196 Bothalia 42,2 (2012) Martínez-AzorínMartínez-Azorín et al.: 467 ((2011):2011): Baker: t. 335 ((1872).1872). baria ABH,ABH, BOLBOL,, BNRHBNRH,, GRA,GRA, J,J, K,K, KEI,KEI, KMG,KMG, NBG,NBG, Epitype, designated by Martínez-Azorín et al.: 467 NH, NU, PEU, PUC, UFH, and WIND, for allowing us (2011): South Africa, Eastern Cape, Somerset East, sum- toto study theitheirr AAlbucalbuca collections. Thanks to P. Wilkin,Wilkin, mmitit of Boschberg, northeast of Bloukop, in shallow soil O. Weber and L. Walsingham (Royal Botanic Gardens, oonn rocky places, 1 574 m, 10 May 2010, V.R.V.R. CClarklark & Kew) for supplying images of some AlbucaAlbuca collectionscollections M. MMartínez-Azorínartínez-Azorín (GRA, epi.; ABH, K, PRE, isoepi.) at Kew. We also thank F. Martínez-Flores who kindly helpedhelped us with the fi gures of AlbucaAlbuca lleaves.eaves. Diagnostic characters: Albuca tenuifolia ccanan be eeas-as- iilyly identifi ed by its hypogeal and proliferous, irregularly ccompressedompressed bulbs; fi liform and straight leaves that beabearr REFERENCESREFERENCES mminuteinute and sessile papillae (only visible under micro- sscope);cope); glabrous peduncle and pedicels; erect pale yel- BAKER,BAKER, J.G.J.G. 1872. Albuca tenuifolia. In W.W. Saunders,Saunders, RefugiumRefugium botanicum 5(2)5(2) t. 335. Van Voorst, Paternoster Row, London.London. llow-greenow-green fl owers; strongly hooded inner t epal tips; all BAKER,BAKER, J.G.J.G. 1873. Revision of the genera and species of ScilleaeScilleae andand sstamenstamens bearing fertile anthers; subglobose ovary with ChlorogaleaeChlorogaleae. JournalJournal of the Linnean Society. Botany 13(68):13(68): divergent paraseptal crest; and a narrowly obpyramidal 209-292.209-292. ttrigonousrigonous style. BAKER,BAKER, J.G.J.G. 1874. Description of new species of ScilleaeScilleae andand ootherther LiliaceaeLiliaceae. JournalJournal of Botany, British and Foreign 12: 363-369.363-369. The characteristic form of the inner tepal tips, BAKER,BAKER, J.G.J.G. 1897. AlbucaAlbuca. In W.T. Thiselton-Dyer, Flora Capensis 6:6: 451-462.451-462. Reeve & co.,co., LondonLondon.. sstronglytrongly hooded and resembling a mitre (Figure 2a) HILLIARD,HILLIARD, O.M. & BURTT,BURTT, B.L. 1982. Notes on some plants of ttogetherogether with the erect fl owers, place A. tenuifolia in AA.. southernsouthern Africa, chiefl y from Natal: IX. Notes from the Royal ssubg.ubg. MMitrotepalumitrotepalum ((== AA.. secsect.t. BrBrancionaanciona). FForor furtherfurther BotanicBotanic Garden, Edinburgh 40: 247-298.247-298. details and discussion on subgeneric arrangement of MCNEILL,MCNEILL, J.,J., BARRIE,BARRIE, F.R.,F.R., BURDET,BURDET, H.M.,H.M., DEMOULIN,DEMOULIN, V.,V., HAWSWORTH,HAWSWORTH, D.L.,D.L., MARHOLD,MARHOLD, K.,K., NICHOLSON,NICHOLSON, D.H.,D.H., AAlbucalbuca see Müller-Doblies ((1987,1987, 19951995).). PRADO,PRADO, J.,J., SILVA,SILVA, P.C.,P.C., SKOG,SKOG, J.E.,J.E., WIERSEMA,WIERSEMA, J.H. & TURLAND, N.J. (EDS.).(EDS.). 