Taxonomic revision of the tropical African group of Carex subsect. Elatae (sect. Spirostachyae, Cyperaceae)

Taxonomic revision of the tropical African group of Carex subsect. Elatae (sect. Spirostachyae, Cyperaceae) by Marcial Escudero1& Modesto Luceño2 1 Pablo de Olavide University, Ctra. de Utrera km 1, E-41013 Seville, Spain. [email protected] 2 Author for correspondence: Department of Molecular Biology and Biochemical Engineering, Pablo de Olavide University, Ctra. de Utrera km 1, E-41013 Seville, Spain [email protected]; fax: + 34 954349813 Anales del Jardín Botánico de Madrid Vol. 68(2): 225-247 julio-diciembre 2011 ISSN: 0211-1322 doi: 10.3989/ajbm.2256 Abstract Escudero, M. & Luceño, M. 2011. Taxonomic revision of the tropical African group of Carex subsect. Elatae (sect. Spirostachyae, Cyperaceae). Anales Jard. Bot. Madrid 68(2): 225-247. The tropical African monophyletic group of Carex subsect. Elatae (sect. Spirostachyae) is distributed in continental tropical Africa, Madagascar, the Mascarene archipelago, and Bioko Island (32 km off the coast of West Africa, in the Gulf of Guinea). The first monographic treatment of this Carex group, as well as of the tribe Cariceae, was published by Kükenthal (as sect. Elatae Kük.). Recently, the first molecular (nrDNA, cpDNA) phylogeny of Carex sect. Elatae has been published, which also included the species of sect. Spirostachyae. In the resulting consensus trees, most species of sect. Elatae were embedded within core Spirostachyae and so this section was joined with sect. Spirostachyae as subsect. Elatae. Within subsect. Elatae, several groups were described, one of which was termed the “tropical African group”. Here we present a taxonomic revision of this group, based on more than 280 vouchers from 29 herbaria as well as in field trips in Tropical Africa. In the revision, we recognise 12 species (16 taxa) within the tropical African group, and so have somewhat modified our previous view, in which 10 species, 12 taxa were listed. One new species from Tanzania is included in this treatment, C. uluguruensis Luceño & M. Escudero. Several combinations are made, C. cyrtosaccus is treated as a synonym of C. vallis-rosetto and, finally, the binomial C. greenwayi has been recognised. Keywords: Bioko, Elatae, Carex, Cyperaceae, Madagascar, Mascarene Islands, Spirostachyae, tropical Africa. Resumen Escudero, M. & Luceño, M. 2011. Revisión taxonómica del grupo tropical africano de Carex subsect. Elatae (sect. Spirostachyae, Cyperaceae). Anales Jard. Bot. Madrid 68(2): 225-247 (en inglés). Las especies de la subsección Elatae (sección Spirostachyae) del género Carex que se distribuyen por África tropical continental, Madagascar, el archipiélago de las Mascareñas y la isla de Bioko (a 32 km del litoral de África occidental, en el golfo de Guinea) forman un grupo monofilético. El primer tratamiento taxonómico de este grupo de cárices, así como de la tribu Cariceae en su conjunto, fue elaborado por Kükenthal (sección Elatae Kük.); recientemente, se ha publicado el primer estudio de filogenia mo - lecular (nrDNA, cpDNA) de los táxones de este grupo, junto con la inmensa mayoría de las restantes especies que Kükenthal incluyó en las secciones Elatae y Spirostachyae. Salvo escasas excepciones, los táxones incluidos se agruparon en un clado (“core Spirostachyae”) que contiene las especies de las dos secciones antes aludidas. En congruencia con estos resultados, la sección Elatae fue incluida en la sección Spirostachyae como susbsección Elatae. Dentro de dicha subsección se observaron varios clados; uno de ellos, el denominado “grupo de África tropical”, es el objeto del presente trabajo, en el que presentamos una revisión taxonómica basada en el estudio de más de 280 pliegos conservados en 29 herbarios, así como en trabajos de campo llevados a cabo en África tropical. Como resultado del análisis de dichos materiales hemos realizado una profunda reorganización taxonómica, fruto de la cual admitimos 12 especies (16 táxones) dentro del grupo de África tropical. Se describe además una nueva especie del NE de Tanzania, C. uluguruensis Luceño & M. Escudero. Asimismo, se llevan a cabo varias combinaciones nomenclaturales, el nombre de C. cyrtosaccus es considerado sinónimo de C. vallis-rosetto, y se reconoce el binomio C. grennwayi. Palabras clave: Bioko, Elatae, Carex, Cyperaceae, Madagascar, islas Mascareñas, Spirostachyae, África tropical. 2256 carex_Maquetación 1 28/11/2011 11:46 Página 225 Introduction The tropical African group of Carex subsect. Elatae (sect. Spirostachyae) is a monophyletic group according to nrDNA phylogeny, but not according to cpDNA phylogeny (Escudero & Luceño, 2009). This group is distributed along continental tropical Africa as well as Madagascar and the Mascarene and Bioko Islands. Lamarck (1792) described the first species of this Carex group from Reunion Island (C. borbonica); other taxa were subsequently described: C. boryana (Schkuhr, 1806), C. petitiana and C. simensis (Richard, 1850), C. fischeri, C. vallis-rosetto, and C. lon gipedunculata (Schumann, 1895). Clarke (1902) treated the continental species of this group in his Flora of Tropical Africa (seven species in total, one of them, C. cyrtosaccus, as new species; Table 1), but he did not refer them to any supraspecific category. The first monographic treatment of this Carex group, as well as of tribe Cariceae, was published by Kükenthal (1909). He included 19 species in the new sect. Elatae Kük., eight of which (15 taxa) were from the tropical African group (Table 1). In this monographic work, one new species (C. mildbraediana) as well as five new varieties and two new combinations were published (Table 1). Subsequently, Kükenthal (1914, 1925, 1934; Table 1) published numerous modifications to his original taxonomic treatment of sect. Elatae (Kükenthal, 1909, Table 1), in which one subspecies, six new varieties and two new forms were described and a new combination was made. Nelmes (1938) revised the group and described four new taxa (three species and one variety) and published a new combination. Subsequent treatments were local, as in the Flora of West Tropical Africa (Hooper & Napper, 1972; two species), or The Sedges and Rushes of East Africa (Haines & Lye, 1983; seven species, Table 1). More recently, Luceño (1992) and Luceño & Castroviejo (1993) studied five taxa which were included in sect. Elatae by Kükenthal (1909) but none were from the tropical African group. Escudero et al. (2008) made the first molecular (nrDNA) phylogeny of Carex sect. Spirostachyae in a study that also included seven species previously treated in sect. Elatae by Kükenthal (1909), one of which was from the tropical African group (Carex mannii). These seven species were included in the core Spirostachyae in the molecular phylogeny. Escudero & Luceño (2009) performed the first molecular (nrDNA, cpDNA) phylogeny of Carex sect. Elatae that also included the species of sect. Spirostachyae. Again, most species of sect. Elatae were embedded within core Spirostachyae and accordingly, sect. Elatae was included in sect. Spirostachyae as susbsect. Elatae. Therefore, the new M. Escudero & M. Luceño taxonomic concept of this section includes 11 species traditionally considered in sect. Spirostachyae and 16 species traditionally included in sect. Elatae (Escu - dero & Luceño, 2009). Within subsect. Elatae, several groups were described, among them the tropical African group (Escudero & Luceño, 2009). This group contained ten species, eight (ten taxa) from continental tropical Africa and three species from Madagascar, and the Mascarene and Bioko Islands (one of them in the continent too). In addition, two new combinations were made (C. mannii ssp. tho - masii and C. mannii ssp. friesiorum) (Table 1). Finally, a checklist of sub-Saharian African Carex species has been recently published (Gehrke, 2011). The general goal of this study is to provide a taxonomic treatment of this controversial tropical African group of Carex. The specific aims are: 1) to establish the taxa that belong to this group, 2) to specify the taxonomical level of the taxa (species or subspecies), 3) to delineate the morphological differences between the taxa and provide taxonomic keys, 4) to describe the new taxa which are necessary, and 5) to perform a nomenclatural revision. Material and methods More than 280 herbarium vouchers were studied for the present taxonomic revision (see Appendix 1). These materials are from 29 herbaria (B, BM, BR, C, E, EA, GOET, H, HUH, K, L, LD, LISC, M, MA, MO, NBG, NU, NY, O, OXF, PRE, U, UPOS, UPS, US, W, WAG, WU; Index Herbariorum, http://sweetgum.nybg.org/ih/). In addition the website JSTOR (http://www.jstor.org/) which contains images of type material of African species was frequently consulted in the present revision. Seventy morphological characters were considered in the study, comprising 37 qualitative characters and 33 quantitative characters. We paid special attention to the characters previously stated as important in the taxonomy of sect. Spi ros tachyae (Luceño & Escudero, 2008). Quantitative characters were measured using a stereoscopic bi no cular Nikon SMZ645. Taxonomic treatment After revision of the herbarium material, and taking into account our previous molecular results, 12 species (16 taxa) have been accepted. We describe a new species from Tanzania (C. uluguruensis Luceño & M. Escudero) and we include an undescribed species, Carex spec. nov., endemic from Kitulo Plateau (Tanzania). Additionally, one new subspecies (C. fischeri ssp. recedens) has been proposed. The morphological 226 Anales del Jardín Botánico de Madrid 68(2): 