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A Taxonomic Review of the Dry-Fruited Species of Anemone (Ranunculaceae) in Southern Africa

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A Taxonomic Review of the Dry-Fruited Species of Anemone (Ranunculaceae) in Southern Africa Bothalia 43,1: 1–13 (2013) A taxonomic review of the dry-fruited species of Anemone (Ranunculaceae) in southern Africa J.C. MANNING* and P. GOLDBLATT** Keywords: Anemone L., classifi cation, nomenclature, Ranunculaceae, southern Africa, taxonomy ABSTRACT The three dry-fruited species of Anemone sect. Pulsatilloides subsect. Alchemillifoliae (Ranunculaceae) from southern Africa are reviewed, with full descriptions and nomenclature, including complete synonomy, taxonomic history with nomen- clatural corrections, ecological notes, and distribution. A. tenuifolia (L.f.) DC. from the Cape Floristic Region is segregated as ser. Pinnatifoliae from the two summer rainfall species, A. caffra (Eckl. & Zeyh.) Harv. and A. fanninnii Harv. ex Masters, which remain in ser. Alchemillifoliae, emphasising the strong vegetative differences between the two series. INTRODUCTION intimate relationship to the southern African Anemone species with dry fruits, with which it shares apomor- Generic relationships of tribe Anemoneae DC. phic pantoporate pollen, and all of the southern African (Ranunculaceae–Ranunculoideae) have been the sub- species are now associated as subsect. Alchemillifoliae ject of intensive morphological and molecular investi- (Ulbrich) Hoot of A. sect. Pulsatilloides DC. (Hoot et al. gations (Hoot et al. 1994; Ehrendorfer & Samuel 2001; 2012). The fl eshy-fruited species, which constitute ser. Schuettpelz et al. 2002; Hoot et al. 2012), resulting in Knowltonia (Salisb.) J.C.Manning & Goldblatt within the recent expansion of the circumscription of Anemone subsect. Alchemillifoliae, have been well monographed L. to include the small segregates Barneoudia C.Gay (3 by Rasmussen (1979) [as the genus Knowltonia]. This is spp.), Hepatica Miller (7 spp.), Knowltonia Salisb. (8 not the case with the dry-fruited southern African spe- spp.), Oreithales Schldl. (1 sp.) and Pulsatilla Miller (± cies, which, as yet, have been treated only incompletely 38 spp.) (Tamura 1993). Second largest genus of tribe or superfi cially (De Candolle 1824; Pritzel 1841; Harvey Anemoneae after Clematis L., Anemone s. l. is diag- 1860; Ulbrich 1906), most recently as part of a wide- nosed by the presence of one or more leafy, cauline ranging review of all austral species of Anemone (Ziman involucres beneath the fl ower, and a perianth comprised of imbricate, petaloid sepals only. It includes ± 200 spe- et al. 2006). Although very valuable, this latter treat- cies distributed throughout the world, primarily in the ment is nomenclaturally incomplete and also perpetuates Northern Hemisphere with only a modest representa- some errors, notably the incorrect name and authors for tion on the southern continents, where ± 30 species are A. tenuifolia (L.f.) DC., as well as containing some mis- recorded mainly from montane regions with a temperate takes in typifi cation. We provide a complete review of climate (Ziman et al. 2006). The austral representatives the taxonomy and nomenclature of the dry-fruited spe- are concentrated in South America (12 spp.) and south- cies of Anemone from southern Africa, including full ern Africa (11 spp.), with A. thomsonii Oliver endemic synonomy, taxonomic history, ecological notes and dis- to east tropical Africa and a handful of additional species tribution. in Indonesia and Australasia (Hoot et al. 2012). Taxonomic relationships among the dry-fruited south- Anemone in southern Africa traditionally included ern African species of Anemone were fi rst formalised just three species with dry fruits, but has since been by Ulbrich (1906), who segregated A. tenuifolia [as A. enlarged to 11 spp. by the recent transfer of the eight capensis] in the monotypic ser. Pinnatifoliae; retain- fl eshy-fruited species of Knowltonia (Manning et al. ing A. caffra and A. fanninii, the two rosulate species 2009). Although previously treated in Anemone by some with unlignifi ed caudices and palmate leaves, in ser. authors, this close-knit group of species was generally Alchemillifoliae. This treatment, with an emphasis on retained as a distinct genus on the basis of its compound morphology, was followed by subsequent authors until infl orescences of relatively small fl owers and its bac- recently when Hoot et al. (2012) weighted phyloge- cate or berry-like fruits. Phylogenetic analysis of plastid netic considerations by including all three species in an and nuclear DNA sequence data has now confi rmed its expanded ser. Alchemillifoliae within subsect. Alchemil- lifoliae, arguing that the two leaf morphs of A. tenuifolia differed more from one another in DNA base sequences * Compton Herbarium, South African National Biodiversity