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IAWA Journal, Vol. 14 (1), 1993: 35-53 BARK STRUCTURE AND IAWA Journal, Vol. 14 (1), 1993: 35-53 BARK STRUCTURE AND INTERGENERIC RELATIONSIllPS 01<' SOME SOUTHERN AFRICAN CASSINOIDEAE (CELASTRACEAE) by Robert H. Archer1 andAbraham E. van Wyk H.G.W.J. Schweickerdt Herbarium, Department of Botany, University of Pretoria, Pretoria, 0002 Republic of South Africa Summary At present Cassine in southern Africa Wyk 1992). Established by Loesener (1892), is treated in a wide sense (s.I.), including this subfamily accommodates those species amongst others Allocassine p. p., Cassine of Celastraceae with indehiscent fruits, usu­ s. str., Crocoxylon, Elaeodendron, Lauridia, ally drupes or berries. and Mystroxylon. A comparative anatomical There is considerable diversity of opinion study was made of mature bark representing regarding the delimitation and recognition of 16 southern African species of Cassine s.l., genera among the species of Cassine s.1. and the monotypic Allocassine, Hartogiella Various authors (Loesener 1894, 1942; Rob­ and Maurocenia (all members of the subfam­ son 1965, 1966) advocate smaller, more ho­ ily Cassinoideae). Six bark types are distin­ mogeneous genera, while Davison (1927), guished on the basis of the type of scleren­ Hou (1962), Codd (1966), Kostermans chymatous elements in the secondary phloem; (1986) and current herbarium practice at the presence or absence of styloid crystals, e1as­ National Herbarium, Pretoria (PRE) (Gibbs tic threads, and sclerified phelloderm; strati­ Russell et al. 1987), prefer to treat Cassine in fied homogeneous phellem; and degree of a wide sense, so as to include Allocassine rhytidome development. These correlate to a p.p., viz. A. tetragona, Crocoxylon, Elaeo­ considerable extent with the generic subdivi­ dendron, Lauridia, and Mystroxylon. For the sion of Cassine s.l. proposed by Loesener last two decades the treatment of Cassine in (1942) and Robson (1965). On the basis of a broad sense has received strong support bark anatomy and other evidence, it is pro­ worldwide (e.g. Dyer 1975; Mabberley 1987; posed that the circumscription of Cassine be Jessup 1984). The accumulation of new evi­ restricted to include only the southern African dence from morphology and anatomy, as species C. peragua and C. parvifolia, and pos­ weH as other sources, might accomplish a sibly Hartogiella. Crocoxylon, Elaeodendron, better understanding of generic limits in the Lauridia and Mystroxylon should be reinstat­ Celastraceae. ed or maintained, although with some modifi­ Comparative bark anatomical studies on cation of the originally defined generic limits. members of the Celastraceae are scanty. Roth (1973, 1981) describes the bark anatomy of Key words: systematic bark anatomy, taxon- woody neotropical species of Celastraceae. omy, Cassine s.l., Cassinoideae, Celas­ Zahur (1959), in a comprehensive survey of traceae. secondary phloem structure in dicotyledons, studied a few species of Celastraceae, but the Introduction anatomical description of this family was ap­ This study forms part of a multidiscipli­ parently accidentally omitted from this publi­ nary approach towards a taxonomic revision cation. Observations on celastraceous bark of the southern African members of subfam­ structure have also been provided by Weber ily Cassinoideae (Archer 1990; Archer & Van (1913), Thorenaar (1926) and Chung & Park 1) Present address: National Botanical Institute, Private Bag X101, Pretoria, 0001 Republic of South Africa. Downloaded from Brill.com10/04/2021 12:50:06AM via free access 36 IAWA Journal, Vol. 14 (1), 1993 (1975). However, no information on the ana­ Bark sampIes were removed at approxi­ tomieal structure of the bark of Cassine could mately 0.5 m height from vertical boles not be found in the literature. less than 100 mm in diameter, and fixed in As an aid towards the classification of FAA (Johansen 1940). Standr.rd procedures woody taxa, the potential taxonomie signifi­ for wood anatomy were used to prepare trans­ cance of bark anatomy has been neglected. verse, radial and tangential seetions, 15-20 The bark anatomy of celastraceous species 11m thiek, from unembedded bark sampIes on hitherto investigated by us, suggests that bark a Reichert sliding rnicrotome. These were sup­ structure could contribute considerably to­ plemented by hand-cut sections. wards the delimitation of taxa, particularly at Sections were double stained in safranin 0 the generic level. In southern African Celastra­ and fast-green FCF (Johansen 1940), and ceae, bark structure is often more useful for mounted permanently in enteIlan (Art. 7961, diagnostic purposes than is wood structure E. Merck, Darmstadt). The phloroglucinol/ (Archer 1990 & unpublished results). hydrochloric acid test and Sudan black B Because of their ethnobotanical impor­ were used for detecting lignin and suberin re­ tance, there is a demand for bark anatomical spectively (Jensen 1962). information on African trees. This would fa­ Hand-cut seetions of the non-collapsed cilitate the identification of bark fragments secondary phloem, mounted in resorcin blue which are extensively traded by tradition al (O'Brien & McCully 1981) or aniline blue healers. Many trees with sought-after barks (Eschrich & Currier 1964), were viewed using are vulnerable to overexploitation. A database bright-field and fluorescence optics to exam­ for the identification of barks is needed, not ine sieve plates. Macerates of a few sampies only for the proteetion of tree species, but were prepared with Schulze's solution (Mc­ also to exert control over the identity of barks Lean & Ivimey-Cook 1941). sold or administered to the public. Barks of Drawings were made using a camera luci­ several species of Cassine s.l. are extensively da. Observations of the rhytidome pose a used in traditional medicine, especiaIly those problem because its brittleness causes it to be containing conspieuous orange pigments. easily tom apart and lost during sectioning. In this paper the comparative bark anat­ To study this tissue, clean-cut pieces of bark omy of the southern Afriean members of the were exarnined under a dissecting rnicroscope Cassinoideae, comprising Cassine s.l. and (Van Wyk 1985). Histochemical testing for the three monotypie genera, Allocassine, Har­ lignin was carried out by applying phloroglu­ togiella and Maurocenia is described and its cinol/hydrochloric acid directly to the bark taxonomie significance assessed. Although surface. placed under the Cassinoideae by Loesener Descriptive terms follow Whitmore (1962a, (1892, 1942), Pleurostylia, with its dry b), Roth (1981) and Van Wyk (1985). The fruits, is not considered to be closely related definition of inner bark foIlows Whitmore to the other southern African Cassinoideae. It (1962a). Terminology of the various scleren­ has therefore been excluded from this study. chymatous elements foIlows the proposals of Special emphasis is placed on bark features Parameswaran (1980). The terms conducting for evaluating previous proposals to subdivide and non-conducting phloem were replaced by Cassine s.l. into several, aIlegedly more na­ non-collapsed and collapsed phloem respec­ tural, genera (e.g. Loesener 1894, 1942; Rob­ tively as suggested by Trockenbrodt (1990). son 1965, 1966). Results Materials and Methods Barks of the investigated species can be Fifty-three sampIes of mature bark, from grouped into six distinct anatomical types. 16 species of Cassine s.l., Allocassine, Har­ The principal structural differences between togiella and Maurocenia were studied using the types are summarised in Table 1. Descrip­ mainly light microscopy. Voucher specimens tions of the various bark types, as weIl as the are housed in the H. G. W. J. Schweickerdt details of the taxa in which they are found, Herbarium (PRU) and listed under Results. are given below. Downloaded from Brill.com10/04/2021 12:50:06AM via free access Archer & Van Wyk - Bark of Cassinoideae 37 Table 1. Summary of the diagnostic bark anatomical features in the six bark types recognised (+ = present; - = absent). o(l o(l s:! s:! l:i i2 s:! ": <::l ~ -2 s:! erd ~-2 .~ Genera proposed by -c:t: :] .~ <::l ~ .5 ~ . ... Archer (1990): ~ ~ f t:i t: ] i: :2l ] ~\..) ~ J 8:il ~ ~ Bark type: Al A2 B C D E Sclerenchyma Large primary sclereids + + Fibres + + Sclerenchyma absent in both non--collapsed and collapsed phloem + Chambered crystalliferous sclereids + + Crystals and other features Styloids + Septate crystal strands + Elastic threads + Cell width of widest phloem rays (near cambium) 6 6 4 3 1 (2) 1 (2) Periderm Sclerenchymatous phelloderm + Number of periderm layers 2-5 2-3 2-3 2-3 3-8 Persistent periderm + Bark type Al (Figs. 1,7,9, 10, 15, 16,23, Ion Eck!. & Zeyh.): Archer 303,312,366.­ 24) C. matabelicum (Loes.) Steedman (alterna­ Diagnostic characters: Large fibre-sclereids tively placed under Elaeodendron Jacq. f.): and/or sclereids abundant throughout the sec­ Venter & Archer 184, 185. - C. papillosa ondary phloem; sclereids in the non-dilating (Hochst.) Kuntze (alternatively placed under phloem often clustered in fusiform groups Elaeodendron): Archer 217,224,344. - C. (radial longitudinal section), also in the dila­ transvaalensis (Burtt Davy) Codd (alterna­ tation zone (transverse section); phelloderm tively placed under Crocoxylon): Archer 215, crystalliferous and weakly sclerified, the 222, 245, 246, 325, Pienaar & Archer 1338. latter elements irregularly arranged, or in a General: Bark surface longitudinally fis­ more or less continuous ring; rhytidome weIl sured; fissures narrow to broad, often cutting developed. deep into the rhytidome
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