225-247, julio-diciembre 2011. ISSN: 0211-1322. doi: 10.3989/ajbm: 2256 2256 carex_Maquetación 1 28/11/2011 11:46 Página 226 Taxonomic revision Carex tropical African group 227 Anales del Jardín Botánico de Madrid 68(2): 225-247, julio-diciembre 2011. ISSN: 0211-1322. doi: 10.3989/ajbm: 2256 Table 1. Taxonomic treatments by Clarke (1902), Kükenthal (1909, 1914, 1925, 1934), Nelmes (1938), Haines & Lye (1983), Escudero & Luceño (2009) and Gehrke (2011). Clarke (1902) Kükenthal (1909) Kükenthal (1914+, 1925*, Nelmes (1938) Haines & Lye (1983) Escudero & Luceño (2009) Gehrke (2011) 7 spp. (1 new sp.) 8 spp. 15 taxa (1 new sp., 1934×), × 1 new var., + 1 new f., 9 spp., 11 taxa (3 new spp., 7 spp. 10 spp., 12 taxa 9 spp., 11 taxa 5 new var., 2 new comb.) * 1 rearrangement, 1 new f., 1 new var., 1 comb.) (2 new comb.) 1 new ssp., 5 new var. C. vallis-rosetto C. vallis-rosetto K. Schum. × C. longipedunculata subsp. C. vallis-rosetto K. Schum. C. mannii E.A. Bruce C. mannii E.A. Bruce C. elgonensis Nelmes K. Schum. var. vallis-rosetto cuprea var. atrennata Kük. C. simensis Hochst. C. vallis-rosetto + C. longipedunculata C. cyrtosaccus C.B. Clarke C. thomasii Nelmes C. mannii ssp. thomasii C. mannii E.A. Bruce ex A. Rich. var. purpurea Kük. K. Schum.f. recedens Kük. as species dubiae (Nelmes) Luceño & M. Escudero C. longipedunculata C. boryana Schkuhr * C. vallis-rosetto K. Schum. C. greenwayi Nelmes C. elgonensis Nelmes C. mannii subsp. C. mannii ssp. thomasii K. Schum. var. boryana (incl. var. purpurea Kük.) (= C. vallis-rosetto friesiorum (Kük.) (Nelmes) var. purpurea Kük.) Luceño & M. Escudero Luceño & M. Escudero C. petitiana A. Rich. C. boryana .* C. vallis-rosetto C. thomasii Nelmes C. vallis-rosetto K. Schum. C. elgonensis Nelmes C. mannii ssp. friesiorum var. simplicissima Kük f. ramosa Kük. (Kük.) Luceño & M. Escudero C. fischeri K. Schum. C. cyrtosaccus C.B. Clarke * C. vallis-rosetto C. elgonensis Nelmes C. cyrtosaccus C.B. Clarke C. vallis-rosetto K. Schum. C. petitiana A. Rich. var. heterostachya Kük. C. boryana Schkuhr C. fischeri K. Schum. * C. mildbraediana C. mildbraediana Kük. C. petitiana A. Rich. C. cyrtosaccus C.B. Clarke C. preussii K. Schum. var. friesiorum Kük. C. cyrtosaccus C. longipedunculata * C. mildbraediana C. longipedunculata C. simensis Hochst. C. petitiana A. Rich. C. simensis Hochst. C.B. Clake K. Schum. var. alpicola Kük. K. Schum. ex A. Rich. ex A. Rich. var. longipedunculata C. longipedunculata * C. fischeri var. basiandra Kük. C. simensis Hochst. . C. simensis Hochst. C. fischeri K. Schum. var. preussii (K. Schum.) Kük ex A. Rich. ex A. Rich. C. longipedunculata * C. longipedunculata C. preussii K. Schum. C. mildbraediana Kük. C. ninagongensis var. ninagongensis Kük. subsp. cuprea Kük. (Kük.) Nelmes C. simensis Hochst. ex. A. Rich. * C. simensis C. preussii C. fischeri K. Schum. C. vallis-rosetto K. Schum var. simensis var. longispitata Kük. var. camerunensis Nelmes 2256 carex_Maquetación 1 28/11/2011 11:46 Página 227 M. Escudero & M. Luceño differences of this new subspecies were noted in Escu - dero & Luceño (2009), but this new taxon was not formally described in that study. The populations of C. petitiana from Zimbabwe and Malawi have been referred to C. petitiana subsp. attenuata (Kük.) Luceño & M. Escudero, C. cyrtosaccus has been treated as a synonym of C. vallis-rosetto, and the binomial C. greenwayi has been accepted following Nelmes (1938). KEY TO THE TROPICAL AFRICAN GROUP OF CAREX SUBSECT. ELATAE 1. Plants with rhizome creeping, with ± long internodes; spikes never gynecandrous..................................... 8. C. spec. nov. 1. Plants caespitose, rhizome with short internodes; very exceptionally rhizome creeping, with ± long internodes, but then gynecandrous spikes are present ........................................ 2 2. Utricles clearly curved .......................... 12. C. vallis-rosetto 2. Utricles straight .................................................................. 3 3. Most female glumes with an arista or mucro longer than 2.5 mm ................................................................ 2. C. boryana 3. Most female glumes with an arista or mucro shorter than 2.5 mm .................................................................................... 4 4. Beak of the utricle truncate or bidentate (dorsal sinus rarely more than 0.1 mm deep) ................................................... 5 4. Beak of the utricle more or less bifid (dorsal sinus more than 0.2 mm deep) ..................................................................... 6 5. Leaves more than 11 mm wide; at least some of the spikes arising in groups of three .................... 7. C. mildbraediana 5. Leaves less than 10 mm wide; all spikes arising in pairs or singly ................................................... 11. C. uluguruensis 6. Female glumes dark blackish-purple without midribs or with midribs that are indistinct and thin ............. 9. C. elgonensis 6. Female glumes rarely dark blackish-purple, but then with distinct midribs that are straw coloured or greenish ................ 7 7. Basal sheaths and sheath of the lowest leaves purplish-red to dark purplish or blackish tinted ........................................... 8 7. Basal sheaths and sheath of the lowest leaves straw coloured, brown or reddish-brown .................................................. 12 8. All spikes androgynous (only a few male flowers at the top) and homomorphic or sub-homomorphic (similar number of male flowers at the top of all spikes); leaves wider than 8 mm ................................................................... 5. C. greenwayi 8. Some spikes not androgynous and homomorphic or subhomomorphic; when so, leaves up to 8 mm wide .................. 9 9. Female glumes hyaline, brown or reddish brown ............... ..................................................................... 4. C. fischeri 9. Male glumes medium to dark purple or blackish ........... 10 10. Female glumes blackish; basal sheaths blackish and coriaceous .......................................................... 10. C. simensis 10. Female glumes brown to dark purplish; basal sheaths variable ............................................................................... 11 11. Leaves markedly coriaceous ........................ 1. C. borbonica 11. Leaves non-coriaceous ..................................... 6. C. mannii 12. Two or more lateral spikes gynecandrous (sometimes these spikes bear only a few male flowers at the base); spikes widely terete or clavate; utricle beak aculeolate (very exceptionally smooth) ........................................................ 9. C. petitiana 12. None, or only one, lateral spike gynecandrous (rarely two); spikes narrowly terete or clavate; utricle beak smooth ........ ..................................................................... 4. C. fischeri 228 Anales del Jardín Botánico de Madrid 68(2): 225-247, julio-diciembre 2011. ISSN: 0211-1322. doi: 10.3989/ajbm: 2256 Table 1. Taxonomic treatments by Clarke (1902), Kükenthal (1909, 1914, 1925, 1934), Nelmes (1938), Haines & Lye (1983), Escudero & Luceño (2009) and Gehrke (2011). Clarke (1902) Kükenthal (1909) Kükenthal (1914+, 1925*, Nelmes (1938) Haines & Lye (1983) Escudero & Luceño (2009) Gehrke (2011) 7 spp. (1 new sp.) 8 spp. 15 taxa (1 new sp., 1934×), × 1 new var., + 1 new f., 9 spp., 11 taxa (3 new spp., 7 spp. 10 spp., 12 taxa 9 spp., 11 taxa 5 new var., 2 new comb.) * 1 rearrangement, 1 new f., 1 new var., 1 comb.) (2 new comb.) 1 new ssp., 5 new var. C. simensis var. stolonifera *C. simensis var. ninagongensis C. cuprea (Kük.) Nelmes C. boryana Schkuhr C. boryana Schkuhr (Boeck.) Kük. (Kük.) Kük. (= C. simensis var. stolonifera (Boeck.) Kük. = C. longipedunculata subsp. cuprea Kük.) C. simensis var. mauensis Kük C. borbonica Lam. C. borbonica Lam. var. borbonica C. borbonica var. pallidior Kük. C. mildbraediana Kük. 2256 carex_Maquetación 1 28/11/2011 11:46 Página 228 Species descriptions 1. Carex borbonica Lam., Encycl. 3: 387. 1792. Ind. loc.: “Commerson a trouvé cette spèce dans l’isle de Bourbon”. Lectotype (designated here): P 307290 (Herb. Lamarck) photo!; isotypes: P 346058 photo!, P 346059 photo! = Carex typhoides Bory, Voy. îles Afrique 3: 24 (1804). Unknown type. Rhizome caespitose, with short internodes. Stems 15-59 cm, stout, trigonous, green to yellowish-green, smooth, sometimes purplish-red spotted. Leaves 10- 18 cm × 4-8 mm, keeled, sometimes slightly plicate in the upper half, rigid and markedly coriaceous, scabrid on the edges and apical parts; lower sheaths dark purplish-red; ligule 0-5.5 mm, obtuse to acute; anteligule edge concave to flat. Inflorescence 6.2-18.5 cm, lowest internode 0.8-3.5 cm and second one 0.5-2.5 cm. Lowest bract 5.5-17 cm × 2.5-6 mm, longer or shorter than inflorescence; sheath 1-7.5 cm long, the inner side purplish-red. Spikes 3-10, heteromorphic to subhomomorphic, with 0-2 male or gynecandrous spikes at the apex, and 3-9 female lateral spikes; male or gynecandrous spikes 3-6 cm × 6-12 mm, terete to widely ellipsoid or obovoid, the terminal one sessile or with a peduncle up to 1.2 cm; female spikes, 1-6.5 cm × 6- 10 mm, arising singly, dense-flowered, terete to obovoid, sessile or with a peduncle up to 1(3.5) cm. Male glumes (4.5)5.5-9 × 1.2-2 mm, oblong-elliptic to oblong-obovate, dark purplish-red with a wide, green or straw coloured midrib, mucronate or aristate, with a mucro or arista up to 1.7 mm or, more rarely, obtuse. Female glumes (4)5.5-6.5 × 1.1-2.2 mm, ovate-elliptic, dark purplish-red with a wide, green or straw co lour - ed midrib, with an arista or mucro 0.5-2.3 mm. Utricles (4)4.5-6.2 × (1.2)1.4-2(2.1) mm, ellipsoid-trigo- nous, straight, plurinerved (with 2 more prominent veins), tapered to beak, brown with purplish speckles to entirely purplish; beak (0.9)1.2-1.6 mm, deeply bifid, with ± similar dorsal and ventral sinus 0.4-0.6 mm, densely aculeolate. Achenes 2.2-2.3 × 0.9-1.2 mm, trigonous, elliptic or elliptic-obovate in outline. 