s. Mc. J. Bot. 1999,65(2) 177-181 177

Systematic studies in subfamily Celastroideae () in southern : reinstatement of the genus Gymno~poria

Marie Jordaan* and A.E . van Wyk 1 'National Botanical Institute, Private Bag X101, Pretoria, 0001 Republic of South Africa 'H G.W.J. Schweickerdt Herbarium, Department of Botany, University of Pretoria, Pretoria, 0002 Republic of South Africa

Received 28 .h/~~' 199~ : revised J I January JYfJi)

Gymnosporia (Wight & Arn.) Hook. r. nom. Ganserv., IS recognised as a genus on its own, comprising all the sp iny members currentty included in the genus Molina 5.1. Diagnostic characters of Gymnopona include the presence of brachyblasts and spines, in fascicles on older branches or alternate on young ones, inflorescence a monochasium, subdichasium or dichasium, and mostly unisexual. is an Old World genus comprising about 80 species and subspecies, occurring in most of Africa, and adjacent islands, southern Spain, the near . , , , extending to the Far East, Malesia, Papua New Guinea, the Philippines, , Queensland (Australia) and the Po lynesian Islands. An amplified generic description is provided. Differences and similarities between the spiny genera of Celastroideae {PutterUckia End!', Gloveria M.Jordaan, Gymnosporia (Wight & Am.) Hook. f. and Maya Griseb.J are tabulated.

Keywords: Celastraceae, Gloveria, Gymnosporia, Maytenus, Moya, Putterlickia, southern Africa, .

.. To whom correspondence should be addressed , (E-mail: [email protected]) .

Introduction seed), on the basis of ovary and secd characters. [n section t::IICC­ The Celastraceae (including Hippocrateaceae) is a family of lastrus the ovary is free from the disc and the seed completely trees, or climbers, compri sing about 85 genera (Brummitt enclosed by a fleshy ari!. This section corresponds to th e genus 1992). The family is widespread, but concentrated in subtropical Ce/aslrus as circumscribed today (Ding Hou 1(55). It occurs in and tropical regions. Subfamily Celastroideae (Loesener 1892, eastern Asia, Oceania, both Americas and Madagascar. between 1942; Scholz 1964; Takhtajan 1980), a natural assemblage of 40° Sand 47° N. In section GYlJll1ospor;a, which corresponds to taxa, is characterised by dry loculic idall y dehiscent capsules and Kunth's ( 1825) unnamed genus. the ovmy is half immersed in seeds with an arit. [n recent years the African, Madagascan, Asi­ the disc, with the seed completely or incomplete[y enclosed by atic and A ustralian members of subfamily Celastroideae (Celas­ an ari!. traceac) have been classified under Calha Forssk. ex Scop., Don ( 1832) delineated the order Celastrineae tribe Euonymeae Ce/aslrus L., Meisn., A.faytelllls Molina, A10ya as with arillatc seeds and simple leaves and placed the spiny Griseb., Polyc:ordia Juss., Pterocelaslrus Meisn. and Pulff!r/ickia African, European and Asiatic mcmbers of Celaslrlls in the genus Endl. Many genera huve a limited distribution range, e.g. Ce/as­ Calha Forssk. , a genlls with hermaphroditic fl owers. He recog­ II"lIS occurs in Eurasia, Madagascar. Malesia, Australia and North ni sed Ce/astl"Jls KUllth ( 1825) I/UII L. (excluding C scam/ells L.) as Ameri ca but is absent from the African continent; Denilamia is a genus with dioecious plants and ovaries which are 2-4-10cular confined to Australia; Polycardia to Madagascar; lv/oya to South with two ovules per locule. DOll (1832) also recognised ..\.fa)'If!lllIS America; Pteroce/aslrllS and Putter/ickia to southern A fr ica and as an exclusively Ameri can genus, characterised by polygamous Catha to Africa and SW Arabia. l\laytenus s./., consisting ofl1on­ flowers, one ovule per locule and the lack of spines. Endlicher spiny and spiny shrubs or trees, is currently considered to be the (1840, 1841) elaborated on Dan's (1832) description of tribe most widespread genus in the subfamily, occurring in both the Euonymeae, including all members with loculicidall y dehiscent Old and New World. The purpose of the present paper is to capsules. This resulted in the recognition of two more genera, review the development of generic concepts in the Celastroideae namely Pferoce/aslrus Meisner (1837) for members with protu­ briefly and to present arguments for the reinstatement o f the berances on the capsule and the newly described genus Pllfwr/ic­ genus Gymnospuria. kia for species with more than two ovul es per [ocu [e. Walpcrs (1842) maintained the genus PII/(er/ickia. but still followed Don Historical review ( 1832) in placing the spiny Asiati c species of Celasfl"lls under Members of the Old and New World genus Maylf!nlfs and Old Catha. However, he returned Don's European and Africllll mem­ World genera Gymno.\poria and PlIllerlickia are all placed in bers ofCat/Ja to Celaslms Kunth non L. Ce lastrus L. by 1110st early allthors, e.g. Linnaeus (1753, 1754), Hooker (1862) elevated section Gyllll1osporia Wight & Arn. to Miller (1768), Linn,eus filius ( 1781), Lamarck (1785), Thun­ generic rank. This genus is geographically restricted to the Old berg (1794,1823), Willdenow ( 1798), Burchell ( 18 24), De Can­ World. The generic name Gymllosporia has subsequently been dolle (1825), Ecklon & Zeyher ( 1834- 1835), Meyer (1843), conserved against the following /lom;na rejiciellcia (Farr f! t al. Schl echtend al (1846), Sonder (1860), Oliver (1868), Kuntze 1979): Calha Forssk. ex Schreb. (1789); Ellcenl,-"s C. Presl (1891, 1898) and Wood (1908). Kunth (1825), however, pointed (1844) ex End!. (1850); Polyacrllllhlls C. Presl (1844). out that some Ctdaslrlls species have a combination of peculiar Distinguishing characters lI sed by Hooker ( 1862) to separate characters and that they may well constitute a new genus, Gyn11losporia fr om MaYlemfs are as fo llows: "I ocule with two although he did not formally propose a name for it. ovules; plants often spi ny: inflorescences cymose Reporting on the Indian species of Ce/astrus, Wight and Gymnosporia. Locu[e with 1 (rarely 2) ovules; plants without Arnott (1834) divided Celastrlls into two sections, Eucelaslrlls spines; flowers sol itary, or inflorescences fllsci cu late or cymose and Gymnosporia (from the Greek gynlnOS :::: naked, and spora:::: - .Maytelllls'. He gave the geographical distribution of 17K S. Afr. J. Bot. 1999. 65(2)

