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A Taxonomic Revision of Elaeodendron Jacq. (Cassinoideae: Celastraceae) in Africa

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A Taxonomic Revision of Elaeodendron Jacq. (Cassinoideae: Celastraceae) in Africa s. Air. 1. Bot. 1998.64(2) 93 A taxonomic revision of Elaeodendron Jacq. (Cassinoideae: Celastraceae) in Africa R.H. Archer' and A.E. van Wyk' ' National Botanical Institute, Private Bag X101 , Pretoria, 0001 Republic of South Africa e-mail: [email protected] l H G.W.J. Schw€lckerdt Herbarium, Department of Botany, University of Pretoria, Pretoria, 0002 Republic of South Africa Recen'!!d 16 ,lfllY 199 7: revised 23 Sepfemher /()() 7 The heterogeneous genuS;, Cassine L. s.l. is subdivided into smaller, more natural genera, following new evidence from macromorphology, palynology and anatomy. Elaeodendron Jacq. , one of the segregate genera, is here revised for Africa. The redefined genus Elaeodendron IS cosmopolitan, with consists of 30 to 40 species of shrubs and trees distributed in Africa, Asia, AustralaSia and central America, Eight species of EJaeodendron are recognized in Africa. A new combination, E. transvaaJense (Burtt Davy) R.H . Archer (:::: Cass;ne transvaaJensis) is proposed. The other species are EJaeodendron aquifolium (Fiori) Chiov" E buchananii (Loes.) Loes. , E. croceum (Thunb,) DC., E. matabeJiclIm Loes ., E. schJechterianum (Loes.) Loes. and E. zeyheri Spreng. ex Turcz. The correct application of the previously misapp:ied and confused name E. croceum is discussed. Keywords: Cassine, Cassinoideae, Celastraceae, EJaeodendron, taxonomy, 'To whom correspondence shou ld be addressed. Introduction Rllbenlia Juss.: 378. 452 (1789); Gille!.: 408 (1791). Type: II. This paper emanates from a multi~disciplinary study towards a o/ivina J,F. Gmel. [Elacodendrot1 orientale Jacq.]. taxonomic revision of the southern African Cassinoideae (Archer Schrebera Retz.: 25 (1791) 11011 L. nee Schreb., nee Roxb. Type: & Van Wyk 1996a). It has long been a matter of dispute whether S. albens Retz. to treat Cas.\"ine L. in a wide sense (e.g. Ding Hou 1962; Koster ~ mans 1(86), or to recognize a number of' segregate genera, Porlenschlag;a Tratt.: 250 (1818). Type: P australis TraIt. including Elaeodem/ron Jacq. (e.g. Loesener in Die natOrlichen Neerija Roxb.: 86 (1814); Carey el Wallich in Roxb.: 444 Ptlanzenfal11ilien, 1942a). Following Archer (1990) and Archer (1824); Carey in Roxb.: 646 (1832). Type: N dicho/ama Roxb. and Van Wyk (1992. 1993a, 1993b, J996a), five distinct southern A friean genera, the African/Madagascar AlystroxJJlol1 and the Crocaxyloll Eck!. & Zeyh., 128 (183 4/5); Harv.: 50 (1838); N. cosmopolitan E/aeodel1dron, can be conclusively recognized on Robson: 40 (1965); 390 (1966); N. Robson & Sousa: 36 (1969). Type: C cxcelsum Eckl. & Zeyh. 110m i/leg (E1aeodendron cro~ evidence frol11l11acrolllorphology. palynology, and anatomy. CelllJl (Thunb.) DC.), Our present concept of lassille s. str largely follows Loesencr (19420), Robson (1965. 1966. 