2n =? Oct.-May. Dry, sunny and sandy places on volcanic soils; 2250-2400 m; Reunion [and Mauritius?]. Comments: Some specimens display intermediate features between C. boryana and C. borbonica, suggesting hybridization, and probably introgression, between these two taxa. We have observed in the field (Le Maïdo, near Malfate circus) both pure species in their typical habitats, with neighbouring populations Taxonomic revision Carex tropical African group growing in intermediate habitats (open and stony scrublands of Philippia montana) whose individuals showed a gradient of characters between the putative parentals. This hybridization hypothesis is in agreement with our molecular data (Escudero & Luceño, 2009). Many of these morphologically intermediate specimens have been described as different taxa: – Carex borbonica var. pallidior Kük., Engl. (ed.), Pflanzenr. IV, 20(38): 655. 1909. Sine ind. loc. Lectotype (designated here): Boivin 997 K! – Carex boryana var. rigidifolia Boeck., Linnaea 41: 285. 1877. Sine ind. loc. Lectotype (designated here): Boivin 996 UPS! – Carex musei Steud., Syn. Pl. Glumac. 2: 205. 1855. Ind. loc.: “ins. Borbon” [Reunion]. Lectotype (designated here): s.c. (P 541690 photo!). Although the type material of C. musei (P 541690!) is very similar to pure individuals of C. boryana, the short female glume aristas and most spikes arising near the apex of the stem lead us to think it is an introgressive form of the latter species with C. borbonica. Carex borbonica was thought to be also distributed on Mauritius island by Kükenthal (1909; “Mauritius (in herb. Lamarck!)”). However, all studied materials of C. borbonica from Lamarck’s herbarium were collected from Reunion where this species grows on volcanic soil above 2250 m, while on Mauritius the highest point is the Piton de la Petite Rivière Noire (828 m). Moreover, although Baker (1877) included C. bor- 229 Anales del Jardín Botánico de Madrid 68(2): 225-247, julio-diciembre 2011. ISSN: 0211-1322. doi: 10.3989/ajbm: 2256 Fig. 1. Distribution of Carex borbonica. 2256 carex_Maquetación 1 28/11/2011 11:46 Página 229 bonica in his Flora of Mauritius, this author indicated: “I have not seen Mauritian specimens”. For all these reasons, the references of this species for Mauritius are most likely mistaken. Selected specimens FRANCE. Reunion, Piton de la Fournaise, Plaine des Sables, 2250 m, 9-11-1966, H.J. Schlieben, 10894 ( BR, PRE, M 124775, B 100240041). Reunion, Plateu des Basaltes (sur Plaine des Remparts-Région du Volcan), 2400 m, 30-5-1966, T. Cadet, 451 (K). Reunion, 10-1875, I.B. Balfour (K). Reunion, just below Maidho W of St. Paul, 2-11-1968, C. Barclay, 1334 (K). Reunion, 15-2-1847, Boivin, 23bis (K). Reunion, Bory, 1003 (BM 898021). Reunion, Plaine des Remparts, Commerzon Crater, 2280 m, 1-1-2009, M. Luceño & M. Guzmán, 1ML09 (UPOS 3635). 2. Carex boryana Schkuhr, Beschr. Riedgräs. 2: 43, fig. 191. 1806. Ind. loc.: “In Insula Franciae”. Lectotype (designated here): J.B.G.G.M. Bory de St. Vicent [Reunion] (HAL 103608, Herb. C. Schkuhr, photo!); isotypes: P 459767 photo! = Carex subulata Wahlenb., Kongl. Vetensk. Acad. Nya Handl. 24: 152. 1803, nom. illeg., non J.F. Gmel., Syst. Nat. 2(1): 138. 1791 (homonyme post.). = Carex boryana var. minor Boott, Ill. Carex 3: 111. 1862. Ind. loc.: “Ile de la Réunion”. Lectotype (designated in Gehrke, 2011) L.H. Boivin (P 459763 photo!); isotype: P 459765 photo!; paratype: Boivin, 997 p.p. (P). = Carex giraudiana Steud., Syn. Pl. Glumac. 2: 209. 1855. Ind. loc.: “Ile Bourbon”. Lectotype (designated here): H. Giraudy (P 459766 photo!). Rhizome densely caespitose, with short internodes. Stems 75-163 cm, ± stout, trigonous, smooth, green to yellowish-green, sometimes purplish-red spotted. Leaves 25-48(63) cm × (2)4-15 mm, keeled to plicate, slightly scabrid on the edges and apical parts; lower sheaths dark purplish-red; ligule 2-2.5(9) mm, obtuse; anteligule edge concave to convex. Inflorescence (14)26-80(90) cm, lowest internode 8-30(45) cm and second one 1-24.5 cm. Lowest bract 17.5-56 cm × 2-9 mm, longer or shorter than the inflorescence; sheath 2.3-12.5(20.5) cm long, the inner side purplishred. Spikes 5-14, sub-homomorphic, sometimes heteromorphic, all of them androgynous, with the male part increasing in length along the stem, exceptionally 1-5 whole male spikes at the top; terminal spike 1-9.5 cm × 2-8(12) mm, linear to narrowly terete, sessil M. Escudero & M. Luceño or with a peduncle up to 1.2 cm; lateral spikes, 1.5- 9.5(11) cm × 4.5-8 mm, arising singly, in pairs or in groups of three, lax to subdense-flowered, linear to narrowly terete, usually bearing a few short branches at the base of the largest spikes, sessile or with a peduncle up to 4(6) cm. Male glumes 5.5-8.5(10) × (0.8)1-1.8 (2.1) mm, oblong-lanceolate, brown to reddish-brown with a wide, green or straw coloured midrib, aristate, with an arista 0.5-2.5 mm. Female glumes 4.5-8(9) × (0.8)1-1.6(2.2) mm, oblong-lanceolate to ovate, brown to reddish-brown with a wide, green or straw coloured midrib, with an arista or mucro (0.5)2.5-4.5(8) mm. Utricles (4)4.5-5.5 × 1.2-1.6 mm, ellipsoid-trigonous, straight, plurinerved (with 2 or more prominent veins), gradually or abruptly narrowed into a beak, light-green or brown with purplish-red speckles; beak (1)1.2- 2.5 mm, deeply bifid, with ± similar dorsal and ventral sinus 0.4-0.6 mm depth, smooth or aculeolate. Achenes (1.7) 2.2-2.5 × 1.1-1.4 mm, trigonous, oblongellipsoid (exceptionally oblongo-bovoid or subspheroid). 2n = 68. Sept.-Jan. In wet forests and other shady places; 1300-2350 m; Reunion, [Mauritius and Madagascar ?]. Comments: Kükenthal (1909) described C. boryana var. boryana as distributed in Reunion, Madagascar (“Perville P! in herb. Musei Paris”, material not seen in the present study) and Mauritius (no materials were cited for this island). In addition, Baker (1877) included this taxon in his Flora of Mauritius. However, we have not seen any material of C. boryana from Madagascar or Mauritius, although the presence of this species on the latter island is plausible. No Carex material studied by us from Madagascar 230 Anales del Jardín Botánico de Madrid 68(2): 225-247, julio-diciembre 2011. ISSN: 0211-1322. doi: 10.3989/ajbm: 2256 Fig. 2. Distribution of Carex boryana. 2256 carex_Maquetación 1 28/11/2011 11:46 Página 230 belongs to subsect. Elataeexcept perhaps the specimen “P. de la Bathie 13391 [at K], Ankaratra, Tsiafajavona Mts”; but this material is somewhat similar to C. elgonensis and therefore very different to C. boryana. Some of the specimens used by Kükenthal (1909) to describe C. boryana var. simplicissima have been included in our concept of C. manni (Mann, 1478 pho- to!), whilst others (Boivin 997 K!) are intermediate forms between C. boryana and C. borbonica. Selected specimens FRANCE. Reunion, Boivin, 996 (UPS). Reunion, le Tévelave, near Les Avirons, Westcoast, Mistforest, 1300 m, 4-11-1966, H.J. Schlieben, 10834a (PRE). Reunion, 1837, M. Richard, 530, (K). Reu - nion, 24-1-1952, Spach (K). Reunion, 1867, Hooker (K). Reunion, 11-1921, Hienthmay, 10162 (M 124764). Reunion, Herville, Nos- si.Bi, 2-12-1906, C.H. D’Alleizette (L 625308). Reunion, 9-1851, M. Boivin (US 2493537). Reunion, 1877, R.J. Shuttleworth (BM 898005). Reunion (C). Reunion, Curepipe, 21-11-1888, H.H. Johnston (E 219443). Reunion, between La Plaine des Cafres and La Paline des Palmistes, Col de Bellevue, 1615 m, 31-12-2008, M. Luceño & M. Guzmán, 127ML08 (UPOS 3732). Reunion, Piton de la Fournaise, crater, 2350 m, 1-1-2009, M. Luceño & M. Guzmán, 2ML09 (UPOS 3636). Reunion, cirque Cilaos, Bras Sec, 1315 m, 4-1-2009, M. Luceño & M. Guzmán, 13ML09 (UPOS). Reunion, Les Makes, La Fenetre, 1650 m, 5-1-2009, M. Luceño & M. Guzmán, 14ML09 (UPOS 3637 to 3641). Reunion, W cirque Malfate, Le Maido, 2200 m, 6-1-2009, M. Luceño & M. Guzmán, 21ML09(1) (UPOS). Carex borbonica × boryana . FRANCE. Reunion, 6-2-1856, Boivin, 997 (K). Reunion, Cirque de Cilaos, Forest-trail to Piton des Neiges, 2260 m, F.I. van Nek, 2203 (WAG 112766). Reunion, W cirque Malfate, Le Maido, 2200 m, 6-1-2009, M. Luceño & M. Guzmán, 20ML09 (UPOS 3724 to 3729). 3. Carex elgonensis Nelmes in Bull. Misc. Inform. Kew 1938: 245-246. 1938. Lectotype (designated in Haines & Lye, 1983): G. Taylor 3474 (BM 898027!); isotype: K 363578 photo! Rhizome caespitose, with short internodes. Stems 23-107 cm, ± stout, trigonous, smooth, green to yellowish-green. Leaves 8-21 cm × 3-6 mm, keeled to plicate, some rigid, slightly scabrid on the edges and apical parts; sheaths straw coloured to dark purplish to blackish; ligule 2.5-4 mm, obtuse or acute; anteligule edge concave to convex. Inflorescence 8.5-27.5 cm, lowest internode 3-11.5 cm and second one 1.5-8 cm. Lowest bract 6.5-20 cm × 2-5 mm, shorter than or as long as the inflorescence; sheath 1-7.5 cm long, the inner side purplish-red. Spikes 5-7, heteromorphic, 1-3 male spikes at the upper part and 3-6 female lateral spikes, frequently with a few male flowers at the top; male spikes 1-4 cm × 4-8 mm, linear to terete, the terminal one sessile or with a peduncle up to 2.3 cm; lateral female or androgynous spikes, 2.5-5.5 cm × 7-10 mm, arising singly (exceptionally in pairs), subdenseflowered, oblong-terete to ovoid-terete, peduncle up Taxonomic revision Carex tropical African group 231 Anales del Jardín Botánico de Madrid 68(2): 225-247, julio-diciembre 2011. ISSN: 0211-1322. doi: 10.3989/ajbm: 2256 Fig. 3. Distribution of Carex elgonensis: R, Rwanda; B, Burundi; S, Somalia. to 5 cm. Male glumes 5.5-9.2 × 1.2-1.8 mm, oblong, dark purplish to blackish without midrib or with a very narrow and straw coloured midrib, acuminate or aristate, acumen or arista up to 1.6 mm. Female glumes 4.5-6.5 × 0.9-1.9 mm, oblong-lanceolate, dark purplish to blackish, without midrib or with a very narrow and straw coloured midrib, acuminate or aristate, acumen or arista up to 1 mm. Utricles 4-6.2 × 1-1.5 mm, ellipsoid to obovoid-trigonous, straight, plu rinerved, gradu ally to abruptly narrowed into a beak, straw coloured in the lower portion and dark purplish to blackish in upper parts, sometimes entirely dark purplish or blackish; beak 0.8-1.9 mm, bifid, with ± similar dorsal and ventral sinus 0.2-0.5 mm depth, smooth or with disperse and short pricklets. Achenes 2-2.3 × 0.9-1.1 mm, trigonous, ellipsoid to obovoid. 