C)'IIIIIU.I'{Jo/';a as Spain. Africa, Asia and Australia, and that of African Celastraceae, was by Davison (1927). [n recent years the .I/uylellils as tropical America and Australia. These generic con­ vi ew that G)'Illl1o,';porio is congeneri c with :v/aYTc!I1f1s has been cept s were followed by subsequent workers such as Szyszylow­ generally uncritically accepted and followed by botanists work­ iez ( 1888), Sim ( 1907), Loesener (1893, 1896 ), Engler and Gilg ing on the 1\ frican flora, fo r exnmple Ex ell and Mendonc;a ( J954) (1 91 2). Schonlnnd ( 1919), Marloth (1925), Davison (1927) and (Angola), Keay and Bl akelock (1958 ) (Flora of West Tropical Adamson (1950). Africa reg ion ), Wilczek (1960) (Democratic Republi c of the In his monumen lal worldwide revision o f the Celastraceae, Congo, Rwanda and Burundi), Marais (1960) (Flora of southern LO l.! sencr (1892. 1942) was the first to establish subfamilies and Africa region), Ding HOll (1962) (Fl ora malesiana region), Rob­ tribes in the family . He re ferred all the Old World spiny mem­ son ( 1966) (F lo ra zambesiaca region), Sousa (1968) (Gui nca­ bers of Celastroidcae with short and long shoots and fasciculate Bissau), Robson and Sousa (1969) (Mozambique), Guillarmod leaves to subfamily Cel.:lstroideae, tribe Eucelastreae. Loesener (197 1) (Lesotho), Ross ( 1972) (KwaZulu-Natal), Villicrs (1975) { 1()42) recognised GYI1lJ1 usporia as an Old World genus, '\fayte­ (Cameroon), Compton (1976) (Swaziland), Troupin (1982) l iltS as a genus occurring in both the O ld and New World and (Rwanda), Bond and Goldblatt ( 1984) (Cape Floristic Region), ('c/a.vlrlls as be ing absent from Africa. A ll owing for the limited Jessup ( 1984) (Australia), Sebsebc (1985, 1989) (Sudan, Ethio­ knowledge on Illany genera (hat existed at that time, the classifi­ pia, Eritrel\, Djibouti, Somalia, Saudi Arabia, Yemen and Oman) catio n proposed by Loe::;ener ( 1942) is a su itable starting point and Robson (1994) (Uganda, Kenya and Tanzania). for ~xprcssing the broad outlines of generic relationships within Marais ( 1960 ) and Robson (1965, 1966), who were mainly the family . followed in the two most recent lists of African species E.xell and Mendon,a (1952) and Exell (1953) argued that (A rn old & De Wet 1993; Lebrun & Stork 1992), had a very Loesencr's (1942) separ(lti on of Gymnosporia from IHaylenllS, broad species concept. Only J 2 spiny members of ,Haylenlls are on the basis of the presence of either thorns or the infl orescence recognised in the Flora of southern Africa region (Arnold & De borne on short shoots ill C;Yl1ll1ojporia, appeared artificial. They Wet 1993) and 30 in the Flora of tropical Africa, south of the proposed an amalgamati on o f the two genera into a widely ci r­ Sahara (Lebrun & Stork 1992). cumscribed ,\lay/emls ,d. Brenan (1953) shared the latter view, and added that ,\/aytelllls had previously been segregated from Reinstatement of Gymnosporia (J)'/J/I/Ospori{/ by what he considered rather vague characters, A recent comparative study on the spiny (spines regarded as such as the prevalence of uniovulate locules and bilocular ova­ modified stems) members o f the Celastroideae (Celastraceae) on ries, whereas species with biovulate locules and with spines were a world wide scale (Jordaan 1995; Jordaan in prep.) provided placed in G)!llIIlOs/}()ria. abundant evidence, both macroscopic and microscopic, for the Ding H Oll (1955) examined a very large number of herbarium segregati on of Maylelll/.\· .d . into two natural, more homogene­ specimens of Ce/astrll.