Christoph Oberprieler The Systematics of Anthemis L. (Compositae, Anthemideae) in W and C North Africa Abstract Oberprieler, Ch.: The Systematics of Anthemis L. (Compositae, Anthemideae) in W and C North Africa. - Bocconea 9: 1-328.1998. - ISSN 1120-4060. Anthemis L., a mainly Mediterranean and SW Asian genus of the Compositae-An­ themideae, is revised far the W and C North African part (Marocco, Algeria, Tunisia, Libya) of its distribution range. On the basis of detailed morphological studies ali species are fully de­ scribed and illustrations of habit, leaves, involucral bracts, and pales are presented, along with photographs of mature achenes under SEM and of transverse sections of achenes under LM. Keys far the deterrnination of ali species, subspecies, and varieties are provided. Complete synonymies are given, inc1uding the typification of the names as far as possible. Oistribution maps of ali taxa studied are presented. Chromosome numbers and karyotypes far most taxa are provided and discussed. In the area covered, Anthemis is represented by 25 species belonging to two subgenera, four sections, and four series. Three taxa are described as new to science: A. maritima subsp. pseudopunctata Oberprieler, A. stiparum subsp. intermedia Oberprieler, and A. zaianiea Oberprieler. The following ten new combinations are established: A. abylaea (Font Quer & Maire) Oberprieler, A. maroeeana subsp. aguilarii (Maire & Sennen) Oberprieler, A. mauritiana subsp. faurei (Maire) Oberprieler, A. peduneulata subsp. atlantica (Pomel) Oberprieler, A. pedunculata subsp. clausonis (Pomel) Oberprieler, A. peduneulata var. discoi­ dea (Boiss.) Oberprieler, A. peduneulata subsp. turolensis (Pau ex Caballero) Oberprieler, A. punctata subsp. kabylica (Battand.) Oberprieler, A. stiparum subsp. sabulieola (Pomel) Oberprieler, A. tenuiseeta subsp. jahandiezii (Maire) Oberprieler. Multivariate statistical methods (principal component analysis) and/or analyses of random amplified polymorphic ONAs (RAPO) were used to assess morphological and genetic varia­ tion and to address problems of species delimitation in the Anthemis boveana group and the A. pedunculata - A. punetata complex. Contents 1. Introduction .......... ......... .. ......... .. .. .. .. .. .. ........... ........................... ......... ...... .. ....... .. ...... 6 2. Material and methods ........................... .. ....... .. ... ................................. .. ... .... ...... ........ 7 3. Taxonomic history ............. .. ...... .. ........... .. .... ... ...... .. ........... .. .................... .. .. ..... .. ..... 9 4. Generic relationships .... ..... .. ........ .. ...... .. ....... .. .. .. ............. ....... .. ..... ....... .. .. ........... .... lO 5. Supraspecific taxonomy ............ .. ..... :................................. .. .. ... .. ..... ....... .. .. .. ........... 12 6. Phytogeography .... .. ... .................................... .. ..... ....... .. ....................... .. .. ............. .. 14 7. Modes of speciation within Anthemis in N Africa ................ .............. ..................... 16 8. Delimitation oftaxa - Concepts of species, subspecies, and varieties .... .. .... ...... .. .... 18 9. Morphological features ............................................................................................ 20 lO. Chromosomes ....... .... ... .... ........................... .... ......... .. ...................... ........... ..... ... .. ... 35 Il. Pollen morphology ...... ......... .. ....... .. .......... .... .. .. ....... .......... ..... ............... .. .. ............ .. 66 6 Oberprieler: Anthemis in N Africa 12. Species delimitation in the Anthemis boveana group .......................................... ... 76 13 . Species delimitation in the Anthemis pedunculata - Anthemis punctata complex ... 85 14. Systematic descriptions and keys .. .. ... ....................... ..... ........... ....................... .... .. 107 Key to the N African species .... .. ................ ... ........ .... .... .. ....... ..... .. .. ............ .. 109 Clé des espèces de l'Afrique du Nord ..... ..... .. .... ... .......... .. ..... ... ... .............. .. 111 . Anthemis subg. Anthemis ............... .......... .. .................... .................. .. ............ 114 Anthemis sect. Anthemis ..................................... .. .................. .. .... .. 114 Anthemis sect. Hiorthia .. ................................. .. ....................... .. ..... 227 Anthemis sect. Maruta ..... ................................. .. ........ ............... .... .. 297 Anthemis subg. Cota .. .. ............ .. ... ................. ... .. ................ ... ........................ 