda & Salvador Talavera Departamento de Biología Vegetal y Ecología, Universidad de Sevilla, E-41080 Sevilla, Spain. Corresponding author: [email protected] Abstract Resumen Cano-Maqueda, J. & Talavera, S. 2011. A taxonomic revision of Cano-Maqueda, J. & Talavera, S. 2011. Una revisión taxonómi- the Campanula lusitanica complex (Campanulaceae) in the ca del complejo Campanula lusitanica (Campanulaceae) en la re- Western Mediterranean region. Anales Jard. Bot. Madrid 68(1): gión occidental mediterránea. Anales Jard. Bot. Madrid 68(1): 15-47. 15-47 (en inglés). The systematics of the annual species of the Campanula lusita- En este trabajo se revisa la sistemática de las especies anuales del nica complex in the Western Mediterranean is reviewed using complejo Campanula lusitanica en el occidente del Mediterrá- ITS sequences, karyology and morphology of all species of the neo usando secuencias ITS, cariología y análisis de los caracteres complex. The information provided by these three sources is morfológicos de todas las especies del complejo. La información consistent, and the species are reorganized into two groups with proporcionada por estas tres fuentes de caracteres es congruen- the rank of section: Campanula sect. Rapunculus Boiss. and te, y las distintas especies se reorganizan en dos grupos princi- Campanula sect. Decumbentes, the latter described as new in pales con categoría de sección, que se corresponden con los dos this work. These sections comprise two well-defined subclades subclados bien definidos: Campanula sect. Rapunculus Boiss. y in the phylogenetic analyses. Sect. Rapunculus is composed, in Campanula sect. Decumbentes, descrita como nueva en este the W Mediterranean region, by C. lusitanica L. and C. matriten- trabajo. En el Mediterráneo occidental, la sect. Rapunculus está sis A. DC., both with 2n = 18 chromosomes, and C. cabezudoi formada por las siguientes especies anuales: C. lusitanica L. y Cano-Maqueda & Talavera, C. specularioides Coss., C. transta- C. matritensis A. DC., con 2n = 18 cromosomas, y C. cabezudoi gana R. Fern., and C. broussonetiana Schult., all with 2n =20 Cano-Maqueda & Talavera, C. specularioides Coss., C. transta- chromosomes. In the Iberian Peninsula, Sect. Decumbentes gana R. Fern. y C. broussonetiana Schult., con 2n = 20 cromoso- com prises two endemic species, C. decumbens A. DC. with mas. En la Península Ibérica la sect. Decumbentes está represen- 2n = 32 chromosomes and C. dieckii Lange with 2n = 28 chro- tada por C. decumbens A. DC., con 2n = 32 cromosomas, y mosomes. In C. decumbens a new subspecies is described: C. dieckii Lange, con 2n = 28 cromosomas, ambas endémicas de C. decumbens subsp. baetica Cano-Maqueda & Talavera, which la Península Ibérica; de C. decumbens se describe una subespecie occurs in the Guadalquivir valley. In the formal systematic part nueva: C. decumbens subsp. baetica Cano-Maqueda & Talavera, we provide a key to identify these annual species of the Western taxon del valle del Guadalquivir muy bien diferenciado morfoló- Mediterranean, with a description and typification, photographs gicamente de la subsp. decumbens. En la parte sistemática se of flowers and fruits, distribution maps, and comments on the proporciona una clave para la identificación de estas especies habitat for each taxon. anuales del oeste del Mediterráneo, así como una descripción y tipificación, fotografías de las flores y frutos, mapas de distribu- ción, y aspectos de la ecología para cada uno de los táxones. Keywords: phylogeny, rn DNA ITS, karyology, systematic, typi- Palabras clave: filogenia, secuencias ITS, cariología, sistemá- fication. tica, tipificación. Introduction there are also some annual herbs, the latter mainly in The genus Campanula comprises about 420 species the Mediterranean. This genus has a high morpholog- distributed mainly in temperate regions of the North- ical complexity that is reflected in the different classi- ern Hemisphere (Lammers, 2007a, b). They are usu- fications that have been proposed. ally perennial herbs, although some are shrubby, and Based mainly on the morphology of the calyx, A. de 2274 campanula:Maquetación 1 13/06/2011 12:09 Página 16 16 J. Cano-Maqueda & S. Talavera Candolle (1830, 1838) divided the genus Campanula C. peregrina L. (E Mediterranean, including Cyprus, into two sections: sect. Medium A. DC., with calycine Lebanon and SW Turkey and S Anatolia); they are appendages, and sect. Eucodon A. DC., without them. hispid perennial plants with large infundibuliform This classification was followed by most nineteenth flowers, arranged in spikes or panicles, with capsules century authors (e.g. Endlicher, 1838; Willkomm, opening by three pores of middle position, and with 1868; Bentham & Hooker, 1876; Rouy, 1908). In con- winged seeds. trast, Boissier (1875) divided the genus into two sec- Sect. Alaria contains a single species, C. pterocaula tions based on fruit dehiscence: sect. Medium A. DC., Hausskn., endemic to N and C Anatolia (Turkey); it is with capsules opening by basal pores or valves, and a biennial glabrous plant, with winged stems, large sect. Rapunculus Boiss., with apical or middle position flowers arranged in spiciform inflorescences, and cap- pores or valves. This classification was followed with sules opening by apical pores. some modifications by Fedorov (1957, 1976). Sect. Rapunculus contains around 50 species dis- Perhaps the taxonomic treatment with most insight tributed mainly in the Mediterranean region; it com- was that by Damboldt (1976) during the preparation prises perennial, biennial or annual plants, glabrous of the Flora of Turkey. He divided the genus Campa - or tomentose, with wingless stems, the inflorescence a nula into six subgenera: Campanula, Megalocalyx panicle, or sometimes spiciform, the capsules opening Damboldt, Sicyocodon (Feer) Damboldt, Roucela (Du- by apical or middle position pores, and wingless mort.) Damboldt, Brachycodonia (Fedorov) Damboldt seeds. and Rapunculus (Boiss.) Kharadze. The subgenus Recent molecular phylogenies (Eddie & al., 2003; Campanula includes plants with 3 or 5 stigmas, with or Park & al., 2006; Roquet & al., 2008, 2009; Cellinese & without calycine appendages, and with the capsules al., 2009; Borsch & al., 2009; Haberle & al., 2009; Ste- opening by basal or middle position pores, or indehis- fanovic ´ & Lakušic, ´ 2009) have revealed that the genus cent. This is the most complex subgenus, with 12-15 Campanula is paraphyletic. Studies by Roquet & al. recognized sections and it contains most of the species (2008) using ITS sequences, show that Campa nula of the genus. According to Sáez & Aldasoro (2003), sect. Pterophyllum together with the genera Musschia subgenera Megalocalyx and Sicyocodon are annual Dumort. and Gadellia Schulkina form a clade (Muss- herbs with flowers with calycine appendages and with chia clade) that is sister to the Campanula core. capsules opening by basal valves; the two subgenera The Campanula core consists of two subclades: one differ in the size of style: very long (exceeding 35 mm) formed by the subgenera Campanula, Megalocalyx in the subgen. Sicyocodon, and short (less than 15 mm) and Roucela, together with other genera of Campanu- in the subgen. Megalocalyx. The around 15 species of laceae; and a second subclade comprising subgenus these two subgenera are distributed in the Mediter- Rapunculus (sections Rapunculus and Alaria) and sub- ranean Basin, SW Asia and Macaronesia. The sub- genus Brachycodonia, again with some other genera of genus Roucela comprises annual plants, with flowers Campanulaceae. without calycine appendages and capsules opening The annual species of sect. Rapunculus sensu by basal valves; it consists of 5 species distributed Damboldt have two centers of diversity: the Iberian throughtout the E Mediterranean and SW Asia, with Peninsula and W Morocco, and the E Mediterranean the exception of C. erinus L. which has a wider distrib- (Greece and Turkey). The annual species of the Iber- ution. The subgenus Brachycodonia comprises only ian Peninsula (which include many species names, C. fastigiata Dufour, an annual plant with axillary in- viz. C. lusitanica L., C. broussonetiana Schult., C. trans - conspicuous flowers, without calycine appendages, tagana R. Fern., C. matritensis A. DC., C. decumbens and with capsules opening by three apical valves; A. DC., C. specularioides Coss. C. cabezudoi Cano- C. fastigiata occurs on gypsum soils in Spain, N Africa, Maqueda & Talavera and C. diekii Lange) have re- C Asia and Transcaucasia. ceived diverse treatments in recent taxonomic studies, The subgenus Rapunculus includes annual and i.e as varieties of Campanula patula L. (Pau, 1921; perennial plants, without calycine appendages, with a Cuatrecasas, 1929), as subspecies or varieties of C. lusi- large campanulate or infundibuliform corolla, and tanica L. (Pau, 1924; Sáez & Aldasoro, 2001), or as va- with capsules opening by apical or middle position rieties of C. decumbens A. DC. (López-González, pores. Damboldt (1976, 1978) divided this subgenus 1979-1980). In all cases most taxa have been relegated in three sections: Pterophyllum Damboldt, Alaria to infraspecific status. Damboldt and Rapunculus. Sect. Pterophyllum in- However, our detailed morpho-geographical