\', GYI11IJOspOl';a and May/eulls and com­ ous genera. We consequently propose that Gyml1osporia be rein­ pared many characters. He concl uded that these three genera stated as a genus, comprising all the spiny members currently were distinct, and proposed their recognition on the basis of a placed in AfayteJ1l1s .d. . thus following the view of Loesener combination of several characters, not a single character on its (1942) and Ding Hou ( 1955). Maylelllls is provisionally retained OW I1 . Exell (195 3), B lakelock ( 1954) and Marais ( 1960), how­ in a 'stri ct sense' to accommodate all the nonspiny members. It ever, claimed that there seemed to be no single character state, remains, however, a heterogeneous assemblage, perhaps best fur­ nor any combination of characters, constant enough to justify the ther subdivided into a number of segregate genera. Morphologi­ retention of G.1'l1Il1os{Jo/'ia. They also agreed that CetC/slms was a cal si m ilarities and differences between the spin y genera of the good genus. and that it was absent from the A frican continent. Celastroideae are sUlllmari sed in Table I. Cetasfrlls is characterised by a scandent habit, an ovary free from Maytelllls s. sIr. d iffers from Ciymnusporia in the absence of the di sc. a persistent ccntral axis of the capsule, and a frequently brachybJasts and spines, alternate leaves w hi ch are never in true racemose or paniculate inflorescence. fascicles, flowers always bi sexual, and inflorescences which are Di ng Hou (1962) changed his mind and stated that the differ­ solitary, fasc iculate (reduced cymes), racemose (reduced pani­ ences between GYlJJl1osporia and iHaylenlls as tabulated in his cles) or variously cymose (though apparently never dichasial paper of 1955 cannot be maintained, especially not in the African cymes). Plants of Maytel1l1S s. stl', are mainly small to large forest flora, as was also concluded by Marais (1960). Ding Hou ( 1962) trees, only a few species remaining low shrubs w hen growing retained onl y ('e/aSlrlfS and Maylel1llS, distingui shed by a combi­ near the coast or in arid reg ions, whereas only a few members of nation of three characters: habit, degree of ad nation of ovary and Gymnosporia are med ium -sized trees, mostly not exceeding 10 disc. and structure. m. In AfayteJlJ/s s. Sir. the seeds are completely or almost com­ The first major taxonomic treatment of spiny Celastroideae pletely enclosed by an aril , while different states occur in Gym­ \Vas by Thunberg in hi s Flora capel1sis (1823). Ecklol1 & Zeyher l1ospo";a: from completely enclosed to seeds with the ari l ( 1834- 1835) were the 'first to note the extreme variability of Lin­ reduced to a basal rim . nacus's (1753) C hIlxf/'olius. Don (1832) placed all the species Gymnosporia has more characters in common with the spiny under ( 'ol/W; Presl (J 844) described two new genera, namely genus PlIlterlickia than with the members of l10n spiny Maylc!l1l1s £ IICcnfrll.\· and Pn/yacafllhll.'i, both now included in Gyll1nmpo­ s SIr. Al though GY11Jnosporia is morphologicall y diverse. it is ria, established by J.D. Hooker (1862). The second main revision easily recognised by a combination of five characters: (i) the was by Sonder (1860) in Flora capensis, a work in which all presence of spines and brachyblasts; (ji) functionally unisexual known spiny Celastroideae were treated under Ce/astrus. flowers; (iii ) a dichasial inflorescence type; (iv) bi-ovulate and Szyszylowicz ( 1888) was the first to place all the species under (v) 2-4-locular ovaries. Plant material of Gymnosporia has been GYlllllmjJona and Pltllel'fickia, the latter described by Endlicher studied from all parts of its range and it is here reinstated as a dis­ (1840). Loesener' s publications on the Celastraceae (1892-1942) tinct genus. arc very important as he attempted to include all known taxa in In habit, , inflorescence and fruit characters Gymno.\poria his monographs of the family. He was the first to transfer all bio­ is very s imilar to Plillerlickia, a mainly southern African genus vulate spiny members to Gymnosporia. (Table I). These two genera are disti nguished mainly on the The Illost recent comprehensive taxonomic revision of the basis of floral sexuality and the number of oyules per locul e. Cdastroideae for southern A frica, and in fact a ll the southern Pulterlickia has bisexual fl owers and usually 6- 12 ovules per S. Ali'. J. Bot. 1999, 65(2) 179

