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A Taxonomic Survey of Lotus Section Pedrosia (Leguminosae, Loteae) Graeme Sandral, Margarita V

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A Taxonomic Survey of Lotus Section Pedrosia (Leguminosae, Loteae) Graeme Sandral, Margarita V © Landesmuseum für Kärnten; download www.landesmuseum.ktn.gv.at/wulfenia; www.biologiezentrum.at Wulfenia 13 (2006): 97–192 Mitteilungen des Kärntner Botanikzentrums Klagenfurt A taxonomic survey of Lotus section Pedrosia (Leguminosae, Loteae) Graeme Sandral, Margarita V. Remizowa & Dmitry D. Sokoloff Summary: A taxonomic review of the section Pedrosia (excl. sect. Rhyncholotus) is presented. Members of the section occur in the Macaronesian Islands, north-western part of Africa and in the Mediterranean region. Centers of diversity are Canary Islands, Cape Verde, Madeira and Morocco. Twenty nine species of the section are recognized; their diagnostic characters are discussed and extensively illustrated. An identifi cation key to species of the section is given as well as morphological descriptions of species and subspecies. Leaf morphology was found to be a particularly useful character in species discrimination. Many species are quite variable, and their identifi cation is only possible using a suite of several characters. Furthermore, various morphological characters have different taxonomic weight in different regions (e.g. Cape Verde and Morocco). The following taxonomic decisions are adopted: (1) Lotus azoricus is synonymous with L . argyrodes; (2) L . tenellus is accepted as a species distinct from L . glaucus, with an area covering Tenerife and Gran Canaria; (3) all material examined from Hierro (Canary Islands) is identifi ed asL . sessilifolius; (4) L . hillebrandii is treated as a subspecies of L . campylocladus; (5) L . spartioides is synonymous with L . holosericus. Representatives of the section Pedrosia from Lanzarote and Fuerteventura are closer to those from Madeira than to plants from other Canary Islands. Keywords: Fabaceae, Leguminosae, Loteae, Lotus, Pedrosia, taxonomy, determination key, typifi cation, comb. nov. The genus Lotus includes agriculturally important plants such as L . corniculatus, and L . pedunculatus, as well as the model legume L . japonicus and ornamentals (e.g. L . berthelotti and L . maculatus). In recent times increased interest in the placement of Lotus and its species using molecular and morphological data has stimulated a closer examination of these genetic resources. Such studies need a sound taxonomic basis, however, Lotus is a taxonomically diffi cult genus (GILLETT 1958; HEYN & HERRNSTADT 1968; KRAMINA 1999a,b, 2006; KRAMINA & SOKOLOFF 2004). In particular, no detailed worldwide taxonomic study of the whole genus has been produced since BRAND’s (1898) monograph. However, in more recent times taxonomic and phylogentic studies of Lotus have been undertaken, for example, the phylogeny of Lotus has been studied by means of cladistic analyses based on morphological (ARAMBARRI 2000) and molecular data (ALLAN et al. 2003, 2004; DEGTJAREVA et al. 2006). In addition, KIRKBRIDE (1999) published a worldwide list of Lotus species, while KRAMINA (1999b, 2006) published taxonomic studies of the L . corniculatus and L . angustissimus groups. Kramina (pers. comm.) is also preparing a monographic revision of the whole section Lotus which includes many agriculturally important species. Australian and Pacifi c Lotus species have been revised recently by KRAMINA & SOKOLOFF (2004). The present paper contributes to this ongoing taxonomic revision of the genus Lotus through the taxonomic survey of its section Pedrosia. The section Pedrosia, which is sometimes accepted as a subgenus (MONOD 1980; MADER & PODLECH 1989; KRAMINA & SOKOLOFF 1999), includes about 30 species that are mostly perennial, some of which are rare or endangered, e.g. Lotus kunkelii and L . mascaënsis (BRAMWELL & BRAMWELL 2001). They cover a wide range of ecological conditions including littoral areas through to high- 97 © Landesmuseum für Kärnten; download www.landesmuseum.ktn.gv.at/wulfenia; www.biologiezentrum.at G. S ANDRAL , M. V. R E M I Z OWA & D. D. S OKOLOFF Table 1: Geographic distribution of species in the Lotus sect. Pedrosia. 28. 29. 25. 26. 27. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 6. 7. 8. 9. 10. 11. 12. 13. 1. 2. 3. 4. 