Institute, than did A. caffra from A. fanninii. There is no com- Private Bag X7, Claremont 7735, South Africa; Research Centre for pelling morphological basis for this treatment, since A. Plant Growth and Development, School of Life Sciences, University of KwaZulu-Natal, Pietermaritzburg, Private Bag X01, Scottsville 3209, tenuifolia is intermediate between ser. Knowltonia and South Africa. E-mail: [email protected]. ser. Alchemillifoliae in having the compound foliage of ** B.A. Krukoff Curator of African Botany, Missouri Botanical Garden, the former but the sericeous, fusiform achenes of the lat- P.O. Box 299, St. Louis, Missouri 63166, USA; Research Centre for ter. The molecular topology retrieves ser. Knowltonia Plant Growth and Development, School of Life Sciences, University of KwaZulu-Natal, Pietermaritzburg, Private Bag X01, Scottsville 3209, as sister to a clade in which A. tenuifolia is in turn sis- South Africa. E-mail: [email protected]. ter to A. caffra + A. fanninii and is thus consistent with MS. received: 2012-07-06. either classifi cation. Signifi cantly, however, the cladistic 2 Bothalia 43,1 (2013) branches subtending A. caffra / A. fanninii, the two leaf caffra (Eckl. & Zeyh.) Harv. morphs of A. tenuifolia, and the members of ser. Knowl- tonia in the molecular phylogram presented by Hoot et Plants with horizontal or ascending caudices. Leaves al. (2012) are all of similar length, indicating compara- palmate or pinnately compound, toothed. Infl orescence ble levels of sequence divergence of the three lineages simple or once- or twice-compound. Flowers with from their last common ancestor. This observation, cou- numerous (10–20) ± linear to narrowly elliptical sepals. pled with the clear morphological and ecological dif- Achenes numerous, usually four times longer than wide, ferences among the lineages, prompts us to revert to glabrous or tomentose, dry or fl eshy. Pollen pantoporate. the segregation of the dry-fruited species in two series, 11 spp., temperate southern Africa with the fl eshy-fruited species comprising a third series. This classifi cation, which incorporates both phenetic and Key to series phylogenetic information, seems more appropriate to us than the alternative that stresses the slightly closer phy- 1a. Leaves palmately lobed . ser. Alchemillifoliae logenetic relationship between the dry-fruited species 1b. Leaves ternately compound (rarely simple): over their obvious morphological diversity. We do not 2a. Infl orescence simple, 1-fl owered (rarely with up to two consider ser. Knowltonia further here, and readers are secondary, involucellate fl owers); tepals larger, 20–40 referred to the taxonomic revision by Rasmussen (1979) mm long; fruits fusiform, dry, densely silky ser. Pinnatifoliae and the revised generic placement by Manning et al. 2b. Infl orescence compound, several- to many-fl owered; tepals smaller, 10–20 mm long; fruits ovoid or ellipsoid, (2009) for further details. ± fl eshy, glabrous to puberulous . ser. Knowltonia [see Rasmussen (1979) for taxonomic account] MATERIALS AND METHODS Key to dry-fruited species (ser. Alchemillifoliae and Pinnatifoliae) All relevant types were examined, as well as all spec- imens from BOL, MO, NBG, PRE and SAM (acronyms 1a. Subshrub with woody caudex; leaves ternately decom- pound, sclerophyllous, glabrescent, margins revolute . after Holmgren et al. 1990), the herbaria housing the . 1. A. tenuifolia most comprehensive collections of southern African spe- 1b. Rosulate perennials; leaves palmate, leathery, persistently cies. All species were also studied in the fi eld. pubescent, margins plane: 2a. Plants smaller, peduncle 100–200 mm long; leaf blades (40–)60–100(–130) mm diam., mostly cleft less than TAXONOMY halfway, usually thinly pubescent above and glabrescent or thinly pubescent beneath, mainly along nerves, rarely Anemone L., Species plantarum: 538 (1753). Type: densely villous beneath, petioles 40–150 mm long; fl ow- A. coronaria L. ers 1(2), sepals dimorphic or ± monomorphic, outer usu- ally only thinly sericeous beneath . 2. A. caffra Knowltonia Salisb.: 372 (1796). Type: Knowltonia 2b. Plants larger, peduncle 200–800 mm long; leaf blades rigida Salisb., nom. illegit. = Anemone knowltonia Burtt 150–350 mm diam., mostly cleft more than halfway, thickly velvety above and densely villous beneath, peti- Davy oles 200–700 mm long; fl owers (1)2–4, sepals dimor- phic, outer ± densely sericeous or villous beneath 3. A. fanninii Perennial, rhizomatous herbs, rarely shrublets. Leaves basal, alternate, simple and palmate or terna- ser. Pinnatifoliae Ulbrich, Botanische Jahrbücher fur tely compound, toothed, petiole base sheathing; fl ower- Systematik, Pfl anzengeschichte und Pfl anengeographie ing stem with 1–5 whorls of 2–4, partly fused leaves. 37: 239 (1906). Type: A. capensis (L.) DC., hom. illegit. Infl orescence 1-fl owered or of di- or tri-chasial, umbel-
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