307 Anthemis sect. Cota ... ...................... .............. .. ......... ......... ...... ........ 307 15. Acknowledgements .... .. .. ....... ... ........................... ................... ........................... ..... 313 16. References ..... ...... .................... .. .............................. .............. ............. .. ... ........... ... 314 17. Index to scientific names .. .. ....... .. ...... .. .. .. .................... ........................... .. .. .. ........ 324 1. Introduction The genus Anthemis belongs to the family Compositae Giseke, alternatively named As­ teraceae Dumort. It is the name-giving genus of the Anthemideae Cass., a tribe which presently consists of 109 genera (Bremer & Humphries 1993; adding Castrilanthemum recently described by Vogt & Oberprieler 1996) and shows an extratropical, mainly Old­ World-centred distribution. According to the most recent phylogenetic reconstruction of the family (Bremer 1996), the Anthemideae are a member of the subfamily Asteroideae and form the sister group to the Astereae. While Heywood & Humphries (1977) indicate around 130 species in the genus, more recent estimates give a total of c. 210 annual, biennial, or perennial species (Bremer & Humphries 1993). Since Candolle's account of Anthemis in his Prodromus (Candolle 1838), the genus has never again been subject to a complete revisionary treatment. However, quite comprehen­ sive regional treatments like Boissier's (1875) Flora orientalis account and Eig's (1938) studies on the orientai species of Anthemis, along with revisions made in the course of the large flora projects of the last 40 years, have resulted in our up-to-date understanding of the genus over a large part of its distributional range, viz. Europe (Fernandes 1975a, 1975b, 1976, 1983), E Europe and W Asia (Fedorov 1961), Turkey (Grierson & Yavin 1975), the Near East (Feinbrun-Dothan 1978), the Flora iranica area (Iranshahr 1986), and Saudi Arabia (Ghafoor & AI-Turki 1997). Additional information was contributed by revisionary treatments of defined infrageneric groups (A. ammanthus group, Greuter 1968; A. sect. Maruta, Yavin 1970; A. sect. Anthemis, Yavin 1972; A. tomentosa group, Geor­ giou 1990) and by geographically limited biosystematic revisions (Bulgaria: Kuzmanov & al. 1981 and Thin 1983; Spain: Benedf i Gonzalez 1987). In N Africa, as compared to other regions within the distributional range of Anthemis, the taxonomy of the genus is poorly understood, and a criticaI revision is lacking. While there are flora treatments available for Algeria (Quézel & Santa 1963), Tunisia (Pottier­ Alapetite 1981), Libya (Alavi 1986), and Egypt (Tackholm 1974), for Morocco there is only a checklist (Jahandiez & Maire 1934). The main aim of the present study was to fill Bocconea 9 - 1998 7 this gap by elaborating a complete taxonomic treatment of the genus for the area covered by Maire's Flore de l'Afrique du Nord (Maire & al. 1952-1987), i.e. the territories of Morocco, Algeria, Tunisia, and Libya. 2. Material and methods This revision is based partly on plant material collected during field trips to Morocco and Tunisia in 1992-1995, which is deposited at the herbarium of the Botanic Garden & Botanical Museum Berlin-Dahlem (B). In June 1992, the study and collection of Anthemis in N Morocco was enabled by the participation in the "Iter Mediterraneum V" sponsored by the Organization for the Phyto-Taxonomic Investigation of the Mediterranean Area (OPTIMA). Further collections in Morocco (April-May 1993, May 1995) and Tunisia (May 1994) were made during field trips organised by the Botanic Garden and Botanical Museum Berlin-Dahlem. Additionally, the study is based on herbarium material from the following public her­ baria, abbreviated as in Holmgren & al. (1990) and Holmgren & Holmgren (1996), and private collections: B Botanischer Garten & Botanisches Museum Berlin-Dahlem BC Institut Botanic de Barcelona BC-Sennen Institut Botanic de Barcelona (Herbarium F. Sennen) BM-Clifford The Natural History Museum, London (Herbarium G. Clifford; microfiche) C-Vahl Botanical Museum and Library, University ofCopenhagen (Herbarium M. Vahl; microfiche) FI Museo di Storia Naturale, Museo Botanico, Firenze G Conservatoire et Jardin botaniques de la ville de Genève GOET Systematisch-Geobotanisches Institut der Universitlit Gèittingen K Royal Botanic Gardens, Kew (London) LINN The Linnean Society of London (Herbarium C. Linné; micro­ fiche) MA Rea! Jardfn Botanico, Madrid MPU-AtN Institut de Botanique Montpellier (Herbier de l' Afrique du Nord) MPU-Braun-Blanquet Institut de Botanique Montpellier (Herbarium J. Braun­ Blanquet) MPU-Dubuis Institut de Botanique Montpellier (Herbarium A. Dubuis) MPU-Weiller Institut de Botanique Montpellier (Herbarium M. Weiller) MPU-Sauvage