5. Lotus arborescens Lotus jacobaeus Lotus purpureus Lotus brunneri Lotus latifolius Lotus glaucus Lotus dumetorum Lotus tenellus Lotus sessilifolius Lotus kunkelii Lotus arinagensis Lotus mascaënsis Lotus spartioides Lotus campylocladus Lotus emeroides Lotus callis-viridis Lotus loweanus Lotus macranthus Lotus agryrodes Lotus lancerottensis Lotus assakensis Lotus pseudocreticus Lotus creticus Lotus chazaliei Lotus arenarius Lotus maroccanus Lotus eriosolen Lotus jolyi Lotus tibesticus Distribution Azores Santa Maria + São Miguel + Terceira + Graciosa São Jorge + Pico + Faial Flores Corvo Madeira Islands Madeira + + + + Desertas + + Porto Santo + + + + Salvage Islands + Canary Islands Lanzarote + Fuerteventura + + Gran Canaria + + + + + Tenerife + + + + + + La Palma + La Gomera + + El Hierro + Cape Verde Santo Antão + + São Nicolau + + São Vicente + Sal + Boa Vista + + Maio + Santiago + + Fogo + + Brava + Mainland Spain & + + Portugal Mainland Africa + + + + + + + + + Remaining part of + Mediterranean region altitude pine forests and have a center of diversity in the Macaronesian Islands (tab. 1) of the Atlantic Ocean (i.e. Cape Verde, the Canaries, Salvage Islands, Madeira, and the Azores). The section is also present in mainland Africa and in the Mediterranean region. In Macaronesia there are only three sections (Canaria, Rhyncholotus, Pedrosia) of the tribe Loteae which includes 18 genera (SOKOLOFF 2003a) exhibiting considerable diversification. All three sections belong 98 © Landesmuseum für Kärnten; download www.landesmuseum.ktn.gv.at/wulfenia; www.biologiezentrum.at A taxonomic survey of Lotus section Pedrosia Figure 1: Flower of Lotus berthelotii. Scale bar = 1 mm. to the genus Lotus, and the section Pedrosia is more diverse than the other two. The section Canaria is taxonomically isolated (DEGTJAREVA et al. 2006) and was probably established in the Canaries before the radiation of Pedrosia. Other genera of the tribe do not have species endemic to the Macaronesian Islands, except for Anthyllis, which has a single and morphologically poorly differentiated endemic species on Madeira. The most important diagnostic character of the section Pedrosia is the presence of a tooth on the ventral side of the stylodium (style) (KRAMINA & SOKOLOFF 1999). This feature is shared with the section Rhyncholotus, which species (L. berthelotii and its allies) are endemic to the Canary Islands. Previously this section was accepted as Lotus sect. Heinekenia (nom. illeg.) by BRAND (1898), as a distinct genus Heinekenia by CHRIST (1888) and KUNKEL (1974), as a subgenus Rhyncholotus by MONOD (1980) and as section Rhyncholotus by SOKOLOFF (2003a). The section Rhyncholotus differs from the section Pedrosia as it includes plants that have a much more specialized corolla structure (fig. 1), that is linked with an adaptation to pollination by birds. Phylogenetic data clearly show monophyly of a clade comprising all studied members of Pedrosia and Rhyncholotus (ALLAN et al. 2004; DEGTJAREVA et al. 2006). Relationships within this clade are not completely resolved, however, it seems that Rhyncholotus is embedded within the paraphyletic Pedrosia. Molecular data (nuclear ribosomal ITS) alone does not show monophyly of Rhyncholotus (ALLAN et al. 2004) while analyses of a combined data set (nrITS plus morphology) revealed its monophyly (DEGTJAREVA et al. 2006). This could imply, that species of Rhyncholotus should be better placed into the broadly accepted section Pedrosia. We, however, do not pursue this amalgamation here as the phylogeny of the Lotus berthelotii-group is not well resolved and, in addition, it may be reasonable to keep some sections of Lotus paraphyletic (see DEGTJAREVA et al. 2006 for discussion). These questions, however, are deserving of further study. Another uncertainty in the section Pedrosia is the taxonomic position of Lotus creticus. This species was traditionally placed into section Lotea (Ball & Chrtkova-Žertova 1968). It was commonly believed, that L . creticus, as well as other species of the section Lotea, do not possess a tooth on 99 © Landesmuseum für Kärnten; download www.landesmuseum.ktn.gv.at/wulfenia; www.biologiezentrum.at G. S ANDRAL , M. V. R E M I Z OWA & D. D. S OKOLOFF the stylodium. However, KRAMINA & SOKOLOFF (1999) described the presence of a small tooth on the stylodium in some specimens of L . creticus. They suggested, that L . creticus can be transferred into the section (subgenus) Pedrosia. VALDÉS (2000) also described a tooth on the stylodium of L . creticus, but placed the species in the section Lotea. Molecular phylogenetic studies so far have not clarified these disagreements. For example, ALLAN et al. (2003, 2004) found L . creticus nested within the section Lotea while DEGTJAREVA et al. (2006) found its placement within the Pedrosia-Rhyncholotus clade. Certainly more molecular phylogenetic data are needed to clarify the placement of L . creticus. In this study, we place L . creticus tentatively in the section Pedrosia as our new morphological observations confirm earlier findings byK RAMINA & SOKOLOFF (1999), that L . creticus is morphologically and ecologically similar to L . pseudocreticus of the section Pedrosia. Providing a more definitive assessment of the position ofL . creticus will be the subject of further studies by the authors and their colleagues in Moscow. Species-level taxonomy of the section Pedrosia is problematic. Since BRAND’s (1998) monographic
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