2006. INTERNATIONALINTERNATIONAL CODECODE OFOF AAlbucalbuca shawiishawii Bakerr in Journal of Botany (London) BOTANICALBOTANICAL NOMENCLATURE (VIENNA(VIENNA CODE)CODE) ADOPTADOPT-- 12: 367 (1874). Lectotype, designated by Hilliard & EDED BY THE SEVENTEENTHSEVENTEENTH IINTERNATIONALNTERNATIONAL BOTANICALBOTANICAL BBurtt:urtt: 281 (1982): South Africa, Northern Cape, Coles- CONGRESS VIENNA,VIENNA, AUSTRIAAUSTRIA,, JULY 20020055. GARTNERGARTNER VER-VER- LAG, RUGGEL, LIECHTENSTEIN [REGNUMREGNUM VEGETABILEVEGETABILE berg, SShawhaw s.n. (K000257312, lecto.—photo!)lecto.—photo!).. 146].146]. MANNING,MANNING, J.C. & GOLDBLATT,GOLDBLATT, P.P. 2003.2003. AlbucaAlbuca. In G.G. Germishui-Germishui- Diagnostic characters: AAlbucalbuca shawii iiss ccharacterizedharacterized zen & N.L. Meyer, Plants of Southern Africa: an annotated by its hypogeal, ovoid, and mostly solitary bulbs; fi li- checklist. Strelitzia 14: 1054-1056. National Botanical Institute,Institute, form leaves with usually coiled apexes; leaves, pendun- Pretoria.Pretoria. ccle,le, and pedicles with evident, stipitate glands; nodding MARTÍNEZ-AZORÍN,MARTÍNEZ-AZORÍN, M.,M., CLARK,CLARK, V.R.,V.R., CRESPO,CRESPO, M.B.,M.B., DOLD A.P.A.P. & BARKER N.P. 2011. The rediscovery of Albuca tenuifo-tenuifo- yellow-green fl owers; inner tepals with a slight apical lia, an orophilous species from the eastern Great Escarpment in hhoodood and a triangular structure facing downwards; only SouthSouth Africa. Nordic Journal of Botany 29: 465-470.465-470. tthehe three inner stamens bearing fertile anthers, the out- MÜLLER-DOBLIES,MÜLLER-DOBLIES, U. 1987. Infrageneric taxonomy of AlbucaAlbuca (Hya- ers commonly lacking anthers; oblong ovary with almost cinthaceae).cinthaceae). Abstract XIV International Botanical Cong ress p. 332.332. Berlin.Berlin. sstraighttraight paraseptal crests; and the shorter and widely MÜLLER-DOBLIES,MÜLLER-DOBLIES, U.U. 1995. Enumeratio Albucarum (Hyacinthace- oobpyramidalbpyramidal trigonous style. ae) Austro-Africanarum adhuc cognitarum. 2. Subgenus Falcon- era ((Salisb.)Salisb.) Baker emend. U.M.-D 1987. Feddes Repertorium The inner tepals tips of AA.. shawii are neither strongly 106: 353-370.353-370. hhoodedooded nor hinged (Figure 2b), a character which, MÜLLER-DOBLIES,MÜLLER-DOBLIES, U. 2012. The enigma of Albuca tenuifolia BakerBaker resolved.resolved. Feddes Repertorium 121: 285-298.285-298. ttogetherogether with the nodding fl owers, place this species in THIERS,THIERS, B.B. 2012. Index herbariorum: a global directory of public her- A. subg. FFalconeraalconera ((== AA.. secsect.t. FFalconeraalconera) ((cf.cf. Müller- baria and associated staff. [continuously updated]. New York. Doblies 1987, 19951995)) The species is the type of AA.. secsect.t. Bot.Bot. Gard. Virt. Herb.
* CIBIO (Instituto(Instituto Universitario de la Biodiversidad),Biodiversidad), Universidad de Alicante, P.O. Box 99, E-03080 Alicante, Spain. [email protected]@ua.es (au(au-- ACKNOWLEDGEMENTSACKNOWLEDGEMENTS thor for correspondence). **** SelmarSelmar Schonland Herbarium, Department of Botany, Rhodes Uni-Uni- This work was partly supported by the Fundación versity,versity, Grahamstown 6140, South Africa. Ramón Areces (Spain). We thank the curators of the her- MS.MS. received: 2012-01-31.2012-01-31.