2n =? Sept.-Feb. Banks of streams and borders of peat bogs; 3290-3810 m; Kenya and Uganda (Mt. Elgon). Comments: Nelmes (1938) described C. elgonensis from Kenyan Mount Elgon. Nevertheless, other authors, such as Haines & Lye (1983), considered C. mildbraedinana var. friesiorum Kük. to be a synonym of C. elgonensis. Therefore, they considered C. elgonensis as distributed in all Kenyan uplands. Following Nelmes (1938), we have considered C. elgonensis as endemic to Mount Elgon, and in contrast to Haines & Lye (1983), we think that C. mildbraedinana var. friesiorumshould be treated as a subspecies of C. mannii (see below). Selected specimens KENYA. Mt. Elgon, eastern slope above Japata estate, Ericacious belt, 3200 m, 9-5-1948, O. Hedberg, 854 (UPS, K). Trans 2256 carex_Maquetación 1 28/11/2011 11:46 Página 231 Nzoia, Mt. Elgon, c. 9 miles SW of the Suam Saw Mills, 3810 m, 22-12-1967, O.M. Mwangangi & F. Kariuki, 373 (K, LISC). Nyanza, Trans-Nzoia, Mount Elgon, 3290 m, 22-2-1935, G. Taylor, 3474 (BM 898027). Mt. Elgon National Park, Kambi, Mtamayua, 3561 m, 22-7-2007, M.L. Buide & al., 31UPO-K (UPOS 3273). UGANDA. Mt. Elgon, W slope above Butadiri, along the track via Mudangi through the Caldera, Hagenia Hypericum zone, on moist ground near “Mountain Hut”, 3200 m, 4-12-1967, O. Hedberg, 4555 (UPS). 4. Carex fischeri K. Schum. in Engl. (ed.), Pflanzenw. Ost-Afrikas, C: 130. 1895. Ind. loc.: “Abori” [Kenya]. Lectotype (designated in Gehrke, 2011): Fischer 640 (B 100240038!); isotype: K 363588 photo! = Carex simensis var. mauensis Kük. in Engl. (ed.), Pflanzenr. IV, 20(38): 654. 1909. Holotype: s.c. (B 100240026!). = Carex fischeri var. basiandra Kük. in Notizbl. Bot. Gart. Berlin-Dahlem 9: 314. 1925. Ind. loc.: “Mount Elgon: Bambus-Zone, 11000 ft”. Lectotype (designated Gehrke, 2011): R.E. Fries & T.C.E. Fries 1462 (UPS); isotypes: B 100240034!, K; paratypes: R.E. Fries & T.C.E. Fries 659 (UPS), R.A. Dummer 3465 (B 100240036!). Rhizome caespitose, with short internodes. Stems (30)40-90(115) cm, ± stout, trigonous, smooth, green to yellowish-green. Leaves (15)24-50(65) cm × 3- 8.5(10) mm, keeled to plicate, ± rigid, ± scabrid; sheaths dark brown, reddish-brown or purplish-red; ligule 1-8(12) mm, obtuse or acute; anteligule edge concave to convex. Inflorescence (11.5)13-37(54) cm, lowest internode 4-14(20) cm and second one (1)2.2- 10(13) cm. Lowest bract (8)15-41(52) cm × 2.5-7(8) mm wide, longer or shorter than the inflorescence; sheath 2.5-7.5(10) cm long, the inner side dark brown, reddish-brown or purplish-red. Spikes 4-10, heteromorphic, 1 male or gynecandrous terminal spike, exceptionally 2 male or gynecandrous spikes at the top, and 3-9 female lateral spikes; male or gynecandrous spikes (2)2.5-5.5(6) cm × 2-6.5(8) mm, linear to terete or clavate (when gynecandrous), sessile or with a peduncle up to 1.5(3.5) cm; female spikes, (1)2-6.5(8) cm × 4-9(10) mm, arising singly, lax- to dense-flow- ered, largely terete, exceptionally bearing a few short branches at the base of largest spikes, peduncle up to 5(8) cm. Male glumes (4)4.5-7.5(8.5) × (0.8) 1-2 mm, oblong-ellipsoid to oblong-obovoid, membranaceous, brown, reddish-brown or purplish with straw or green coloured midrib, acuminate, mucronate or aristate, acumen, mucro or arista up to 2(2.3) mm, exceptionally obtuse. Female glumes (2.8)3-5.5(6.5) × M. Escudero & M. Luceño232 Anales del Jardín Botánico de Madrid 68(2): 225-247, julio-diciembre 2011. ISSN: 0211-1322. doi: 10.3989/ajbm: 2256 (0.8)1-2(2.2) mm, ovate-elliptic, hyaline, brown, reddish-brown or purplish with straw or green coloured midrib, acuminate or aristate, acumen or arista up to 1.7(2.2) mm. Utricles (3)3.3-5.5(6) × 1.1-1.7(2) mm, ellipsoid, straight, pluri ner ved, taper ed to beak, brown or greenish-brown with reddish or purplish speckles; beak (0.8)1.1-1.7(2) mm, ± deep ly bifid, ± similar dorsal and ventral sinus 0.2-0.5 mm depth, smooth. Achenes 1.9-2.6 × 0.9-1.3 mm, trigonous, ellipsoid or ellipsoid-obovoid. 2n = 68, 70, 72. Jan.-Dec. Wet and open or shady places (forests, meadows, swamps, peat bogs, stream banks); 2150- 4300 m; D.R. Congo, Ethiopia, Kenya, Rwanda, Sudan, Tanzania, and Uganda. Comments: In view of the variability of the studied materials, we have considered two subspecies. However, we have seen several specimens that exhibit intermediate morphological features, but most of these materials can be classified in one of the two races. It is also interesting to note that some individuals studied show intermediate characters between this species and C. mannii (e.g. spikes arising in pairs and aculeolate beaks and red basal sheaths). Some additional studies may be necessary to know if these forms are the result of sporadic hybridization with the latter species or a third race of this highly polymorphic species. Fig. 4. Distribution of Carex fischeri subsp. fischeri: R, Rwanda; B, Burundi; S, Somalia. 2256 carex_Maquetación 1 28/11/2011 11:46 Página 232 We would like to emphasise that although the type materials of Carex fischeri var. basiandra: R.E. Fries & T.C.E. Fries 2210 (K 363579 photo!) and R.E. Fries & T.C.E. Fries 1462 (K 363580 photo!) each consist of only a few utricles, we are confident of their inclusion in this species. KEY TO THE SUBSPECIES OF CAREX FISCHERI 1. Female glumes hyaline, pale brown or light reddish-brown, never purplish; leaves non-rigid ........................................... ............................................. 4a. C. fischeri subsp. fischeri 1. Female glumes, dark reddish-brown or purplish; leaves rigid .................................. 4b. C. fischeri subsp. recedens 4a. Carex fischeri subsp. fischeri Stems (30)40-90(115) cm long, ± stout. Leaves up to 8.5(10) mm wide, slightly scabrid on the edges and apical parts; basal sheaths dark brown, reddishbrown or purplish-red. Inflorescence (11.5)17-37(54) cm, lowest internode 4-14(16) cm and second one (1)3-9.5(10.5) cm. Lowest bract 2.5-7 mm wide; sheath (2.5)4-7.5(10) cm long, the inner side dark brown, reddish-brown or purplish-red. Spikes 5-10; female spikes, 4-7(8) mm wide, lax-flowered to denseflowered, terete, peduncle up to 5(8) cm. Female glumes (2.8)3-4.8(5.5) × (0.8)1-1.6(1.8) mm, hyaline, pale brown to reddish-brown. Utricles (3)3.3-4.8(5) long, light brown to greenish-brown with reddish or purplish speckles. 2n = 68, 70, 72. Jan.-Dec. Wet and open places; 2350-3450 m; Kenya, Sudan, Tanzania, and Uganda. Comments: Plants collected in Tanzania are usually bigger in all their parts. Selected specimens ETHIOPIA. Mt. Uociacia, about 15 Km W of Addis Ababa, 2500 m, 27-10-1965, W.J.J.O. de Wilde & B.E.E. de Wilde-Duyfjes, 8542 (WAG 112501, WAG 112502). Sidamo, 82 Km S of Agere Selam along road to Kebre Mengist, 2150 m, J.J.F.E. De Wilde & M.G. Gilbert, 317 (WAG 112496, WAG112497). Addis Abeba, about 2 Km NE of Italian Embassy, 2400 m, 31-7-1965, W.J.J.O. de Wilde & B.E.E. de Wilde-Duyfjes, 7409 (WAG 112505, WAG 112504, WAG 112505). About 25 Km W of Addis Ababa, 2600 m, 24-4-1965, W.J.J.O. de Wilde & B.E.E. de Wilde-Duyfjes, 6423 (WAG 112506, WAG 112507). KENYA. Nyanza, Londiani, Tinderet Forest Reserve, Camp 6, about 6 Km SSE of Timboros Station, 2800 m, R.A. Maas Geesteranus, 5475 (MO, PRE, L 625310). Rift Valley, Nakuru, eastern Mau Forest Reserve, camp 10, 2650 m, 30-8-1949, R.A. Maas Geesteranus, 5966 (MO 1731309, PRE, L 625311, WAG 112520, US 2247255). Mt. Kenya occid., 2300 m, 29-12-1921, R.E. Fries & T.C.E. Fries, 493 (BR, B 100240023). Mt. Aberdare, 2300 m, 12-3- 1922, R.E. Fries & T.C.E. Fries, 2210 (B 100240035). Nyanza, Trans-Nzoia, Mount Elgon, 2400 m, 19-2-1935, G. Taylor, 3424 (MO 2128965). Mt. Kenya, Westhang, bei Met.