Table 1 Summary of selected morphological similarities and differences between the spiny genera of Celastroideae

Clu.lr;Jctcr PUfferiickia Gloveria GY/JIJ/osporia il"foya

Africa. Madagascar. S Spain. Arabia. Pakistan. Indomale:-;ia. Phillipines. mainly castem parts ofS Little Knroo and Australia, Papua New Guinea and Argentina, Bolivia, Para· Geographical distribution A II-lea and S Mozambique Namaqualand the Polynesian islands guay and Umguay

suffrutices. sllnlbs or Illcdium~s i zed Habit shmbs Of cl imbers shrubs or smnlllrccs Ire!.':s shrubs or small trees

in axils of leaves or bradlY­ in axils of leaves or in axils of leaves or brachyblasts. or terminating lateral Spines blasts brachyblasls tenninating brachyblasL<; branches

Number of nodes Otl a spine (modi lied stems) one more than one usually one none

l:lltire. or with few teeth in dis- entire or with lew teeth in distal or Lt:armargim; ta l hall' entire both halves entire or with lew teeth

lamina surface glabrous glabrous glabrous or puberuJous glabrcscent or pubescent

Intlort:scencc di chasium dichasiulll monocbasium Uf dichasi l1tll fasciculate or solitary

Sexuality offlowcrs bisexual bisexual mainly unisexual unisexual

OV31)' 3-loeular 3-loeular (2)3(4)-loeular 2(3)-locular Locult: 6- 12- ovulate 3--6-ovulatt: 2-t)\'ulatc 4-ovulate smooth, glabrou s, rugose. vei ned or smooth, glabrescent or Capsuk surface smooth and glabrous smooth and glabrous pub!.':rulous pubescent