1989) and Robson el al (1994) Cassine Loes.: 214 (1892), pro parte excl. typum; Davison 2: and necessitates many name changes back to the original 326 (1927) pro parle; sensu allet plllr., 11011 L. Type: C. peraguQ basionyms in Elaeoc/endnm for Illost species of em'sine .d, from L. type prop. cons. other parts of the world. Fortunately, most species have been wel! knowll under Elaeodendro}l, and in most cases correct com­ Cassine L. sect. Ellelacodendroll Loes.: 215 (1892). binations already exist. Approximately 40 species of Elaeodell~ Telemac/zia Urb.: 334 (1916). Type: lrinitensis Urb. (Elaeo­ droll can be recognized from central America, Africa and :r dendron allstrale VenL). Australasia. Eight taxa are confined to Africa, excluding the Mascarenes and Madagascar, and are here revised. One species, Pseudocassille Brcdell: 330 (1937); Com pIon: 338 (1976). Type: E/aeodelldron orientale Jacq. , which was treated as a cultivated P. fransl'oalellSis (Burtt Davy) Bredel!. species in Flora Zambesiaca (Robson 1966), seems to exist in that Flora area as a single tree in the Mutare Botanical Garden, Evergreen or rardy deciduous shrubs to trees; unarmed, glabrous, Zimbabwe. A synopsis of the known species of Elaeodendrol1 ,vithout elastic threads (Irans~1. 4-poly'isoprene) in bark and leaves: worldwide is in preparation and will be published elsewhere. bark with layers of ydlow pigment {l(;casionally present or hardly Herbarium specimens cited here are arranged by the quarter­ disccmible. Branchlels subangular to tcrete, knticcls usually promi ~ degree reference system of Edwards and Leistner (1971 ). One nent. Leaves opposite to sUboppositc. or occasionally alternate, spi~ specimen is cited per quarter degree square. Localities north of railed to fasciculate: stipuks minute. ± I mm long, ± triangular. the equator, or west of Greenwich are indicaled by the letters N marcescent. inflorescences usually pedunculate. dichasial. Flowers or W after the longitude or latitude respectively. Codes following lIsually bisexual. occasionally unisexual. sometimes with pctaloid the geographical divisions of the Flora (?f Tropical East .· ~ff'ica staminodes in female tlowers; 3-, 4- or 5~merous (except ovary), (Polhill 1988) have been added in the case of specimens cited for pedicellate. Sepals equal, subcircular. Petals cream to greenish, the latter region. oblong to ovate. entire or with ventral projections, spreading. Stamens erect to spreading, arising inside margin of disc or from Systematic treatment sinuses in the margin of the disc: anthers introrse or extrorse. dehisc~ Elll(!oticntiroll Jacq., leones Plantarull1 RarioruJl1 t. 48 (1782); ing by longitudinal slits. {)iSC entire to subentire and convex. fleshy. Murray: ::!41 (1784), as Elaeodcndrll1Jl; Jacq. f.: 36 ( 1787), as or tlat and quadrangular. with sinuses at the point of stamen inser~ F/aeodclldri: Loes.: 223 (1897); 172 (1942a); Thonner: 331 tion, nat 10 concave. Ovary ± Ih immersed in and adnate to the disc. (1915): N. Robson: 385 (1966); N. Robson el al. : 29 (1994). 2~. 3~ or 4~locular. with two erect collateral ovules per locule: style Type: E. oriemale Jacq. short to absent. st igma inconspicuous. Fruit drupaceous, spheroid to 94 s. Mr. J. Bot. 1998, 64(2) ellipsoid, white to yellow; stone narrowly ellipsoid to ellipsoid, sur­ 6b Leaf margin glandular-crenate; lam ina pak face smooth or with grooves. Seeds brownish, narrowly ellipsoid, yellowish green: so uthern tropical Africa . ovoid, flattened to triangular, postchalazal vascular bundles onen 6. E. mataheficunl present, endosperm present; embryo erect with cotyledons fleshy, Ib Flowers with ovary isomerous \vith other fl oral whorls. 3- or 4- ell iptic or ovate. locular ...... ... 7 71.1 Flowers 4-meroll s: Eastt!1"Il Cape & KwaZulu-Natal. ra re in E/aeodendron is one of the largest genera in the Celastraceae, Mozamb iq ue and Northern Prov ince . ...... .... 7. £. :;eyhen consisting mostly of shrubs or trees. An estimated 30 to 40 spe­ 7b Flowers 3-maolls: wi desprt!ad in southern Africa. cies occur in central America, Africa, Asia and Australasia. .. 8. E. t1"(lIlsvaalensC' Some members are commercially exploited or have potential for wood production. 1. Elaeodendron CrOCelll11 (Tbuob.) DC. Prodromus 2: 11 The African species of Elaeodendron are more frequent in (1825) [as Elaeodendran' l; Pappe: 10 (1 854); Sond.: 468 (1860) southern Africa and the eastern lowland parts of the continent. pro parle; Marloth: 154 (1925); Archer & Condy: 58 ( 1995) . Three species. E. buchananii. E. matabelicuin and E. trans­ Type: Thul1berg, sheet 3807 in Herb. Thu nb. (UPS!, holo). voalense, also extend to the drier parts of western tropical Africa. Much uncertai nty exists in recent li terature concerning the lIex cracea Thunb.: 32 ( 1794a); 1: 169; 2: 109 ( 1794b [original correct spelling and author citation of the name Elaeodendron Swedish edition, 1788]); Thunb.: 577 (1818); 159 ( 1823). Cass­ (Archer & Van Wyk 1996b). Kostermans (1986) pointed out that ine crocea (Thunb.) Kuntze: 114 ( 1891); Von Breitenbach: 637 Elaeodendl'um Murray has priority over Elaeodendron Jacq. f. , (1965), pro parle; Coates Palgrave: 510 (1977) pro porle; 11011 while most authors considered either Jacq. f. or Jacq. to be the Davison: 334 (1927). correct author of the generic name. Archer an d Van Wyk Elaeadendrol1 capeme Eckl. & Zeyh.: 127 ( 18 34/5); Graham: (1996b) argue that Elaeadendron Jacq. ( 1782) is validly pub­ 3835 (1841 ); Sand.: 468 (1860); Bak. f.: 45 (19 11 ); Eyles: 404 lished as a plate with an analysis and th at it predates both Murray (1916); Marioth: 154 ( 1925); Loes.: 173 (1942a); N. Robson: ( 1784) and Jacquin f. (1787). 386 (1966). Type: Cape, ' Sylvis Sitsikamma & Krakakamma Robson (1965, 1966) treated E. zeyheri (Cassine Cf·oeca allcl. (Georg, Uitenhage). Flor Jul. Fruct. Febr.', Ecklan & Zeyher S .I1. non Thunb.) and E. transvaalense under the genus Crocoxylon. (SAM!, lecto, here designated, S!, TCD! , Z!). The latter genus is supported by a di stinctive pollen type with a rugulose-reticulate exine structure (Archer & Van Wyk 1992). £Ioeodendron papiJIoslIm Hochst. in Krauss: 305 [( 1844) reprinted: 42 (1846)]; Ettingshausen: 58 ( 1857). Cassine papil­ However, several tropical African and extra-African species are lasa (Hochst.) Kuntze: 11 4 ( 189 1); Davison: 334 (1927) pro intermediate (in pollen structure and fl ower morphology) parle excl. syn. C lacinulata Loes.; Palmer & Pi tman: 1321 between Elaeodendron and Robson's concept of CrocoxylOI1. ( 1973); Vo n Bre itenbach: 228 ( 1974); Coates Palgrave: 511 The key provided below gives some indication of the considera­ ( 1977); Coates Palgrave el 01.: 64 (1985); Pooley: 278 ( 1993). ble variation in the number of fl oral parts in the African members Type: 'sylvis prope Natal, Julio 1839', Krallss 270 (TUB, holo., of Elaeodendron. At thi s stage it is not considered feasible to BM!, K!, MO!, sr, TCD!). recognize Crocoxylon as a segregate genus or even any supraspe­ cHic taxa. Incidentally, the name Crocoxylon is to be regarded as Olea sp. Mellis: 312 ( 1875). a synonym of Elaeadendran, typified by E crocellm (Thunb.) leones: Graham: t. 3835 ( 184 1); Von Breitenbach: 229 (1974); DC. (not Crocoxylon cracea (Thunb.) N. Robson). Coates Palgrave el al.: 64 (1985); Archer & Cond y: t. 21 12 (1995). Key to the African species of Elaeodendron la Fl owers with ovary heteromerous wi th other floral whorls.
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