- Station, 3050 m, 25-9-1980, H. Rehder (M 124770, M 124772). Nyeri, Mount Kenya Taxonomic revision Carex tropical African group National Park, area near Percival’s Bridge along the park road E of Naro Moru, 2740 m, 5-3-1974, G. Davidse, 7031 (WAG 122522, WAG 122521). Bamboo zone, western slopes of Mount Kenya, along the trail from West Kenya Forest Station to summit, British East Africa, 3000 m, 28-9 to 7-10-1909, E.A. Mearns, 1765 (US 631719, C). Bamboo zone, western slopes of Mount Kenya, along the trail from West Kenya Forest Station to summit, British East Africa, 3000 m, 28-9 to 7-10-1909, E.A. Mearns, 1674 (US 631625). Mau – Plateau, 3000 m, 8-5-1906 (B 100240026). West Kenya, Liki- River, in moist Stelle am Bache, 2500 m, 11-2-1922, R.E. Fries & T.C.E. Fries, 1462 (B 100240034). Bamboo zone, western slopes of Mount Kenya, along the trail from West Kenya Forest Station to summit, British East Africa, 3000 m, 28-9 to 7-10-1909, E.A. Mearns, 1736 (C). Central, Nanyuki, Mt. Kenya, W slope, National Park Road (Naro Moru Track), 3450 m, 23-1-1970, L. Ryvarden, 5576 (O). Aberdares National Park, Mutubio road, 3106 m, 26-7- 2007, M.L. Buide & al., 83UPO-K (UPOS 3289). Mt. Kenya National Park, Naro Moru route, 2974 m, 28-7-2007, M.L. Buide & al., 104UPO-K (UPOS 3300). SUDAN. Equatoria, Torit, Mt. Dumuso, 1-6-1950, J.K. Jackson, 1550 (K). TANZANIA. Arusha, Arusha National Park, the crater Mt. Meru, E. side, 2600 m, 23-4-1968, P.J. Greenway & Kanuri, 13502 (MO 1991585). Arusha, Arumeru, Mt. Meru, 2820 m, 11-1-1985, R.E. Gereau, 1667 (MO 3204851, MO 5106243). Tanganyika Territory, Mt. Meru, eastern slopes above Olkakola estate, 3150 m, 31-10-1948, O. Hedberg, 2420 (UPS). Mt. Meru, Crater, montane meadow inside the crater, 2700 m, 23-1-1976, K. Szczepanek & B. Zemanek, 820 (UPS). Kilimanjaro, Morlow above rain gauge, 2800 m, 4-8-1993, JMG, 93521 (K). Kilimanjaro, forest above Kilimanjaro timbers, 2500 m, 13-1-1994, JMG, 9464 (K). Tanganjika, Kilimandscharo, Güterwald, 2360 m, 4-1-1934, H.J. Schlieben, 4487 (M 124774, B 100240046, B 100240048). Kilimandscharo, Urwaldzone zwischen Marangu und der Bismarckhütte, 2800 m, 12-1929, Fr.u.R. Wettstein (M 124773, WU 3029). Kilimandscharo, 3000 m, 2-10-1909, R. Endlich, 626 (B 100240037). UGANDA. Western, Toro, Rwenzori, Namwamba Valley, 2750 m, 6-6-1935, G. Taylor, 2944 (MO 2128967). Ruwenzori, Bujuku Valley, near Bigo camp, along a small path close to the camp site, 3400 m, 22-3-1948, O. Hedberg, 397 (UPS). Kigezi, Mt. Muhavura, W slopes, 3350 m, 7-10-1948, O. Hedberg, 2179 (UPS). Mt. Elgon, eastern slope above Japata estate, 3000 m, 1-3-1948, O. Hedberg, 193 (UPS). Kigezi, Mt. Muhavura, W slopes, 3350 m, 7- 10-1948, O. Hedberg, 2179 (UPS). Mt. Elgon, bamboo forest along the bank of the Sasa River, close to the Bumagaluba - Mountain Hut Trail, 2500 m, 22-2-1974, S. Lisowski, 10936 (BR). Toro U2, Mt. Ruwenzori, near Nyamileju (2 nd) Hut, 3300 m, 30-12-1968, K.A. Lye, 1259 (K). Mt. Ruwenzori, 2870 m, 8-1938, J.W. Purseglove, 325a (K). Mt. Elgon, 3350 m, 1-1918, R.A. Dummer, 3465 (US 1172896, B 100240036). 4b. Carex fischeri subsp. recedens (Kük.) Luceño & M. Escudero, comb. nov. ≡Carex longipedunculata K. Schum. var. nina gon - gensis Kük. f. recedens Kük. in Mildbr. (ed.), Wiss. Erg. Deut. Zentr.-Afr. Exped., Bot.: 53. 1914, basyonimus. Ind. loc: “Vulkan – Gebiet: Karisimbi, Güterwald von Hagenia über dam Bambus”. Lectotype (designated in Gehrke, 2011): Mildbraed 1578 (B 100240021!); isotype: K. 233 Anales del Jardín Botánico de Madrid 68(2): 225-247, julio-diciembre 2011. ISSN: 0211-1322. doi: 10.3989/ajbm: 2256 2256 carex_Maquetación 1 28/11/2011 11:46 Página 233 Selected specimens TANZANIA. Kyimbila, North of Lake Nyasa, 1915, A. Stolz, 2284 (UPS, BR, PRE, C, HUH 246049, H 1559427). Kitulo Plateau, just left of Igoma-Kitulo road, 1 Km beyond Kikondo, 2750 m, 31-12-1969, RW, 530 (K). Kitulo Project, along Makengalima River, 25-11-1967, J. Prins-Lambert, 317 (WAG 112511, WAG 112512). 9. Carex petitiana A. Rich., Tent. Fl. Abyss. 2: 513. 1850. Inc. loc.: “Crescit in montosis provinciae Ouodgerate (Ant. Petit)” [Ethiopia]. Lectotype (designated by Haines & Lye, 1983): Petit s.n. (P 466086, photo!). = Carex longipedunculata K. Schum. in Engl., Pflanzenw. Ost-Afrikas, C: 130. 1895. Ind. loc.: “(Nordseite des Kl. [Kilimanjaro] …)”. “Sumpfige Senke bei der Höhle Noholu, 3200 m”. Lectotype (designated in Gehrke, 2011): Volkens 2015 (B); isotype: K363582 photo! K 363591 photo! = Carex longipedunculata subsp. cuprea Kük. in Notizbl. Bot. Gart. Berlin-Dahlem 9: 315-316. 1925. Ind. loc.: “West-Kenia, Forest Station in Sumpf, 2300 m”. Lectotype (designated in Gehrke, 2011): R.E. Fries & T.C.E. Fries 734 (UPS); isotype: BR 8638984! K; pa ra types: R.E. Fries & T.C.E. Fries 405 (UPS!, K 363585 photo!), R.E. Fries & T.C.E. Fries 659 (S 05- 9680 photo!, UPS!), R.E. Fries & T.C.E. Fries 943 (UPS!), R.E. Fries & T.C.E. Fries 2763 (UPS!). Carex cuprea (Kük.) Nelmes in Bull. Misc. Inform. Kew 1938: 247. 1938. = Carex preussii K. Schum. in Bot. Jahrb. Syst. 24: 340. 1897. Ind. Loc.: “Mt. Cameroon, near Mann’s spring, 2600 m”. Lectotype (designated in Gehrke, 2011): Preuss 727 (B). = Carex preussii var. camerunensis Nelmes in Bull. Misc. Inform. Kew 1938: 247. 1938. Ind. loc.: “British Cameroons, Cameroons Mountain, 2130-3000 m”. Lectotype (designated in Gehrke, 2011; precised here): Mann 2099 (K); isotypes: HUH 246054! K 363492 photo!, K 3634923 photo!, K 363494 photo! = Carex aethiopica var. stolonifera Boeck, Linnaea 41: 286. 1877. Ind. loc.: “Abyssinia, Gaffat (alt. 8000 ped.)”. Lectotype (designated in Gehrke, 2011): Schimper 1298 (K); isotypes: EA, K 363462 photo, M 105734 M. Escudero & M. Luceño photo! US 945614!, Z + ZT. Carex simensis var. stolonifera (Boeck.) Kük. in Engl. (ed.), Pflanzenr. IV, 20(38): 654. 1909. Rhizome caespitose, with short, or very exceptiona - lly long, internodes. Stems (27.5)40-133 cm, ± stout, trigonous, smooth, green to yellowish-green. Leaves (12)15-60 cm × (4)4.8-10 mm, keeled to plicate, rigid, slightly scabrid on the edges and apical parts; sheaths brown to reddish-brown; ligule 1.5-10.5 mm, obtuse or acute; anteligule edge concave to convex. Inflorescence (7)11-43 cm, lowest internode (3)5-18.5 cm and second one (0.2)2-12 cm. Lowest bract 5.5-35 cm × 2-7.5 mm, shorter or longer than the inflorescence; sheath 1.8-7(8) cm long, the inner side brown, reddishbrown or purplish. Spikes 3-9, subhomomorphic to subheteromorphic (rarely heteromorphic), generally all spikes gynecandrous, with the male part increasingly longer as going upwards in the stem, sometimes lowest spikes entirely female (more rarely 1 male terminal and the rest female); terminal spike 3.2-6.5 cm × (5)7- 12 mm, clavate (rarely terete), sessile or with a peduncle up to 3.3 cm; lateral spikes (1)2-6 cm × 7-12 mm, dense-flowered, clavate to widely terete or ellipsoid, solitary or in pairs, exceptionally with a few short branches at the base of the largest spikes, peduncle up to 7.5(12.5) cm. Male glumes (4.5)5.4-7.5(9) × (1)1.1- 2.2(2.4) mm, oblong-elliptic to oblong-obovate, dark brown to dark reddish-brown with straw or green coloured midrib, mucronate or acuminate, mucro or acumen up to 1.5 mm, rarely obtuse. Female glumes 3.4-6(7) × (1.1)1.3-2.6 mm, widely oblong-elliptic, dark brown, reddish-brown or purplish-red, with wide and green or straw coloured midrib, mucro or acumen up to 1(1.5) mm, rarely, obtuse. Utricles (3.7)4-6(6.5) × 1.2-2.2(2.5) mm, ellipsoid-trigonous, straight, tetra-nerved to plurinerved, tapered to beak, greyish-green with reddish or purplish speckles; beak 1.1-1.6(1.8) mm, deeply bifid, with ± similar dorsal and ventral sinus, 0.4-0.6 mm depth, aculeolate (very rarely smooth). Achenes 2.5-3(3.2) × 1.1-1.6(1.9) mm, trigonous, oblong-elliptic to oblong-obovoid. 2n = 70. Jan.-Dec. Open (sometimes shady) and wet places: swamps, wet meadows, streams and bogs; 1800-3600 m; Cameroon, D.R. Congo, Ethiopia, Kenya, Malawi, Nigeria, Tanzania, Uganda and Zimbabwe. Comments: Carex preussii K. Schum. was described from Cameroon and accepted by some authors, such as Hooper & Napper (1972) and Gehrke (2011). Nevertheless, we have not found any difference between materials from western tropical Africa and the remaining typical C. petitiana materials from eastern tropical Africa. 240 Anales del Jardín Botánico de Madrid 68(2): 225-247, julio-diciembre 2011. ISSN: 0211-1322. doi: 10.3989/ajbm: 2256 2256 carex_Maquetación 1 28/11/2011 11:46 Página 240 The putative isotype material of Carex longipedunculata subsp. cuprea Kük. “West-Aberdare: An der Grenze zwischen Bergwald und Bambusregion”, R.E. Fries & T.C.E. Fries 2763 (K 363586 photo!) consists of just a few utricles. KEY TO THE SUBSPECIES OF CAREX PETITIANA 1. Spikes all solitary in the nodes; female glumes brown to reddish-brown .................... 9a. C. petitiana subsp. petitiana 1. At least some spikes arising in pairs; female glumes reddishbrown to purplish-red ... 9b. C. petitiana subsp. attenuata 9a. Carex petitiana subsp. petitiana Stems (27.5)40-95(120) cm, ± stout. Leaves (12)15- 35(42) cm × (4)4.8-8(9) mm. Inflorescence (7)11- 33(37) cm, lowest internode (3)5-15 cm and second one (0.2)2-7.3(9.5) cm. Lowest bract 5.5-23 cm × 2- 4(7.5) mm; sheath (1.8)2.5-7(8) cm long, the inner side brown to reddish-brown. Spikes 3-8, all solitary. Female glumes (3.5)4-6(7) × 1.6-2.6 mm, dark brown to reddish-brown. 2n = 70. Jan.