Ari l colour orange pinkish orange. pinkish. yellow or white red

Percentage orsecd covered by ± 100%, 60%. 30% or only a rim at aril 100% ± 100% the base ± 80%

locule. Glol'eria, a new monotypic genus of spiny Celastroideae genera. Other differences between Moya and the spiny members from southern A frica has been described recently (Jordaan & of Celastroideae are mentioned in Table 1. Van Wyk 1998). It is closely alli ed to Gymnosporia from which Members of GY"111mporia are adapted to both summer and it is disti nguished by always having bisexual flowers and 3-6 winter rainfall climates, as well as semi-desert regions, but are ovules per locu le. Glol'eria is simi lar to Putlerlickia in having absent from the Desert Biome (Rutherford & Westfall 1994). bisexual flowers, bur deviates by its fewer ovules per tocule, brachyblasts main ly on the spines and more than one node per spine (Table I). CYllll1ospol';a is an Old World genus comprising about 80 spe­ cies and subspecies (Jordaan 1999), occurring in most of Africa, Madagascar and adjacent islands, southern Spain, the near Mid­ dle East, Pakistan, India, Sri Lanka, extending to the Far East, Malesia, Papua New Guinea, the Philippines. Taiwan, Queens­ land (Australia) and Polynes ian Islands (Figure I). Members of Gymllosporia arc spiny, much-branched, woody, evergreen per­ ennials, varying from dwarf stoloniferous shrubtets, to medium~ sized shrubs or small trees, usuall y not taller than 10 111 . Plants are ITIostly dioecious and, therefore, are outbreeders. The flowers are dull in colour (whitish, cream, yellow or rarely red), usually with a disagreeable odour attracting mainly flies. The polymor­ phic C. sel1egalensis (Lam.) Loes. is the most widely distributed species, ranging from southern Africa northwards to south ern Spain and eastwards as far as India. The nonspiny members of Maytenlls are distributed mainly in the eastern parts of Africa, Madagascar, Malesia, Australia, and South and Central America. Certain spiny members ofCelastroi­ deae (tour species) fr0111 the arid parts of (Argen~ tina and Bolivia) are placed in the genus A10ya by Grisebach (1874) and l oesener (1942), but in Maytenlls by Lourteig and ,o~----4------+----~------+------+----~------~--I,o O' Doneli (1955). In A10ya there are only two ovu les per locul e as in GYlllllosporia and May tenus, but it can be distinguished by the o 30 120 ISO 180 aril which is visible at flowering stage (Loesener 1942), whereas " the mil is visible only at the fruiting stage in the other two Figure Geographica! range of the genus vymnusporw. 180 s. Afr. J. Bot. 1999.65(2)