ITEACEAE
TAXONOMIC AND NOMENCLATURAL NOTES ON THE GENUS CHORISTYLIS
Since the description of the sub-Saharan African status and family position of the genus and summa- Choristylis and its only species, C. rhamnoides by Har- rize its nomenclatural history. Harvey (1862), Hooker vey in 1842, the family position of this woody shrub or (1865), Mildbraed (1934), Boutique (1964) and Liben small tree has been uncertain. We review the taxonomic (1969) referred Choristylis to Saxifragaceae, and Engler Bothalia 42,2 (2012) 197 (1891, 1930) placed it in Saxifragaceae subfamily Escal- 2–5 united carpels. Although the carpels of Iteaceae are lonioideae. Other authors, including Hutchinson (1969), free initially, they become partly or completely united Verdcourt (1973, 1983), Brummitt (1992), Jordaan along the style (Ge et al. 2002; Kubitzski 2006). (2000), Lötter (2002), Coates Palgrave (2002) and Boon (2010) included Choristylis in the Escalloniaceae. Airy The African Choristylis is notably disjunct from other Shaw (1973) was the fi rst to associate Choristylis with members of Itea, which are exclusively Northern Hemi- Itea in the small family Iteaceae J.G.Agardh (sweet- sphere in distribution, extending from south and south- spire family), a step followed by Takhtajan (1996) and east Asia from the Himalayas to China, Taiwan, Japan, Kubitzki (2006). This is the family placement followed Java, the Philippines, and southeast United States of here. America. Choristylis, however, is similar in habit, leaf, fl ower, fruit, and seed characters to Itea. Members of Saxifragaceae are exclusively herbaceous with exstip- Itea have alternate or spirally arranged, glandular ser- ulate leaves. Escalloniaceae are trees with exstipulate rate or dentate to spinose-dentate leaf margins with the leaves and a parietal placentation, whereas in the woody lateral or tertiary veins ending at the margin in a gland Iteaceae the leaves are always stipulate, placentation is or spine (the same condition as in Choristylis) to rarely axile and the pollen grains are unusual in being 2-porate entire. The leaf venation is seemingly 3-veined from and subisopolar (Erdtman 1955; Agababian 1960). Pol- or near the base in Choristylis and pinnately veined in len in Escalloniaceae is shed as monads and the grains all species of Itea. Both genera have similar, small, 5- are isopolar and 3–5-colporate. Iteaceae also differ from merous, creamy white or greenish yellow fl owers Escalloniaceae in their bitegmic and crassinucellate borne in axillary or terminal panicles or racemes, often ovules. Escalloniaceae have unitegmic and tenuinucel- superposed in groups of two or three. In Choristylis late ovules. The crassinucellate condition was deter- the infl orescences are shorter than the leaves, up to 45 mined in both Itea and Choristylis (Mauritzon 1933, fi de mm long, whereas most Itea species have long, droop- Kubitzki 2006). APG III (2009) place Escalloniaceae ing, elongated panicles or racemes, up to 60–200 mm in an order of its own, the Escalloniales. Fossil fl owers long, much longer than the leaves (Jin & Ohba 2001; resembling those of Iteaceae and dating from the Turo- Siddiqi 2005; Morin 2009). The fl owers in both genera nian stage of the Late Cretaceous have been found in the have a turbinate or obconic calyx-tube adnate to the base Raritan Formation, New Jersey, and leaves ascribed to of the ovary and with fi ve lobes; the petals are valvate Iteaceae and dating to the Eocene have been described and persistent; the stamens are inserted at the margin from the Klondike Mountain Formation, Washington of a hypogynous disc and are included or exserted; the (Hermsen et al. 2003; Friis et al. 2011). ovary is 2-locular, superior to half-inferior; the styles are undivided or ± deeply divided, but apically coherent Molecular analyses place Itea and Choristylis in the at anthesis. The fruit is a septicidally dehiscent capsule core Saxifragales, sister to Pterostemon Schauer (Pter- with a persistent perianth in both genera. Given the gen- ostemaceae) (Savolainen et al. 2000; Fishbein et al. eral morphological similarity between the two genera 2001). Pterostemonaceae (Engl.) Small is a monogeneric and their identical pollen we agree with Kubitzki (2006) family of two species of shrubs native to tropical and that the range of variation of characters, such as ovary subtropical Mexico. This family, formerly associated position (nearly superior to half inferior), the degree of with Grossulariaceae s.l., Escalloniaceae, and Hydrange- fusion of the styles (undivided or ± deeply divided,) and aceae by some authors, is currently included in Iteaceae ramifi cation and length of infl orescences (terminal or by APG III (2009). Studies done by Bohm et al. (1999) axillary panicles or racemes), makes the maintenance of showed that the fl avonoid profi les of Pterostemon con- Choristylis impossible. sist of C-glucosylfl avones and 3-O-glycosides of querce- tin, which resembles very closely that observed in Itea. Bentham (1865) kept Choristylis and Itea separate The presence of C-glucosylfl avones in these two genera when only a few species of Itea were known, and stated provides additional support for the closeness of their that the one Itea species from America (I. virginica L.) relationship. In contrast, Saxifragaceae, Crassulaceae, and one from Japan (I. japonica Oliv.) are deciduous and and other members of the Saxifragales are characterized that their seeds are oblong and fl attened, distinguishing by fl avonol production (Bohm et al. 1988). them from the Asian species, which are evergreen trees and their seeds are fusiform. Since then many more Kubitzki (2006) relegates Choristylis to the syn- species of Itea have been described and Engler (1891, onymy of Itea, a genus with about 19 species, of which 1930) consequently took up these characters mentioned 15 species (ten endemic) occur in south and southeast by Bentham and used them to distinguish two sections Asia and Malaysia (Jin & Ohba 2001), one in Paki- in Itea, namely sect. Deciduae Engl. for the American stan (Siddiqi 2005), one in Japan (Ohwi 1965), one in and Japan species, and sect. Sempervirentes Engl. for the North America (Morin 2009) and one species in Africa Asian species. (Verdcourt 1973, 1983). Members of Iteaceae have spi- rally-arranged or alternate, glandular-dentate, serrate to Although Kubitzki (2006) argued that Engler’s sec- spinose leaves with superposed axillary buds, paniculate tions (1891, 1930) in Itea could not be upheld with the to racemose infl orescences with rather small 5-merous inclusion of Choristylis within Itea, the seed morphol- fl owers, and septicidally dehiscent fruits, with the valves ogy of Choristylis differs enough from the other species often remaining attached by the stigma. The fl owers to warrant a section of its own in the genus Itea. Chori- have a hypanthium, the corolla is valvate, and there are stylis is an evergreen scrambling shrub or tree and the only two carpels. Flowers of Saxifragaceae have four or seeds are irregularly obovoid or oblong-obovoid, fl at- fi ve free sepals and petals, with stamens as many as or tened, sculptured and slightly curved (Verdcourt 1973, twice as many as the sepals, and the pistil consisting of tab. 1,13; Kubitzki 2006, fi g. 70G). We accordingly 198 Bothalia 42,2 (2012) describe a new section within Itea to accommodate the the base, short infl orescences less than 45 mm long, and single African species. seeds which are irregularly obovoid or oblong-obovoid, fl attened, sculptured and slightly curved. TAXONOMY Type species: Itea rhamnoides (Harv.) Kubitzki. Itea L. sect. Choristylis (Harv.) Jordaan, sect. nov. (=Choristylis rhamnoides Harv.) et stat. nov. Choristylis Harv.: 19 (1842). Similar to sect. Sempervirentes Engl. in its evergreen habit but differ- Itea rhamnoides (Harv.) Kubitzki in K. Kubitzki, ing in its leaves seemingly three-veined from or near The families and genera of vascular plants 9: 203
B
E
C
A
D
FIGURE 1.—Itea rhamnoides, A, fl owering twig, × 1; B, leaf with irregular glandular-serrated margin, × 2; C, young fl ower, × 15; D, mature fl ower, × 15; E, fruit, × 15. A & C taken from Thode A318 (PRE); B taken from Dahlstrand 2669 (PRE); D taken from Jordaan 3938 (PRE); E taken from Theron 3648 (PRU). Artist: Daleen Roodt. Bothalia 42,2 (2012) 199 (2006). Choristylis rhamnoides Harv.: 19 (1842), 308 ular, hermaphrodite or polygamous, 5-merous, creamy (1862). Type: South Africa, Eastern Cape, Katberg, white or greenish yellow, sweetly scented; pedicels Brownlee s.n. (TCD, holo.–Aluka image!; K, iso.). 1–2(–5) mm long. Calyx tube (hypanthium) short, ± 1.5 × 1.5 mm, adnate to ovary; lobes triangular, 0.5–2.0 × Choristylis shirensis Baker f.: 13, t. 3/1–6 (1894). 0.3–0.5(–1.0) mm at base, ciliate. Petals ovate, 1.5–3.5 Type: Malawi, Mt. Milanji [Mulanje], 1891, Whyte 53 × 1–2 mm at base, 3-veined, hairy. Stamens slightly (BM–Aluka image!, lecto., designated here; Z–Zürich exserted, hairy; anthers ovoid, hairy. Disc annular. Virtual Herbarium!, isolecto.). [Whyte 53 in the British Ovary half-inferior, bicarpellate, carpels free initially, Museum is chosen as the lectotype as the best represent- 2-locular; ovules numerous, biseriate, with axile placen- ative specimen among the available syntypes]. tation; styles 2, subulate, adherent or diverging, 0.5–1.0 mm long, yellowish, turning brown; stigmas ± 0.5 mm C. ulugurensis Mildbr.: 191 (1934). Type: Tanganyika diam., free or sometimes cohering for a time. Fruit a [Tanzania], Uluguru Mts., southern slope of Lukwangule septicidally dehiscent capsule, campanulate to turbinate Peak, August 1933, Schlieben 4293 (B B100165111– below, conical above, 3–5(–6) × 3 mm, brownish, fi nely Aluka image!, lecto., here designated). [The type col- hairy, slightly ribbed, leathery. Seeds obovoid or oblong- lection of Schlieben 4239 on which Mildbraed (1934) obovoid, fl attened, ± 1.0 × 0.5 mm, reticulate, brownish. based his name consists of two specimens, of which Figure 1. number B100165111 is the better specimen with more fl owering material and therefore chosen as lectotype]. Diagnostic characters: Itea rhamnoides can be rec- ognized by its leaves, which are spirally arranged, ± Evergreen scrambling shrub or small tree up to 7 3-veined at or near the base, with glandular toothed mar- m tall, with slender trailing branches, single- or multi- gins and conspicuous tertiary reticulate venation. The stemmed; stems slender, diverging from base, angu- ovary is half-inferior with two diverging styles, whence lar-ribbed; young branchlets often purplish, covered is derived the name Choristylis, from the Greek word with short hairs. Bark on stems splitting longitudinally. Leaves spirally arranged, glossy green above, paler and choris = separate and stylis = style. The valves of the dull below, hairless or with sparse hairs on the main septicidally dehiscent capsules often remain attached by veins below and often with domatia; lamina ovate or the stigma. oblong to ovate-lanceolate, l5–103 × 9–55 mm, apex Distribution and habitat: Itea rhamnoides grows in acute to shortly acuminate or rarely rounded, base the eastern parts of southern Africa, from the Katberg, cuneate to unequally rounded; margins sharply and Hogsback, Keiskammahoek, and Kentani in the Eastern closely irregular glandular-serrate; seemingly 3-veined Cape, northwards through KwaZulu-Natal, Swaziland from base, but otherwise pinnately veined; midrib and and Mpumalanga to the Soutpansberg in the north. From main veins sunk above, prominent below, fi ne tertiary here it extends further northwards through Zimbabwe, reticulate venation conspicuous; petioles often purple, Mozambique, and Malawi along the mountains to the up to 20 mm long, pubescent. Stipules small, decidu- ous. Infl orescences many-fl owered panicles, shorter than Democratic Republic of the Congo, Burundi, Uganda, leaves, fi nely hairy, up to 45 mm long, in dense clusters; and Tanzania along the East African Rift, but seems to bracts and bracteoles narrowly triangular, 0.5–1.5 mm be absent from the more recent volcanic mountains in long; peduncles 0–12 mm long. Flowers very small, reg- East Africa (Kenya and Ethiopia) (Verdcourt 1983) (Fig- ure 2). The distribution coincides with that of the archi- pelago-like Afromontane Region of Endemism (White
H 1978, 1983). It grows in forest, forest margins, riverine 55 and valley forests, open hillsides, often along streams at medium to high altitudes. S 5 5 Synonymy: Harvey (1862) considered Baeobot- 5 IH 5 5 rys rufescens E.Mey. to apply to Choristyles rham- 5 5 noides, which De Candolle (1844) placed in the genus Maesa, based on a Drège specimen, and Maesa palus- IS 555 5 tris Hochst. (1844), based on a Krauss specimen (Krauss 5 5 5 152), under Choristylis rhamnoides. These synonyms, 55 PH 5 however, belong to Maesa lanceolata Forssk. (Dyer
555 1963). Verdcourt (1973) was the fi rst to place the tropi- 555 PS 55 55555 cal African Choristylis shirensis Baker f. (1894) from 555 555 Malawi and C. ulugurensis Mildbr. (1934) from Tanza- 555 5 555 nia in synonymy under C. rhamnoides. The name Chori- QH 555 5555 stylis virescens Baker f., which appears in The Interna- 5 H QHH THH km tional Plant Names Index [IPNI] as published on the QS same page where Baker described Choristylis shirensis IH IS PH PS QH QS RH RS (Baker 1894) seems to be fi ctitious (Verdcourt 1983). FIGURE 2.—Distribution of Itea rhamnoides, based on specimens We have checked Baker (1894) and also cannot fi nd any housed in the National Herbarium, Pretoria, and in literature mention of C. virescens. We conclude that it is a spelling (Liben 1969 & Verdcourt 1973, 1983). error for C. shirensis. 200 Bothalia 42,2 (2012)
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