-Feb. Open (sometimes shady) and wet places: swamps, wet meadows, streams and bogs; 2000-3600 m; Cameroon, D.R. Congo, Ethiopia, Kenya, Nigeria, Tanzania and Uganda. Selected specimens CAMEROON. Northwest, near Verkovi Village, lower slopes of Mt. Oku, 2600 m, 4-1986, D.W. Thomas & H.L. Mcleod, 6045 (MO 3442595). Bambouto Mountains, ca. 5 Km NW. of the village Biété, 2300 m, 9-5-1964, W.J.J.O. de Wilde & B.E.E. de Wilde- Duyfjes, 2541 (MO 2171215, B 100240029). Southern Cameroons, Bamenda, 2000 m, 17-2-1958, F.N. Hepper & J.R. Charter, 2042 (K, WAG 112529). Bafut-Ngemba Forest Reserve, 2150 m, 20-2- 1958, F.N. Hepper, 2085A (K). Southern Cameroons, Bamenda, 2150 m, 20-2-1958, F.N. Hepper, 2085 (K). Mezam, Bambili Lakes, 2230 m, 16-5-1970, P.J. Bauer, 68 (K). Following from ridge above Bamenda and Bafutngem F.R., 2320 m, 8-7-1962, M. Brunt, 777 (K). Bambouto Mountains, c. 5 Km NW of the village of Biété, 2300 m, 9-5-1964, G.J.H. Amshoff, 1965 (U227743). Est, Kumbo, Mt. Okou, 2700 m, 1-2-1970, C.N.A.D., 1814 (WAG 112528). Kamerunberg, Gras region, 2500 m, 8-5-1938, Domke, 308 (B 100240033). Buea, Manns-Quelle, Abhang, 4-2-1891, Preufs (B 100240031). Bamenda, Mt. Mba, Kokeka, 2130 m, 21-3-1935, D.E. Coombe, 229 (B 100240032). CONGO. Parc National Albert (Virunga National Park), Shaheru, 2700 m, 1-1938, J. Lebrun, 9332 (BR, NY, BM 898009, B 100240039). Parc Natinal Albert (Virunga National Park), Versant E du Ninagongo, cratere du Shaheru, 2100 m, 11-1937, J. Lebrun, 8780 (BR, US 2091181). Kivi Nord, Talsohle von Ngungu, 18 Km SW Sake, 2300 m, 12-8-1954, H.U. Stauffer, 69 (WAG 225343). ETHIOPIA.About 5 Km NE of Addis Abeba, 2400 m, 26-4- 1965, W.J.J.O. de Wilde & B.E.E. de Wilde-Duyfjes, 6431 (WAG 112498, WAG 112499, WAG 112500). Arussi, Chilalo Awraja, Gobe Livestock farm, c. 10 Km N of Koffale, 2700 m, 12-10-1971, M. Thulin, 1466 (UPS). About 10 Km SE of Hagere Selam, SE of Wondo, 3000 m, 13-3-1966, WJ.J.O. de Wilde & B.E.E. de Wilde- Taxonomic revision Carex tropical African group Duyfjes, 10307 (WAG 112493). Schimper, 1298 (US 945614, BM 898016). KENYA. Narok, K. 6, Enesambulia Valley N.E., 2700 m, 29-6- 1969, P.J. Greenway & Kanuri, 13666 (MO 1991712). Mau Range, c. 10 Km N of Timboroa, 2700 m, 2-6-1948, O. Hedberg, 1089 (UPS, K). Mt. Kenya occid., prope Forest Station, 2300 m, 27-1- 1921, R.E. Fries & T.C.E. Fries, 405 (UPS). Mt. Aberdare occid., 3- 4-1922, R.E. Fries & T.C.E. Fries, 2763 (UPS). Mt. Kenya occid., 2300 m, 1-1-1922, R.E. Fries & T.C.E. Fries, 659 (UPS). Mt. Kenya occid., Coles Farm, 2000 m, 16-1-1922, R.E. Fries & T.C.E. Fries, 943 (UPS). Narok, Mbalagai (Masai), Doboti about 9 miles from Cobb’s Gate near the Mau Forest Reserve on the track to Nairagie Ngare, 2900 m, 22-5-1961, Clover & al., 1486 (BR, K). Navaisha, South Kinangop, 2750 m, 3-6-1961, Polhill, 429 (BR, K, NY). Thomson’s Falls to Nakuru, 20 miles from Th. Falls, 2750 m, 14-8- 1952, A.V. Bogdan, 3525 (K). Near Njoro, 2380 m, 9-8-1962, A. Bogdan, 552 (K). NE of Kijabe, Aberdare Forest Reserve, 2500 m, 16-10-1949, A. Bogdan, 2655 (K). 5 miles NW of Timboroa, 2750 m, 17-5-1959, A. Bogdan, 4826 (K). Mdoto Mts., Siruan, 2440 m, 1-1-1959, J.G.B. Newbould, 3373 (K). Trans-Nzoia, Suam Sawmills road, 2200 m, 21-5-1969, D.M. Napper & Tweedie, 2133 (K, WAG 112510, WAG 112509). Rift Valley, Nakuru, eastern Mau Forest Reserve, camp 10, Songei, 2650 m, 1-9-1949, R.A. Maas Geesteranus, 6015 (WAG 112526). Vicinity of Lake Naivasha, British East Africa, 1950 m, 17-7 to 15-8-1909, E.A. Mearns, 899 (US 630929, BM 898014). Bamboo zone, western slopes of Mount Kenya, along the trail from West Kenya Forest Station to summit, British East Africa, 3000 m, 28-9 to 7-10-1909, E.A. Mearns, 2359 (US 632331). Mt. Kenya National Park, Naro Moru route, 2270 m, 28-7-2007, M.L. Buide & al., 123UPO-K (UPOS 3212). Mt. Elgon National Park, 2788 m, 23-7-2007, M.L. Buide & al., 56UPO-K (UPOS 3278). Mt. Elgon National Park, Kambi Mtamayua, 3450 m, 22-7-2007, M.L. Buide & al., 24UPO-K (UPOS 3270). Aberdares National Park, Mutubio West Gate, 3212 m, 27-7-2007, M.L. Buide & al., 100UPO-K (UPOS 3296). Mt. Elgon National Park, 2768 m, 23-7-2007, M.L. Buide & al., 53UPO-K (UPOS3276). Mt. Elgon National Park, Kambi Mtamayua, 3450 m, 22-7-2007, M.L. Buide & al., 24UPO-Kbis (UPOS 3270bis). Mt. Elgon National Park, Kambi Mtamayua, 3561 m, 22- 7-2007, M.L. Buide & al., 29UPO-K (UPOS 3271). Mt. Elgon National Park, Kambi Mtamayua, 3561 m, 22-7-2007, M.L. Buide & al., 30UPO-K (UPOS 3272). NIGERIA. NE State Nigeria, Sardauna, Gangirwal, 2290 m, 24-11-1974, J. Chapman, 3576 (OXF 142981). 241 Anales del Jardín Botánico de Madrid 68(2): 225-247, julio-diciembre 2011. ISSN: 0211-1322. doi: 10.3989/ajbm: 2256 Fig. 12. Distribution of Carex petitiana subsp. petitiana: CR, Central African Replublic; U, Uganda; C, Cameroon; S, Somalia. 2256 carex_Maquetación 1 28/11/2011 11:46 Página 241 UGANDA. Kigezi, 13-2-1945, Greenway & Kanuri, 7121 (BR). Virunga Mountains, Mgainga-Sabinio, 2440 m, 21-11-1934, G. Taylor, 1897 (BM 898026). Ruwenzori, 2440 m, 1893-94, G.F. Scott Elliot, 7578 (BM 898017). 9b. Carex petitiana subsp. attenuata (Kük.) Luceño & M. Escudero, comb. nov. ≡ Carex longipedunculata var. attenuata Kük. in Bot. Not. (Lund) 1934: 82. 1934, basyonimus. Ind. loc.: “Inyanga: prope pagum Inyanga ad rivulum kuhera, c. 1800 m.s.m.” [Malawi]. Lectotype (designated here): T.C.E. Fries & al. 3094 (LD 60956!); isotype: T.C.E. Fries & al. 3094 (S photo!). Stems 50-133 cm, very stout. Leaves 20-60 cm × 8-10 mm. Inflorescence 21-43 cm, lowest internode 7-18.5 cm and second one 4.5-12 cm. Lowest bract 16-35 cm × 3-7 mm, shorter than the inflorescence; sheath 1.8-5 cm long, the inner side reddish-brown to purplish. Spikes 5-9, arising in pairs in at least one node, the rest singly. Female glumes 3.4-5.5 × (1.1)1.3-1.9(2) mm, dark reddish-brown to purplish-red. 2n = 70. Jan.-Dec. Wet and open places: swamps, wet meadows and bogs; 1800-2300 m; Malawi (Nyika) and Zimbabwe (Inyanga). Comments: All specimens studied from Zimbabwe are well differentiated from those of the typical subspecies, but some individuals from Malawi display some intermediate features between the subspecies. Selected specimens MALAWI. Northern, Nyika Plateau, 2300 m, 15-12-1975, E. M. Escudero & M. Luceño Phillips, 530 (MO 2829617). Northern, Nyika Plateau, 2300 m, 7-11-1975, E. Phillips, 145 (MO 2449940). Nyasaland, Nyika Plateau, Lake Kaulime, 4-1-1959, E.A. Robinson, 3028 (M 124768, M 124769). Northern, Nyika Plateau, 2290 m, 6-11-1975, E. Phillips, 118 (WAG 112780, WAG 112779). Northern, Nyika Plateau, 2290 m, 22-11-1975, E. Phillips, 396 (WAG 112782, WAG 112781). Mare Dam in Nyika National Park, 2000 m, 1-1- 1991, S. Laegaard, 16024 (US 3261097, B 100014065). ZIMBABWE. Inyanga, Mare Dam, Inyanga National Park, 6- 1-1972, G.E. Gibbs Russell, 1208 (MO 2653179). Rhodesia austr., Inyanga, ad pedes montis Inyangani, 2000 m, 6-7-1930, T.C.E. Fries & al., 3504 (UPS). Rhodesia austr., Inyanga, ad pedes montis Inyangani, 2000 m, 6-7-1930, T.C.E. Fries & al., 3504 (BR, BM 896204, B 100240030, LD 600955, LD 600956). 10. Carex simensis Hochst. ex A. Rich. In Tent. Fl. Abyss. 2: 514. 1850. Ind. loc.: “Crescit in pratis humidis montis Bouhait, ad fines et supra fines arborum et fruticum, in provincial Semiène, manse Junio (Schimper)” [Ethiopia]. Lectotype (designated in Haines and Lye; precised here): Schimper 1180 (P 466087 photo!); isotypes: P 466088 photo!, P 466089 photo!, MO 3255498!, MO 2128070!, L 625307!, L 625305!, US 945543!, M 105740 photo!, STU 347 photo!, STU 348 photo!, BR 8249845 photo!, WAG 3721 photo! = Carex simensis var. lanuriensis De Wild., Pl. Bequaert. 4: 247. 1927. Ind. loc.: “Ruwenzori, vallée du Lanuri 4000 et 4300 m. et aux environs de 3000 m. d’altitude”. Lectotype (designated in Gehrke, 2011): Bequaert 4520 (BR 8644619 photo!); isotypes: OXF!, PRE, K 363488 photo!, BR 8644626 photo!, S; paratypes: Bequaert 4680 (BR 864461!), BR!, K 363487 photo!, PRE 0594525-0 photo! = Carex karisimbiensis Cherm. in Bull. Soc. Bot. France 82: 344. 1935. Ind. loc.: “volcan Karisimbi (NE. du lac Kivu), 3400-3900 [Congo]”. Lectotype (designated in Gehrke, 2011; precised here): H. Humbert 8557 (P 466090 photo!); isotypes: P 466091 photo!, P 466092 photo!, B 100166188 photo!, BR 8644596 photo!, BR 8644602 photo! = Carex simensis var. longistipitata Kük. in Notizbl. Bot. Gart. Berlin-Dahlem 9: 316. 1925. Ind. loc.: “West-Kenia: Obere Bambusregion im sumpfe, 2800 m”. Lectotype (designated in Gehrke, 2011): R.E. Fries & T.C.E. Fries 1300 (UPS); isotypes: BR 6265427 pho- to!, H 1559429!, S 05-9686 photo!; paratype: R.E. Fries & T.C.E. Fries 1300a (UPS). 242 Anales del Jardín Botánico de Madrid 68(2): 225-247, julio-diciembre 2011. ISSN: 0211-1322. doi: 10.3989/ajbm: 2256 Fig. 13. Distribution of Carex petitiana subsp. attenuata: M, Malawi. 2256 carex_Maquetación 1 28/11/2011 11:46 Página 242 = Carex simensis var. nemorum Chiov. in Ann. Bot. (Rome) 10(3): 407. 1912. Holotype: G. Negri (FT 669 photo!). Rhizome caespitose, with short internodes. Stems (13)22-79 cm, ± stout, trigonous, smooth, green to yellowish-green. Leaves 6.5-31 cm × 3-12 mm, keeled to plicate, very rigid, ± coriaceous, slightly scabrid on apical parts; basal sheaths dark purplish to blackish, coriaceous; ligule 3-23 mm, acute; anteligule edge concave to convex. Inflorescence 4-43.5 cm, lowest internode 0.5-15.5(23) cm and second one 0.5-8(12) cm. Lowest bract 6.5-20 cm × 2-5 mm, shorter than the inflorescence; sheath (0.5)1-7.5 cm long, the inner side deep purple to blackish. Spikes 5-12, heteromorphic, 0-2 male spikes at the top (sometimes the terminal one gynecandrous) and 3-11 female lateral spikes; terminal spike (1)2.5-4.8(7) cm × 2-7(10) mm, terete, sessile or with a peduncle up to 2.5 cm; female spikes, 1.5-6.5(8) cm × 5-10 mm, subdense-flowered, terete to narrowly ellipsoid, arising singly or in pairs, sessile or with a peduncle up to 8(10) cm. Male glumes (4)5-8(9) × (1.2)1.4-2.2(2.8) mm, oblong-elliptic to oblong-obo- vate, blackish with very wide and pale midrib, emarginate-mucronate to mucronate, mucro up to 1 mm, rarely obtuse. Female glumes (3)4-6(6.5) × (1.2)1.4- 2.1(2.5) mm, oblong-elliptic, deep purple to blackish, with very wide and pale midrib, acuminate or aristate, acumen or arista up to 1.6 mm. Utricles (3.5)4.5- 6.5(7.2) × (1.1)1.2-1.6(2) mm, narrowly ellipsoidtrigonous, straight, plurinerved, tapered to beak, greyish-brown to green; beak (0.7)1-1.6(2) mm, bifid, with ± similar dorsal and ventral sinus 0.2-0.6 mm depth, smooth. Achenes (2.2)2.3-2.9(3.2) × (0.9)1-1.2(1.3) mm, trigonous, oblong-ellipsoid. 2n =? Jan.