Species have been recorded from all three subkingdoms of the subentire. Petals mostly whitt:. Crl:alll. ~"dIO\\I. pale green or occa­ Palaeotropical K ingdom (Good 1974) and in all the main sionally pink or red. im hricate ill hud. margin ci liate or lacilliatc. phytochoria in Afri ca (White 1983), but the group is noticeably rarely entire. spreading ()f rctlexed. DISC ill trastarn ina1. convex or absent from higher altitudes in the Afromontane Region. Habi­ concave. 5- 10-l obed. green. yellow or rlo! d. Stamens atlachlo!d to bast: tats include coastal rocky outcrops and sand dunes, karoe scrub, of disc: anthers introrst!. with 2 thecae. ddliscing longitudinally. valley bu shveld , open savanna, rocky or grassy hillsides as well GYlloechlm syncarpous: ovary with quarter to half immersed in disc. as the margins and understorey of afromontane forests. The (2)3(4)-I ocular with 2(3) erect. collateral ovules in each locuk: style group is poorly represented in fynbos. terc:te. short or elongate: stigma (2)3(4)-lobcd. Fruit i.I dry capsule Morley an d Toelken (1983), in Flowering planls in Australia, opening longitudinall y. loculi cidal and septifragal. Cap.mles vari­ separate Maylenlls and Gymnosporia on account of their differ­ ously coloured, globose. obconic-trigollollS. triquetrolls. pyriform or ent inflorescence types and state that Gyn1l1ojporia is represented conic-pyramidal. smooth to rugose. punctate or veined: pl.:ricarp semi fleshy. chartaceolls. coriaceous. \Voody or ridged. Seeds 1-4. by on ly one species, without mentioning a name, presumably C. glossy. reddish brown or black; aril Ikshy or thin. partially \0 com­ mOIl/ana (Roth ex Roem. & Schult. ) Benth. Jessup (1984), in a pletely covering the seed. or reduced to a rim at base of seed. revision of the Celastraceae for th e Flora of Allstralia, considers pinkish. \V hite. yellow. or orange. Gymnosporia mOlltalla a synonym of A,foylell/ls emorgillota (Willd.) Ding Hou, thus fo ll owing Ding Hou ( 1962). GymllD.lpo­ Acknowledgements ria montano, the type species of Gynmosporia, was described We are most grateful to Emsic du Plessis fo r critically reading from eastern India. In recent years it has been considered a syno­ and improving the manuscript. nym ofG. senega/ensis by, for example, Sebsebe ( 1985). The plants occurring in Australia are usually without spines, although References brachyblasts are present on older branches. 1\1. emargillata has cymose inflorescences, which differentiate it from the other spe­ ADAMSON, R.S. 1950. Celaslraceae Lindl. In : Flora of the Cape Penin­ sula, eds R.S . Adamson & T.M. Salter. pp. 564-568.. luta. Cape cies of Maylellus s. sIr. It is stated by Bentham (1863) that the Town. Australian specimens of G. montana are often w ithout spines, as ARNO LD. T.H. & DE WET. B.C. (cds). 1993. Celastraceae. R.H. is the case in India as well as in some tropical A frican specimens Archcr. In : Plants of sou thern Africa: numcs and distribution. ,\lem. of G. senega/ensis. The leaves of the A ustra lian plants are not bal. SlIrv. S. Al;·. 62: -t80-481. coriaceous and glaucous like C. senega/eJ1sis or G. emarginata. BENTHAM. G. 1863. Celastraceae. In: Flora mlstraliensis: a de5cription Furthermore, the Australian plants have 3-locular ovaries and an of the plants of the Australian territory. Vol. I. Reeve. London. aril reduced to the base, whereas G. senega/ensis has 2-locular BLAKELOCK. R.A. 11)54. Nomina gcncrica conservanda rcposila. ovaries and an ariI which partially covers the seed and G. emar­ Taxon 3: 196-11)7. ginala has a 3-locular ovary. This is clearly not the sam e taxon as BOND, P. & GO LD BLATT. P. 1984. Celastraceae. Plants of the Cape C. senegalensis nor is it con specific with the Indian C. emargin­ Flora: a descripti ve catalogue. J. S Afr BOI. Suppl. 13 : 224. ala. At this stage the m aterial from Malesia, Papua New Guinea, BRENAN, J.P.M . 1953. Plants collected by the Vernay Nyassaland the Philippines, Taiwan, A ustralia and the Polynesian Islands is Expedition of 1946. CeJastraceae. Mem. A'. Y. bot. Gdn 8: 236- 238. best pl aced under G. vWel/sis (A.Gray) Seem. BRUMMITT, R.K. (cd.) 1992. Vascul ar plant families and genera. Royal Botanic Gardens. Kcw. Taxonomy BURCHELL. W.J. 1824. Travels in the interior of southern Ati-ica, Vol. Amplified genus description 2. Longman. London. COMPTON. R.H. 1976. Celastraceac. The Flora of S\Vaziland. J. S. Afr. Gymnosporia (Wighl & Am.) Hook.! in Bentham & Hooker, 801. Suppl. I L 334 . Genera plantarum I: 365 ( 1862) emend. Loes.: 87 (1942) (l1 omel1 DA VISON, J.D. 1927. Cc::la." tr.lceae R.Br. Bori1(llia 2: 289-346. cOl1servandum). Type species: G. montana (Roth ex Roem. & DE CAN DOLLE. A. P. 1825. Ce lastrincae. In: Prod romus systemati s Schult.) Benth.: 400 ( 1863) (typ. cons.). natural is regni vegetabili5. Vol. 2. pp. 2-18. Treuttel & WUrtz. Paris. Ce/aslrus L. Section GYIII/l osporia Wight & Am.: 159 ( 1834). DING HOU. 1955. A re vision of the genus Celastrus. Alln. Aftss. bOI Gymnosporia Subgenus 1. Eugymnosporia Loes. Section 1. Gd1l42: 2 15-302. spil10sae Loes.: 207 ( 1892). DING HOU. 1962. Cclastr