-Dec. Open (sometimes shady) and wet places: wet meadows, streams and swamps; 2800-4100 m; D.R. Congo, Ethiopia, Kenya and Uganda. Comments: Some populations, mainly from Ethio - pia but also from Kenya, are formed by individuals that are significantly smaller in all their parts. The names C. simensis var. lanuriensis and C. kari - simbiensis were created to accommodate plants exhibiting some atypical characters but, in our opinion, all such forms should be included within the variability of C. simensis. Selected specimens CONGO. Kivu, Parc des Virunga, Karisimbi, plateu de Rukumi, 3500 m, 22-1-1972, P. van der Keken, 9125 (MO 3763955). Kivu, Parc des Virunga, Karisimbi, plateu de Rukumi, 3500 m, 22- 1-1972, P. van der Keken, 9126 (MO 3763955). Parc National des Virungas, between Mikeno & the Karisimbi volcanoes, 3200 m, 23- Taxonomic revision Carex tropical African group 1-1972, G. Troupin, 14301 (MO 2421974). Bale, Square 84D, between Shashamane and Goba, 3200 m, 28-5-1972, J. Ash, 1696 (K). ETHIOPIA. Montis Simensis Bachit ad fines et supra fines, 23- 6-1838, Schimper, 1180 (MO 3255498, MO 2128070, M 124766, L 625307, L 625305, US 945543, O). Mt. Boruluccu, along road to Ticcio, about 30 Km SE of Asella, 4000 m, 4-7-1965, W.J.J.O. de Wilde & B.E.E. de Wilde-Duyfjes, 9119 (MO 2265995, WAG 112492). Bale, 10-15 Km SE of Goba on road towards Masslo, 3400 m, 16-5-1980, M. Thulin & al., 3716 (UPS, K). Arussi, Kaka, E slope, 3750 m, 7-1980, O. Hedberg & M. Tadesse, 7047 (UPS). Arussi, Chillalo Awraja, Galama Mts. (c. 30 Km ESE of Aselle), about 3 Km E of Boraluco, 3750 m, 6-9-1967, O. Hedberg, 4181 (UPS, K). Arussi, Kaka, E slope, 3750 m, 7-1980, O. Hedberg & M. Tadesse, 7068 (UPS). Shewa, c. 25 Km from Debra Brhan to Akober (Gorebela) near microwave, 3600 m, 4-6-1982, S. Edwards & Class, 502/82 (UPS). Arussi, Chillalo Awraja, Galama Mts. (c. 30 Km ESE of Aselle), about 3 Km E of Boraluco, 3750 m, 8-9- 1967, O. Hedberg, 4224 (UPS). Bale, Bale Mountains National Park, on the crest S of Garba Goracha cap site, 4100 m, 2-11-1973, O. Hedberg, 5659 (UPS). Gojjam, Mts. Choké, vicinity of the upper Ghiedeb valley, 12-8-1957, I.M. Evans & C.L.A. Leakey, 482 (K). Bale, Dinshu, about 157 Km E of Shashamane, along road to Goba, 3300 m, 24-7-1970, J.J.F.E. De Wilde, 6787 (K). Guma, 10- 9-1863, Schimper, 1289 (M124765, US945608, E219439). About 25 Km of Asella, W-slope of Mt. Boroluccu, 3800 m, 11-9-1965, W.J.J.O. de Wilde & B.E.E. de Wilde-Duyfjes, 8124 (WAG 112494, WAG 112495). Harar, Gara Mullata Mt, 3290 m, 10-6-1962, W. Burger, 1902 (US 2480955). In Monte Simenensis Bachit, 22-2- 1852, Schimper, 1988 (GOET, WU 676). KENYA. Nyanza, Trans-Nzoia, Mount Elgon, 3400 m, 24-2- 1935, G. Taylor, 3677 (MO 1611134). Mt. Kenya, K2, Cheramgani Hills, Chepkotet, 3240 m, 13-8-1968, M. Thulin & A. Tigids, 243 (UPS). Aberdare Range, Kinangop, northern slope, Alpine belt, 3750 m, 13-7-1948, O. Hedberg, 1546 (UPS). Mt. Kenya, NW slopes, in the uppermost part of the Hagenia-Hypericum zone, 3400 m, 18-8-1948, O. Hedberg, 1936 (UPS). Mt. Elgon, in the crater (alpine region), near “Maji ya moto”, 3600 m, 21-5-1948, O. Hedberg, 1017 (UPS). Nanyuki, Mt. Kenya, W slope, National Park Road (Naro Moru Track), 3600 m, 10-1-1971, O. Hedberg, 4828 (UPS). Fort Hall, Mt. Aberdare, S slope above S Kinangop, 243 Anales del Jardín Botánico de Madrid 68(2): 225-247, julio-diciembre 2011. ISSN: 0211-1322. doi: 10.3989/ajbm: 2256 Fig. 14. Distribution of Carex simensis: R, Rwanda; B, Burundi. 2256 carex_Maquetación 1 28/11/2011 11:46 Página 243 along the path towards “The Elephant”, 3500 m, 20-9-1967, O. Hedberg, 4330 (UPS). Western, Mt. Elgon, on the ridge W of Suam River, 3560 m, 24-9-1966, S.W. Bie, 66266 (UPS). Bamboo zone, western slopes of Mount Kenya, along the trail from West Kenya Forest Station to summit, British East Africa, 3000 m, 28-9 to 7-10-1909, E.A. Mearns, 1677 (NY, US 631628). Bamboo zone, western slopes of Mount Kenya, along the trail from West Kenya Forest Station to summit, British East Africa, 3000 m, 28-9 to 7-10- 1909, E.A. Mearns, 1706 (NY, BM 898007, HUH 246050). Mt. Elgon, Kimothon Route, 3640 m, 15-3-1983, H. Rehder (M 124778). Bamboo zone, western slopes of Mount Kenya, along the trail from West Kenya Forest Station to summit, British East Africa, 3000 m, 28-9 to 7-10-1909, E.A. Mearns, 1742 (BM 898008, HUH 246051). Obere Bambusregion im Sumpfe, 2800 m, 30-1-1922, R.E. Fries & T.C.E. Fries, 1300 (H 1559429). Aberdares National Park, Mutubio West Gate, 3212 m, 27-7-2007, M.L. Buide & al., 101UPO-K (UPOS 3297). Mt. Kenya National Park, Naro Moru route, 3588 m, 28-7-2007, M.L. Buide & al., 116UPO-K (UPOS 3307). UGANDA.Mt. Elgon, at the Sebei path W of Swam river, 3530 m, 22-5-1948, O. Hedberg, 1043 (UPS). 11. Carex uluguruensis Luceño & M. Escudero, spec. nov. A simili Carex mildbraediana Kük. foliis angustioribus atque spicis lateralibus solitariis aut binis –nec ternis– differt. Ind. loc.: “Tanzania, Morogoro, Uluguru Mountains”. Holotype: H.J. Schlieben 3516 (WAG 112513!); isotypes: B 100240022!, PRE! Rhizome caespitose, with short internodes. Stems 50-90 cm, ± stout, trigonous, smooth, green to yellowish-green. Leaves 13-26 cm × 7-9 mm, keeled to plicate, very rigid and coriaceous, with revolute margins, slightly scabrid on the edges and apical parts; sheaths reddish-brown to purplish-red; ligule 4-6.5 mm, M. Escudero & M. Luceño acute; anteligule edge concave. Inflorescence 8.5-20.5 cm, lowest internode 4.5-11.5 cm and second one 2- 3.5 cm. Lowest bract 8.5-13.5 cm × 3-6 mm, shorter or longer than the inflorescence; sheath 4-5 cm long, the inner side purplish-red. Spikes 5-8, heteromorphic, 1 male (sometimes gynecandrous) terminal spike and 4-7 female lateral spikes; terminal spike 2.2- 3.5 cm × 3.5-4.5(11) mm, terete, sessile or with a peduncle up to 1 cm; lateral spikes, 1-5 cm × 9-10 mm, dense-flowered, terete, arising singly or in pairs, peduncle up to 4.5 cm. Male glumes 5.5-8 × 1.2-2.2 mm, oblong-elliptic, dark purplish-red with very wide and straw coloured midrib, acuminate, acumen up to 1.5 mm. Female glumes 4-7.2 × 1.2-1.8 mm, oblong-ellip- tic, dark purplish-red, with a wide and straw coloured midrib and an acumen up to 2 mm. Utricles 4.3-5 × 1- 1.5 mm, ellipsoid-trigonous, straight, plurinerved, although the veins are not very prominent, tapered to beak, brown to reddish-brown; beak 1-1.6 mm, smooth truncated or bidentated, with ventral sinus deeper than the dorsal one, the latter up to 0.2 mm depth. Achenes 2-2.4 × 0.9-1.1 mm, trigonous, oblong-ellipsoid to oblong-obovoid. 2n =? Feb. Marshy grassland; 2400 m; Tanzania (Uluguru mountains). Comments: Although only a single collection was studied, we consider that these specimens merit status as a new species due to: (1) their morphological differences to the other species of this group, and (2) their geographical isolation (the only other species of this group found in the Uluguru mountains is C. vallisrosetto which is clearly different to C. uluguruensis). Selected specimens TANZANIA: Tanganjika, Morogoro, Uluguru, Lukwangule Plateau, Savannenhügel, 2400 m, 20-2-1933; H.J. Schlieben, 3516 (PRE, L 625309, WAG 112513, B 100240022). 12. Carex vallis-rosetto K. Schum. in Engl. (ed.), Pflanzenw. Ost-Afrikas, C: 130. 1895. Ind. loc.: “(Usb., Rosettothal …)” “An Bachbetten im Gürtelwalde; auch über demselben in Erika-Bestän- den bis 2800 m”. Lectotype (designated in Gehrke, 2011): Holst 3823 (B 100166181!); isotype: K 363446 photo! = Carex cyrtosaccus C.B. Clarke in Thiselton-Dyer (ed.), Fl. Trop. Afr. 8: 524. 1902. Ind. loc.: “Nyasaland, Mt. Milanji”. Lectotype (designated here): A. White s.n. (K 363565 photo!); isotype: BR 898011!; paratypes: A. White s.n. (K 363563 photo!), “Nyasaland, Mt. Zom- 244 Anales del Jardín Botánico de Madrid 68(2): 225-247, julio-diciembre 2011. ISSN: 0211-1322. doi: 10.3989/ajbm: 2256 Fig. 15. Distribution of Carex uluguruensis: B, Burundi; R, Rwanda; S, Somalia. 2256 carex_Maquetación 1 28/11/2011 11:46 Página 244 ba, 4000-6000 ft.” A. White s.n. (K 363564 photo!). Rhizome caespitose, with short internodes. Culms (46)70-190(215) cm, very stout, trigonous, smooth, green to yellowish-green, but densely dark purplishred spotted or tinted towards the base. Leaves 25-76 cm × 7.5-13.5 mm, plicate, with revolute margins, slightly scabrid on margins and apical parts; sheaths dark purplish-red; ligule up to 6 mm, acute or obtuse; anteligule edge flat to convex. Inflorescence 14- 76(98) cm, lowest internode 5-25(31) cm and second one 2.5-17.5(25) cm. Lowest bract 21-40 cm × 3- 7.5(10) mm, longer or shorter than the inflorescence; sheath 3-10.5(15.5) cm long, the inner side purplishred. Spikes 7-22, homomorphic to heteromorphic, 0- 6 male spikes at the top and up to 17 female or androgynous spikes; male spikes 1-6.5(11.5) cm × (2)3- 7(8) mm, terete, sessile or with a peduncle up to 3.3 cm; lateral spikes, (1)2-9(12) cm × 7-11 mm, denseflowered, terete, mostly arising in pairs or groups of three, the rest single, frequently with some short branches at the base of the largest spikes, sessile or with a peduncle up to 6 cm. Male glumes (3.5)4.5-6.5 × 1-2 mm, oblong-lanceolate to oblong-obovate, dark reddish-brown to purplish-red with straw or green coloured midrib, mucronate, mucro up to 1.3 mm, or obtuse. Female glumes (3)3.5-5.7 × 1-2 mm, oblong-lanceolate, dark reddish-brown to purplishred with straw or green coloured midrib, mucronate or acuminate, mucro or acumen up to 1.5 mm, rarely obtuse. Utricles 4-5.5 × 1.2-1.8 mm, ellipsoid to obovoid, clearly curved, tetra-nerved (sometimes pluri nerved), gradually or abruptly narrowed into a beak, greyish-green with purplish-red speckles, sometimes entirely purplish in the upper half, trigonous; beak 1-1.8 mm, bifid, with ventral sinus deeper Taxonomic revision Carex tropical African group than dorsal one, the latter 0.2-0.5 mm depth, smooth or with some scattered pricklets. Achenes 2.5-3.1 × 0.9-1.2 mm, trigonous, oblong-elipsoid to oblongobovoid. 2n =? Jul.-Mar. Wet places: streams and bogs; 1750- 2500 m; Tanzania and Malawi. Selected specimens MALAWI. Southern, Mt. Mulanje, Litchenya Plateau, near C.C.A.P. Hut, 1990 m, 28-11-1986, J.D. Chapman & E.G. Chapman, 8276 (MO 3255780, PRE). Williams falls Zomba Plateau, 21- 10-1986, A.J. Salubeni, 4768 (MO 3917787, PRE). N Nkhata Bay, ENE Chikangawa, 1780 m, 13-10-1978, E. Phillips, 4081 (WAG 112783). 1962, P.J. Tyrer, 746 (BM 898023). Mt. Milanji, 1891, White s.n. (BM 898011). TANZANIA. Same, Kilimanjaro, T3, south Pare Mountains, Chome Forest Reserve (Shengena Forest), 1900 m, 18-11-1999, J. Elia, 130 (MO 5337368, PRE). Iringa, Mufinidi, Luisenda, 1830 m, 24-8-1984, D.W. Thomas, 3577 (MO 3298505, K). Morogoro, Uluguru Mountains, Luwkwangulu Plateau, 2400 m, 19-9-1970, M. Thulin & B. Mhoro, 1060 (UPS). Tanganyika, Morogoro, Uluguru Mts., Lukguangule plateau, above Chenzema Mission, 2500 m, 13-3-1953, R.B. Drummond & J.H. Hemsley, 1511 (BR, NY). Tanganyika, Tanga, Lushoto, Mkuzi Forest Reserve, 18-10- 1962, S.R. Semsei, 3527 (PRE). Tanganyika, Southern, Luwira- Kitega Forest, 25-10-1962, S.R. Semsei, 2550 (PRE). Tanganyika, T.T., Southern Highlands, Kigogo, Mufindi, 1800 m, 7-1959, J. Procter, 1284 (PRE, K). Tanganyika, Rungwe, Ngozi, Poroto Mts., 2100 m., 17-10-1956, H.M. Richards, 6573 (K). Mbeya, Poroto Mts., Ngozi, 1950 m, 17-10-1956, H.M. Richards, 6577 (K). Usambara, Rosetto-Thal an Bächen, 9-1882, Holst (B 100166182). Usambara, 9-1892, Holst, 3823 (B 100266181). Acknowledgements The authors wish to thank S. Martín-Bravo and P. Jiménez- Mejías for critical comments; and M. Laínz for the Latin diagnosis. References Baker, J.G. 1877. Flora of Mauritius and the Seychelles. Reeve. Londres. Clarke, C.B. 1902. Cyperaceae. In: Dyer, T. (ed.), Flora of Tropical Africa 8: 266-527. L. Reeve and Co. London. Escudero, M. & Luceño, M. 2009. Systematics and Evolution of Carex sects. Spirostachyae and Elatae (Cyperaceae). Plant Systematics and Evolution 279: 163-189. Escudero, M., Valcárcel, V., Vargas, P. & Luceño, M. 2008. Evolution in Carex L. sect. Spirostachyae (Cyperaceae): a molecular and cytogenetic approach. Organisms Diversity and Evolution 7: 271-291. Gehrke, B. 2011. Synopsis of Carex (Cyperaceae) from sub-Saha- ran Africa and Madagascar. Botanical Journal of the Linnean Society 166: 51-99. Govaerts, R. & Simpson, D.A. 2007. World Checklist of Cypera - ceae: Sedges. Richmond, Surrey, RBG Kew. Kew. Haines, R. & Lye, K.A. 1983. The sedges and rushes of East Africa. East African Natural History Society. Nairobi. Hooper, S.S. & Napper, D.M. 1972. Cyperaceae. In: Hepper, F.N. (ed.), Flora of West Tropical Africa, Vol. III, Part II, second edi- 245 Anales del Jardín Botánico de Madrid 68(2): 225-247, julio-diciembre 2011. ISSN: 0211-1322. doi: 10.3989/ajbm: 2256 Fig. 16. Distribution of Carex vallis-rosetto: M, Malawi; MA, Madagascar; Z, Zimbabwe. 2256 carex_Maquetación 1 28/11/2011 11:46 Página 245 tion: 278-349. Crown Agents for Oversea Governments and Administrations Millbank. London. Kükenthal, G. 1909. Cyperaceae-Caricoideae. In: Engler, A. (ed.), Das Pflanzenreich IV(20). Engelmann. Leipzig. Kükenthal, G. 1914. Carex longipedunculata f. recedens. In: Friedrichs, A. (ed.), Wissenschaftliche ergebnisse der Deutschen Zentral-Africa-Expedition, 1907-1908: unter Führung Adolf Friedrichs, herzogs zu Mecklenburg: 53. Klinkhardt & Biermann. Leipzig. Kükenthal, G. 1925. Cyperaceae. In: Fries, R.E. (ed.), Wissenschaftliche Ergebnise der schwedischen Rhodesia-Kongo- Expedition 1911-1912 unter Leitung von Eric Graf von Rosen. Band I, Erg. -Heft. 12: 308-312. Kükenthal, G. 1934. Cyperaceae. In: Norlindh, T. & Weimarck, H. (eds.), Beiträge zur Kenntnis der Flora von Süd-Rhodesia. II. Botaniska Notizer (Lund) 1934: 64-83. Lamarck, J.-B. 1792. Encyclopédie Méthodique: Botanique vol. 3: 387. Panckoucke. Paris. Luceño, M. 1992. Cytotaxonomic studies in Iberian and Ma - caronesian species of Carex (Cyperaceae). Willdenowia 22: 149-165. Luceño, M. & Castroviejo, S. 1993. Cytotaxonomic studies in the sections Spirostachyae (Drejer) Bailey and Ceratocystis Dumort. of the genus Carex L. (Cyperaceae), with special reference to Iberian and North African taxa. Botanical Journal of the Linnean Society 112: 335-350. Luceño, M. & Escudero, M. 2008. Carex sect. Spirostachyae. In: Castroviejo, S, Luceño, M, Galán, A, Jiménez-Mejías, P., Cabezas, F. & Medina, L. (eds.), Flora iberica, vol. XVIII: 178- 191. Real Jardín Botánico, CSIC. Madrid. Nelmes, E. 1938. XXXVI-Notes on Carex: IV. The section Elatae in Tropical Africa. Bulletin of Miscellaneous Information Kew 1938: 242-248. Richard, A. 1950. Tentamen florae Abyssinicae, vol. 2: 513-514. Arthus Bertrand. Paris. Schkuhr, C. 1806. Beschreibung und Abbildung der theils bekannten, theils noch nicht beschriebenen Arten von Riedgräsern, nach eigenen Beobachtungen und vergrösserter Darstellung der kleinsten Theile, vol. 2: 43. Wittenberg. Schumman, K. 1895. Carex L. In: Engler, H.G.A. (ed.), Die Pflanzenwelt Ost-Afrikas und der Nachbargebiete: 129-130. Geographische Verlagshandlung Dietrich Reimer. Berlin. Appendix 1 List of numbered exsiccatae The numbers in parenthesis refer to those given in the numerical list of taxa. Amshoff, G.J.H. 1965 (9a) Ash, J. 1696 (10) Bamps, P. 3064 (4b) Balfour, I.B. sn (1) Barclay, C. 1334 (1) Bauer, P.J. 68 (9a) Bequaert, 4680 (6) Bie, S.W. 66246 (5), 66266 (10) Bogdan, A.V. 552 (9a), 2655 (9a), 3525 (9a), 4826 (9a) Boivin, sn (2), 23bis (1), 996 (2) Bory, 1003 (1) Brunt, M. 777 (9a) Buide, M.L. 24UPO-K (9a), 24UPO-Kbis (9a), 29UPO-K (9a), 30UPO-K (9a), 31UPO-K (3), 53UPO-K (9a), 56UPO-K (9a), M. Escudero & M. Luceño 75UPO-K (5), 76UPO-K (6a), 79UPO-K (6a), 83UPO-K (4a), 92UPO-K (6b), 100UPO-K (9a), 101UPO-K (10), 103UPO-K (6b), 104UPO-K (4a), 105UPO-K (6b), 116UPO-K (10), 123UPO-K (9a), 132UPO-K (6b) Burger, W. 1902 (10) Cadet, T. 451 (1) Carvahlo, 3677 (6a) Chapin, J.P. sn (4b) Chapman, J.D. sn (12), 3575 (9a) Charmichael, W. 1549 (5) Clover, 1486 (9a) C.N.A.D., 1814 (9a) Coombe, D.E. 229 (9a) Croat, T.B. 28307 (6a) D’Alleizette, C.H. (2) D’Arcy, W.G. 7574 (4b), 7789 (6), 8280 (6a), 8317 (4b), 8520 (4b) Davidse, G. 7031 (4a, 6a) Drummond, R.B. 1511 (12) Domke, 308 (9a) Dummer, R.A. 3459 (6a), 3465 (4a) Edwards, S. 502/82 (10) Eggeling, W.J. 1070 (6a) Elia, J. 130 (12) Endlich, R. 626 (4a, 5) Evans, I.M. 482 (10) Fries, R.E. 405 (9a), 493 (4a), 659 (9a), 676 (5), 943 (9a), 1158 (5), 1228 (6b), 1300 (10), 1462 (4a), 2210(4a), 2533 (6a), 2552 (6b), 2553 (6b), 2703 (6b), 2763 (9a) Friis, I 853 (6c), 1640 (6c), 3504 (9b), 7150 (6c) Geilinger, 4374 (5) Gereau, R.E. 1667 (4a) Germain, 3723 (4b), 3724 (4b) Greenway, P.J. sn (4a), 7121 (9a), 13666 (9a) Gibbs Russell, G.E. 1208 (9b) Haines, R.W. 274 (4b), 4315 (6a) Hedberg, O. 193 (4a), 397(4a), 854 (3), 1017 (10), 1043 (10), 1089 (9a), 1546 (10), 1600 (6b), 1633 (6a), 1936 (10), 2179 (4a), 2420 (4a), 4181 (10), 4224 (10), 4330 (10), 4555 (3), 4828 (10), 5659 (10), 7047 (10), 7068 (10) Hepper, F.N. 2042 (9a), 2085 (9a), 2085A (9a), 4962 (6b) Hienthmay, 10162 (2) Holst, sn (12), 3823 (12) Hooker, sn (2) Howard, W.J. 32 (6c) Humbert, H. 7999 (6a) Jackson, J.K. 1550 (4a) Johnston, H.B. 1408 (6c) Johnston, H.H. sn (2) JMG, 93937 (5), 9464 (4a), Katende, 3004 (4b) Keken, P. van der 9125 (10), 9126 (4b, 10) Laegaard, S. 16024 (9b) Lebrun, J. 7450 (4b), 8747 (6a), 8780 (4b, 9a), 9332 (9a) Liebenberg, L.C. 1708 (6a) Linder, D.H. 2076 (6a), 2252 (4b), 2364 (4b) Lisowski, S. 10936 (4a), 10956 (6a) Luceño, M. 127ML08 (2), 1ML09 (1), 2ML09 (2), 13ML09 (2), 14ML09 (2), 21ML09 (2) Louis, J. 5404 (4b) Lye, K.A. 5271 (6a), 5287 (6a), 5300 (6a), 1259 (4a) Maas Geesteranus, R.A. 5475 (4a), 5882 (6b), 5966(4a), 6015 (6b, 9a) Marien, W. 678 (6b) 246 Anales del Jardín Botánico de Madrid 68(2): 225-247, julio-diciembre 2011. ISSN: 0211-1322. doi: 10.3989/ajbm: 2256 2256 carex_Maquetación 1 28/11/2011 11:46 Página 246 Mearns, E.A. 899 (9a), 1674 (4a), 1677 (10), 1706 (10), 1736 (4a), 1742 (10), 1765 (4a, 6a), 2359 (9a) Milbraed, J. 966 (7), 1338 (6a), 1578 (4b) Mwangangi, O.M. 373 (3) Napper, D.M. 2133 (9a) Newbould, J.G.B. 3373 (9a) Norman, E.M. 164 (6a) Peter, A. 792 (5), 1298 (5) Phillips, E. 118 (9b), 145 (9b), 396 (9b), 530 (9b), 4081 (12) Piemeisel, R.L. 599 (6b) Polhill, 429 (9a) Preufs, sn (9a) Prins-Lambert, J. 317 (8) Procter, J. 1284 (12) Purseglove, J.W. 325a (4a) Rammeloo, J. 5016 (4b) Reekmans, M. 11054 (7) Rehder, H. sn (4a, 6, 10) Richard, M. 530 (2) Richards, H.M. 6573 (12), 6577 (12) Robinson, E.A. 3028 (9b) Runyinya, B. 738 (6A) Ryvarden, L. 5576 (4a) RW, 530 (8) Salubeni, A.J. 4768 (12) Taxonomic revision Carex tropical African group Saxer, A. 166 (6a) Schimper, 1180 (10), 1289 (10), 1298 (9a), 1988 (10) Schlieben, H.J. 3516 (11), 4487 (4a, 5), 10834a (2), 10894 (1), Scott Elliot, G.F. 7578 (9a) Semsei, S.R. 2550 (12), 3527 (12) Shuttleworth, R.J. sn (2) Spach, sn (2) Stauffer, H.U. 69 (9a), 777 (6a), 836 (4b) Stolz, A. 2284 (8) Szczepanek, K. 820 (4a) Taylor, G. 1897 (9a), 2944 (4a), 3424 (4a), 3474 (3), 3677 (10) Thomas, D.W. 3577 (12), 6045 (9a) Thulin, M. 243 (10), 1060 (12), 1466 (9a), 3716 (10) Troupin, G. 14300 (4b), 14301 (10) Verdcourt, B. 2005 (6b) Volkens, G. 1291 (5), 1342 (5) Wettstein, Fr.u.R. sn (4a) White s.n. (12) Wilde, W.J.J.O. 317 (4a), 2541 (9a), 6423(4a), 6431 (9a), 6787 (10), 7409 (4a), 8124 (10), 8542 (4a), 9119 (10), 10307 (9a) Associate Editor: I. Álvarez Received: 26-III-2010 Accepted: 11-III-2011 247 Anales del Jardín Botánico de Madrid 68(2): 225-247, julio-diciembre 2011. ISSN: 0211-1322. doi: 10.3989/ajbm: 2256 2256 carex_Maquetación 1 28/11/2011 11:46 Página 247