lIominUIll gencricorutn, Vol. 2, p. 760 & Vol. 3, p. 1466. Dohn, Schel­ Excll. A. Fcmandcs & I\. Wi ld, Vol. 2(2), pp. 355-4 18. Crown lema & Holkema, Utrecht. Agents for Oversea Govt.!rnmellt .tllI..! Administrations, Lonuon. GOOD, R. 1974. The geography of the tlmvering plants, edn 4. Long­ ROBSON. N.K.B. & SOUSA. E.P. 1%9. Celastraceac. In; Flora de man, London. Mo!;ambique, Vo!' 48( I): 1-66. Centro de bol[mica. Lisbon. GRlSEBACH, A.RR. 1874. Abh. Konig!. Ges. lViss. Gal/ingen 19: Ill. ROBSON. N.K.B. 1994 Celaslruce.le. ,\!aytel1us. In : Flora of Tropical HOOKER., J.D. 1862. Cclaslrincae. In: Genera plantarum . eds G. East Africa. Celaslracr.!ne. cd. R.M. Po lhil!. pp.I-2 I . Balkcllla. Rolkr­ Bentham & J. D. Hooker, Vol. I, pp. 357-371. Reeve, London. dam. JACOT GUILLARMOD. A. 1971. Celastrnccac. In: r:lora of Lesotho, ROSS. J.H. 1972. Celaslraccac. Thr.! Flora orNata!' Mem. bot. SlIr\,. S pp. 207-208. J. Cramer. Lehre. A/r. 39: 231 - 232. JESSUP. LW. 1984. CcJastraceae. In: Flora of Auslralin. pp.150-164. RUTHERFORD. M.e. & WESTFALL. R.H . 1l}94. Biomes nrsollthcm Australian Govenncnt Publishing Service, Canberra. Africa; an objective categorizatiull. ,\/l'III. hot. SlIrv. S. I~fi' 63: 1- 94. JORDAAN, M. 1995. A taxonomic revision of the spi ny rncmbl.!rs of SCHLECHTENDAL. D.F.L VON. 18-l6. Vergleichungcn Jer von Eck· subfamily Cdastroidcae (Celastraccae) in soulhem Africa. M.Sc. the­ lOll und Zeyher und von Dregc gr.!sammdtcll sGJalrikanischen Pnan­ sis. University of Pretoria. Pretoria. zen mit den Exclllplarcn von Zc)'her's IICLIesten Sammhlllgcn. JORDAAN. M. & VAN WYK , A.E. 1998. Systematic stud ies in su b­ Celastrineae. Linl1aea 19: 623- 624. family Celastroideac (Celastraceae) in southern Africa: C/ul'Cria. a SCHOLZ, H. 1964. CelastroiJcilc. In; A. Eng,lt::r's Syllabus dl.!r Pl1an­ new monotypic genus. S. A.fr. J. Bal. 64: 299-301. zenfamilien, cd. H. Melchior. 12th rev. cdn. Vol. 2. Domtraeger. Bel'· KEAY, R.W.J. & BLAKELOCK, R.A. 1958. Cclastraceat':.ln: f lora of I in- N i kolassee. West Tropical Africa, 2nd cdn, Vol. 1,2, pp. 623-634. Crovm Agents, SC HONLAND, S. 1919. Phanerogamic 110ra o l'the divisions of Uitcn­ london. hage and Port Elizabeth. Mem . hOI. Surv. S Afr. I: 1- 11 8. KUNTH, CS, 1825. Celastrineae R.Br. In: F.W.H.A. Humboldt, AJ.A. SCHREBER, J.C.D. VON. 1789. In : Caroli a Linne. Genera plantarul11. Bonpland & CS. Kunth, Nova genera, Vol. 7, pp. 64- 79. Paris. KUNTZE. CE.O. 1891. Celastraceae. In: Rcy isio generum planlarum, edn 8(a). Vol. L Fnmkfurt am Main. SEBSEBE. D. 1985. TIle genus Maytcnus (Celaslraccac) in NE tropical Vo L L pp . 113-117. AI1h ur Felix. Leipzig. KUNTZE. ('E.O. 1898. Celaslraccae. In; Revisio gent':rum piantanun. Africa and tropical Arabia. ~:Ymb . hot. IIp.wl 15: 81 - 90. Vol. 3, pp. 37- 38. Arthur Felix. Leipzig. SEBSEBE, D. 1989. Celnsl r acea~. Mayll'lIl1s. In : r: lora of Ethiopia. ~ds LAMARCK, J.B.A. P.M. DE. 1785. Encyclopedie methodiquc. Bota­ L Hedberg & S. EdwarJs. Vol. 3. pp. 33 1-338. Addis Abab~ Univer­ nique. Vol. I, p. 66 1. Puris, sity, Eth iopia. LEBRUN, J.-P. & STORK. A.L 1992. Celastraceae. Enumeration des SIM, T.R. 1907. Celnslrineae. In: The t(lI'eSlS and forest nora or the pIal/res djleurs d'Afriqlle rropicafe 2: 147-150. Geneve. Colony of the Capt! of Good Hope. pp. 18 1- 19 I. Taylor & HcnJer­ L1NNAEUS, C 1753. Species piantarum, cdn I. Laun.::ntius Salvius. son, Aberdeen, Scotland. Stockholm. SONDER.O.W. 1860. Celastrineae. In: flora capensis. cds W. H. Har­ LINNAEUS, C 1754. Genera plantarum, edn 5. Laurentius Salvi us. vey & 0, W. Sonder, Vol. I. pp. 452-460. I-lodges & Smith. Dublin . Stockholm. SOUSA. E.P. 1968. Rcvisao das Cdastraceae da Guine Portllgllcsa. LlNNAEUS. C. (filius). 1781. Supplementum plantarutn. Dmnswick, Garcia de Orla 16.2: 21 1- 218. lisboa. Upsala. SZYSZYLOWICZ, l.R. VON. 1888. Polypetalae Discillorae Relunanni­ LOESENER. L.ET. 1892. Cclastraceac. Celastroideae-Eucelastreae. In: anae. pp. 35-37. (Publishd in Ro=pr Sprmv. PO.l'ied=en Wyds Mat.­ Die natiirlichen Pflanzenfamilien. cds A. Engler & K. Pranl!. Vol. Przy", Akad. Umiejeln. UniVerSI/Olis Jage//olJicae (Cracow) 18; 1- 75.) 3(5), pp. 203-21 I. Engelmann, Leipzig. TAKHTAJAN. A.L. 198 0. Outlinc or the c1assi li cation uf !lowcring LOESENER, LE.T. 1893. CelaSlnlceae africanae I. BOI. Jb. 17: 54 1- plants (Magnoliophyta). /J0i. Rev. 46: 225- 359. 546. THUNBERG. C. P. 179 .. . Prodromus plantarulll capcllsiulll ... Part I. LOESENER. LLT. 1896. Celastraceae. Bull. Herb. Boissier 4: 429- Edman, Upsala. 430. THUNBERG. c. r. 1823. Flora capcllsis. ed. J.A. Schultes, 2nd r.!dn. J.(i LOESENER. L.E T . 194 2. Cclastraceae. In : Die natOrlichen Ptlanzen­ Cottae, Stuttgart . familicn, eds A. Engler & K. Prantl, Band 2, 20b, pp. 87-197. TROUP1N, G.M.D.J. 1982. Ce l aslrac~a~. tn: Flore des plantcs ligncuscs Duncker & Humblot, Berlin. du Rwanda. Musee Royal de L' A li'i quc Centrale. Sciences A. & LOURTEIG, O'DONELL, C.A. 1955. Las Celast"ceas de Argen­ Economiques n. 12. pp. 198- 205. T ~rvuren. Belgique. tina y Chile. Aire:,· 1(2): 181-233. Narura, Buenos VILLIERS, J.-F. 1975. Ceiastracces. In: Flore du Cameroun, eJs A. MARAIS, W. 1960. An enumeration orthe Maylenus species of south ­ Aubrevilte & l -F. Leroy. Vol. 19. pp. 30-32. Museum National em Africa. BOlhalia 7: 381 - 386. D'Histoire Naturel lc. Paris. MARLOTH, H.W. R. 1925. The fl ora of South Africa with synoptical WALPERS, W.G. IR42 . Cdas t rin~ac . In : Rcpertorium botanices sys­ table~ o rthe genera of the higher plants, Vol. 2. Darter, Cape Town. tem81icae. Vol. I, pp. 532- 534. I Ioflllcistcr. Leipzig. MEISNER, C.F. 1837. Pl,mtamm vascularium genera. Wcidmanl1ia. WHITE, f. 1983. The vegetation of Alrit: