IAWA Bulletin n.s., Vol. 11 (2), 1990: 173-182

WOOD AND BARK ANATOMY OF LAURACEAE IV. DAHLGRENODENDRON J.J.M. VAN DER MERWE & VAN WYK

by

H. G. Richter* and A. E. Van Wyk**

Summary The southem African tree, Dahlgrenoden­ schmiedia (Van der Merwe et af. 1988). Al­ dron natalense, previously ascribed to the though the ovary of Dahfgrenodendron is genus Beilschmiedia was recently segregated superior, the mature fruit is completely in­ as a monotypic genus, Dahlgrenodendron, cIuded in the accrescent hypanthium (recep­ on account of its distinctive exomorphology, tacular tube), thereby resembling genera such palynology and fruit structure. Dahlgreno­ as Aspidostemon, Cryptocarya, Eusideroxy­ dendron differs from Beilschmiedia in nearly fon, Potoxyfon and Ravensara. Mainly on the all quantitative and qualitative features of basis of this character, and also its 2-celled wood and bark anatomy. The overall struc­ anthers on the stamens, Van der Merwe et af. tural pattern of these tissues does not support (1988) provisionally associated Dahfgreno­ the initial assignment of Dahlgrenodendron to dendron with tribe Cryptocaryeae, subtribe the tribe Cryptocaryeae, nor does it fit any Cryptocaryineae. However, the marked incon- other lauraceous taxon. Within Lauraceae, an isolated non-aligned status is provisionally proposed for the genus. Key words: Wood anatomy, bark anatomy, -, l l~ - , .~.~. taxonomy, Aspidostemon, Beilschmiedia, './ ~. Cryptocarya, Dahlgrenodendron, Laura­ f ceae. / " NATA L q;' ,r'\ . V" Introduction f ~ The monotypic genus Dahlgrenodendron , Ir .v was established by Van der Merwe et al. , Ja -- , -~ (1988), and is based on D. natalense, a spe­ ~ ~ ~ cies first described as Beilschmiedia natalen­ ~fv'<- W' I-Pondoland sis J.H. Ross (1973). ? I I Dahlgrenodendron natafense is an ever­ ~ ,.. .1.. I, .. " green forest tree up to about 20 m high, and '/ l"llllll I is one of several plant species endemie to a ,.. /'" ,e- I ". few restricted areas of Natal Group sand­ stone in Natal and Pondoland in southem Fig. 1. Map showing the geographieal dis­ Africa (Fig. 1). With the total number of in­ tribution of Dahfgrenodendron natalense in dividuals probably not exceeding 200, it is southem Africa. The three small dots signify considered a palaeoendemic on the brink of relictual occurrences of one or two plants natural extinction. Evidence from macromor­ only. Most individuals are confined to patches phology, palynology and fruit structure indi­ of the relict forest in the southem Natall cated that D. natalense was misplaced in Beil- Pondoland sandstone region (large dots).

* Institut für Holzbiologie und Holzschutz, Bundesforschungsanstalt für Forst- und Holz­ wirtschaft, Leuschnerstrasse 91,2050 Hamburg 80, F.R.G. ** H.G.W.J. Schweickerdt Herbarium, Department of Botany, University of Pretoria, Pre­ toria, 0002 South Africa.

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Figs. 2-5. Secondary xylem of Dah/grenodendron nara/ense. - 2-4: Transverse, tangential and radial sections. - 5: Ray heterogeneity, septate fibres and fibre lumina occIuded by organic compounds. - Scale bar = 200 ~m.

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Table 1. Wood and bark specimens exarnined of Dahlgrenodendron natalense (lH. Ross) J.J.M. Van der Merwe & A.E. Van Wyk.

RBHw Coll./No. Origin of specimen Quarter degree No. (Southern Africa) grid reference Wood 18936 Van Wyk 8123 Natal, Ngutu Falls 2930#DB s. n. Van Wyk8256 Natal, Umdoni Park 3030# BC s. n. Van Wyk8258 Natal, Urntamvuna Nat. Res. 3130#AA s. n. Van Wyk8259 idem idem s. n. Van Wyk8260 idem idem Bark s. n. Van Wyk A-542 Natal, Umtamvuna Nat. Res. 3030#CC s. n. Van Wyk A-546 Transkei, Sikuba River Gorge 3130#AA s. n. Van Wyk A-553 Natal, Umtamvuna Nat. Res. idem gruence between different characters of some Schweickerdt Herbarium (PRU). Localities of these genera has led to the suggestion that are given in Table 1, also as quaner-degree the mutual presence of an accrescent hypan­ grid references (Edwards & Leistner 1971). thium may, in at least some taxa, reflect To minimise damage to these rare trees parallel evolution (Rohwer & Richter 1987). wood sampies (mainly sapwood) were ex­ Dahlgrenodendron is unusual in the Lau­ traeted with an increment borer, at 1.5 m raceae in having eonspicuously striate pollen height, from vertieal sterns with diameters grains (Van der Merwe et al. 1988). Striate from 23 to 60 cm (wood sampies) and 14 to grains have not yet been recorded elsewhere 45 em (bark sampIes). In one instance, eross­ in the family or in the Laurales. Henee, Van seetional sampies were also taken from a der Merwe et al. (1988) also suggested that dead 20 cm diameter stern (Van Wyk 8256). Dahlgrenodendron might eonstitute an iso­ Mieroscopie slides of wood sampIes were lated genus in Lauraeeae, perhaps best plaeed prepared following comrnon laboratory pro­ in a new tribe or subtribe of its own. The cedures. Bark material was sectioned after in­ present paper repons on the wood and bark filtration with polyethelene glyeol (PEG, DP structure of Dahlgrenodendron natalense, and 1500) using the method developed by Rupp its possible taxonomie signifieanee. Because (1964) and modified by Richter (1981a). the affinities of Dahlgrenodendron are ob­ Wood and bark sections were double stained seure, this study was undenaken in the hope with aqueous or alcoholie solutions of 1% of establishing more satisfaetorily the rela­ chrysoidin/acridin red and astra blue. tionships of the genus. Tests for fluoreseence, natural saponins This paper is the founh of aseries on the and aluminium follow the methods prescribed systemarie wood and bark anatomy of genera by Miller (1981), with minor modifieations belonging to the Lauraceae (Richter 1981b, (Dyer 1988), and were based on one wood 1985, 1990) and eomplements an earlier sur­ sampie (Van Wyk 8256). vey ofthe family (Richter 1981a). Specifie gravity based on oven-dry weight and volume (for Van Wyk 8256 only) was Material and Methods adjusted to specific gravity at 12% me via The wood and bark descriptions of Dahl­ conversion table (Miller & Baas 1981). grenodendron natalense are based on five Rhytidome structure was studied on trans­ wood and three bark sampies (Table 1), col­ verse surfaees of dry bark sampies, follow­ leeted by A.E. Van Wyk and backed by her­ ing the method outlined previously (Van barium vouchers deposited at the H.G.W.l Wyk 1985).

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Figs. 6-8. Secondary phloem of Dahlgrenodendron natalense. - 6: Bark surface pattern. - 7 & 8: Low magnification overview of younger (Van Wyk A-546) and older (Van Wyk 542) bark with distribution pattern of scIereid bundles. - Scale bar of 6 = 50 mrn; of 7 & 8 = 200 ~m.

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The terminology used in the wood descrip­ thick-walled (lumen ~ double wall thick­ tion adheres to the standards proposed by the ness); mean fibre length around 1100 Ilm, International Association of Wood Anato­ overall range from 700-1400 Ilm; with often mists in its most recent form (IAWA 1989); distinctly bordered pits (3-5 Ilm) commonly bark terminology follows Trockenbrodt confined to radial walls; regularly septate, (1990). Terms used in wood and bark de­ septa few (usuall y 1, rarel y up to 3 per fibre); scriptions but not listed in the terminologies frequently with reddish brown gum deposits. mentioned above (e.g. 'stratified fibres, col­ Growth increment boundaries in some speci­ umnar sclereids') are explained and/or illus­ mens marked by 1-2 rows of radially flat­ trated in the general treatise on Lauraceae tened fibres. (Richter 1981a). Parenchyma inconspicuous, scanty para­ tracheal to weakly vasicentric, definitely not Wood description (Figs.2-5) in concentric bands as reported by Swart & General: No visual distinction between Van der Walt (1986); 4-8 cells per paren­ heartwood and sapwood (largest cross-sec­ chyma strand, non-storeyed. tional sampie 200 mm in diameter); wood Rays non-storeyed; uniseriate, 1-6 cells pale reddish brown and without distinctive high, composed of upright, rarely square odour or taste; specific gravity 878 kg/m3 at cells, and multiseriate (2-)3-5(-7) cells 12 percent moisture content (784 kg/m3 for wide; body cells procumbent with 1-2, rarely oven-dried wood); splinter bums to a grey or up to 4 marginal rows of predominantly up­ black charcoal; fluorescence of heartwood as right cells; mean height of the 25 largest rays weIl as water and ethanol extracts negative; averaging 0.50-0.80 mm between speci­ froth test for natural saponins negative; col­ mens, frequency 5-8/mm. Vessel-ray pits our of water and ethanol extracts reddish commonly large and with much reduced bor­ brown; chrome azurol-S test for aluminium ders, pit outline rounded, oval to irregular in negative. shape to horizontally (gash-like) or diagon­ Anatomy: Wood diffuse-porous, vessels ally extended; occasionally similar to inter­ evenly distributed with occasional tendency vascular pits in procumbent cells. Rays usu­ to larger numbers of smaller vessels along ally replete with reddish brown organic com­ growth ring boundaries (latewood); solitary pounds. and in radial multiples of 2-4, the latter pre­ Secretory (oil or mucilage) cells, crystal­ dominant in most specimens examined; few line deposits and silica not observed in the clusters present; round to oval in outline with secondary xylem of the specimens examined. tendency to angularity, thin-walled; rather frequent, specimen means ranging from 40- 55/mrn2; tangential diameter varying from a Bark description (Figs.6-12) minimum of 30 Ilm (latewood) to a maximum General: Periderms usually 3-6, not un­ of 110 Ilm (earlywood), the specimen means dulating, completely separating living tissues (25 largest vessels) from 70-90 Ilm. Perfo­ from dead; sequential periderms formed as ration plates simple and mostly as wide as discontinuous layers connected to the older vessel elements. Intervascular pitting alter­ layers and usually overlapping one another, nate, pits (3-)4-5(-6) Ilm in diameter (ver­ slightly curved or usually more or less paral­ tical dimension), circular to somewhat poly­ lel to the cambium, radially up to about 4 mrn gonal in outline according to varying degrees apart; occasionally penetrated by the sclereid of crowding; non-vestured, apertures includ­ bundles of the secondary phloem. ed; thin-walled tyloses present but rare (most Rhytidome present and covering the whole material is sapwood); vessellumina frequent­ bole surface, usually 10-20 mm thick, con­ ly occluded by reddish brown organic com­ sisting of 2-6 superimposed rhytidome pounds. layers, each formed by the cutting off of the Fibres non-stratified, polygonal in cross collapsed phloem by weakly scallop-shaped section; rather wide, the largest measuring periderm connected to the preceding periderm from 30 to 50 Ilm in diameter; medium to layer.

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Figs. 9-12: Secondary phloem of Dahlgrenodendron natalense. - 9: Transverse, transition from xylem to phloem in young bark (Van Wyk 546) with sclereid bundle. - 10: Transverse, close-up of sclereid bundles. - 11: Longitudinal (radial) view of sclereid bundle. - 12: Close-up showing sclereid bundle composed of elongated ('columnar') sclereids and sclerification of traversing ray tissue. - Scale bar = 200 J.lm.

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Cut-off phloem tissue becoming dark composed of regular sclereids, particularly in brown; fairly resistent to decay, except the areas between wedges of dilatation tissue. sclereid bundles which soon rot away; secre­ Phloem rays resemb1e wood rays in size tory (oil) cells usually still visible under mag­ and composition and do not change signifi­ nification as minute yellow granules in de­ cantiy in dimensions and direction radially. cayed outer layers of cut-off phloem. When traversing sclereid islands, ray cells Surface fairly slowly scaling or breaking become sclerified. Occasionally wedge-like off in irregularly outlined pieces of rhytidome dilatation zones are formed by tangential ex­ (Fig. 6), each piece being the remnant of one pansion and subdivision ofaxial phloem or more rhytidome layers. Fissures usually parenchyma. narrow, longitudinal and transverse, often Oil cells numerous, scattered throughout cutting deep into the rhytidome layers; slough­ the secondary phloem and the cortex. Small ing taking place continually over the whole cubic, navicular and acicular crystals abun­ surface of a bole; scales apparently maintain­ dant in ray, rare in axial parenchyma. ed for a long time because of the continuous periderm system and the lack of adefinite Discussion sloughing mechanism; the pieces of rhyti­ Based on wood anatomical evidence dome breaking away mainly owing to weath­ Swart & Van der Walt (1986) questioned ering processes and fissuring. Slash dark Ross' decision to assign Dahlgrenodendron pinkish red. natalense to Beilschmiedia. Initially, geo­ Anatomy: Secondary phloem composed graphic distribution and similar vessel mor­ of sieve tubes + companion cells and phloem phology suggested a possible relationship parenchyma in, respectively, decreasing and with native (southern African) Cryptocarya. increasing proportions with radial distance However, a comparative statistical analysis of from the cambium; interspersed with numer­ a wide array of quantitative parameters made ous oval to diamond-shaped sclereid bundles them reject the previous assumption. (Figs. 7, 8) consisting ofaxially elongated The present, more complete assessment of 'columnar' sclereids (Figs. 9-12); few cir­ the secondary xylem complemented by the cular to tangentially extended (cross section) new evidence on phloem structure fully cor­ normal sclereids present either isolated or roborates Swart & Van der Walt's earlier marginally associated with sclereid bundles; doubts and, moreover, Van der Merwe et phloem fibres lacking. In younger bark (Van al.'s decision to assign Beilschmiedia nata­ Wyk A-553) the secondary phloem is delirn­ lensis the status of a new monotypic genus - ited towards the periphery by a continuous Dahlgrenodendron. The overall structural ring of smaIl, irregularly shaped, ± isodia­ pattern of Dahlgrenodendron does not corre­ metric sclereids enclosing isolated groups of spond weH with any of the existing laurace­ primary phloem fibres. In older bark (Van ous taxa. When comparing its xylem struc­ Wyk A-542, A-546) the sclereid ring is still ture with that of the known genera, one finds including traces of primary phloem fibres, for every shared feature one or more that are and all external parts are cut off by sequential not in agreement. A comparison with Beil­ periderms and eventually lost. schmiedia (Table 2) and Cryptocarya, the The sclereid ring is followed by the cortex two genera most often considered possib1e (parenchyma cells with some isolated scle­ relatives, particularly illustrates these differ­ reids) gradually phasing into the last-formed ences: Dahlgrenodendron natalense features periderm. Phellern about 1 mm thick, usually paratracheal parerichyma, albeit weakly de­ pale brown on cut transverse surfaces of veloped, but lacks the (in Lauraceae) highly dried bark sampies. Phelloderm well defined, diagnostic marginal bands that characterise about 0.25 mm thick, initially parenchymatic both Beilschmiedia and Cryptocarya. Fibre with darkly stained contents, but soon with morphology (dimensions, wall thickness; most of the radial and inner tangential cell pits) agrees very weH with Beilschmiedia walls lignified and slightly thickened; oldest except for the septate condition (fibres are layers occasionally strongly sclerified and exclusively non-septate in Beilschmiedia and

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Table 2. Differences in secondary xylem and phloem features between Dahlgrenodendron and Beilschmiedia.

Feature Dahlgrenodendron Beilschmiedia Wood Vessels I pores - tang. diam. range 30-110 Jlm 60-300 Jlm - specimen means 70-90 Jlm 90-220 Jlm - average N/mm2 40-46-55 3-6-12* - diam. interv. pits (3-)4-5(-6) Jlm 8-11 Jlm (vertical) Fibres - arrangement non-stratified horiz. stratified - morphology septate non-septate - contents - gum deposits present absent - tyloses not observed present (most species) Axial parenchyma scanty paratracheal a) paratracheal vasicentric to only aliform b) zonate marginal bands Wood rays - inorganic contents not observed a) small prismatic crystals b) silica (small particles or large aggregates) Sec re tory cells not observed commonly present

Bark Mechanical tissue - phloem fibres absent absent - sclereids axially elongated ± isodiametric - contents absent a) large prismatic crystals, and b) vitrious silica (few species) Parenchyma - lignified parenchyma absent present in most specimens Phloem rays - sclereid formation absent often present in outer phloem

* Beilschmiedia berteroana and B. miersii from temperate South America (Chile) possess smaller (70-90 Jlm) and more frequent (17-25/mm2) vessels and are not included here.

Cryptocarya). Vessel characters such as out­ The secondary phloem of Dahlgrenoden­ line, arrangement, grouping, and perfora­ dron natalense lacks phloem fibres. It shares tion plates nearly match those of Beilschmie­ this (in Lauraceae) highly diagnostic feature dia. Yet size, frequency, and the (in Laura­ with a small number of taxa including Beil­ ceae) highly diagnostic intervessel pit dia­ schmiedia (+Potameia, Triadodaphne), Cryp­ meter (vertical) emerge as distinctly different tocarya (+ Ravensara), Aspidostemon, and a (Table 2). few genera assigned to the more distant tribe

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Cinnamomeae, subtribes Cinnamomineae cate common ancestry or elose relationship. (i.e., Neocinnamomum, part of neotropical Also, the number of anther cells on the sta­ Ocotea) and Anibineae (i.e., Aniba, Licaria) mens must be seriously questioned as a gen­ sensu Kostermans (1957a). However, in erally valid taxonomic character in Lauraceae. combination with the presence ofaxially It has been used consistently in systematic elongated ('columnar' fide Chattaway 1959) treatments of the family (e.g., Pax 1891, selereids the scope of structural agreement is Kostermans 1957a, Hutchinson 1964) in narrowed down to the latter five genera and a delimiting taxa at the subfamilial, tribai and small segment of neotropical Cryptocarya (C. generic level. Bamber & Surnmerville (1979) aschersoniana, C. moschata) from southern report a strong agreement between both bark Brazil (Richter 1981a). In thecontextofsim­ and wood characters of selected genera and ilar fioristic traits of southern temperate Bra­ Pax' elassification in the subfamilies Persoi­ zil and southern Africa it would be valuable deae (4-celled anthers) and Lauroideae (2- to compare the bark structure of Cryptocarya celled anthers). Yet, there are many cases species from both areas to gain a better un­ cited in older (e.g. Mez 1889) and more re­ derstanding of their relationships. cent (Kopp 1966; Van der Werff & Richter Following Kostermans' (1957a) system­ 1985; Van der Werff 1986; Rohwer 1985, atic treatment of Lauraceae Van der Merwe et 1988) studies that do not support the idea that al. (1988) provisionally placed Dahlgreno­ taxa with 2- and 4-celled anthers form distinct dendron in the tribe Cryptocaryeae, subtribe groups. Raij & Van der Werff (1988) discuss Cryptocaryineae. This ineludes all those these cases in detail. They come to the woody members of Lauraceae (excl. Cassy­ conclusion that the use of this readily thoideae) with a superior ovary, but with the observable but artificial character obscures fruit completely ineluded in the accrescent rather than enhances laural taxonomy and hypanthium, and with 2-celled anthers. An should be scaled down in favour of more assignment to Cryptocarya or Ravensara, the convincing arguments, ineluding those de­ two genera comprising Cryptocaryineae, is rived from other than macromorphological ruled out because of distinct differences in evidence. pollen morphology and internal fruit struc­ Hence, although Dahlgrenodendron is char­ ture. However, this provisional placement of acterised by 2-celled anthers and external Dahlgrenodendron was made subject to revi­ fruit characters similar to those of Crypto­ sion pending further studies on the opposite­ carya, its wood and bark anatomy weighs in leaved Madagascan species of Cryptocarya heavily against relegating Dahlgrenodendron, which, in the meantime, were elevated to however provisionally, to the subtribe Cryp­ generic rank as Aspido~temon (Rohwer & tocaryineae. An isolated non-aligned status is Richter 1987). preferred until more evidence from compara­ The segregation of Aspidostemon from tive studies on these and other lauraceous Cryptocarya is based on profound differ­ taxa becomes available. ences in fiower and secondary xylem/phloem structure notwithstanding the external fruit Acknowledgements characters which it shares with Cryptocarya, We are indebted to Stephanie T. Dyer for and also Dahlgrenodendron. It was proba­ conducting some of the wood tests, to Clau­ bly this common trait that led Kostermans dia Muche for preparing the wood and bark (1957b) to place these species in Cryptocarya slides, and also to Tony Abbott for assistance with subgeneric rank. during field work. It is hypothesised that this particular fruit structure common to the taxa assembled in Cryptocaryeae sensu Kostermans (1957a) References represents a derived character state within Bamber, R. K. & R. Summerville. 1979. Lauraceae that has developed parallel in in­ Taxonomic significance of sclerified tissue dependent lines (Rohwer & Richter 1987; in the barks of Lauraceae. IAWA BuH. Richter 1990). Therefore, it might not indi- 1979/4: 69-74.

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Chattaway, M.M. 1959. The anatomy of Richter, H. G. 1985. Wood and bark anatomy bark. VII. Species of Eugenia (sens. lat.). of Lauraceae II. Licaria Aubiet. IAWA Trop. Woods 111: 1-14. Bull. n.s. 6: 187-199. Dyer, S. T. 1988. Wood fiuorescence of Richter, H.G. 1990. Wood and bark anat­ indigenous South African trees. IAWA omy of Lauraceae III. Aspidostemon Bull. n.s. 9: 75-87. Rohwer & Richter. IA W A BuH. n. s. 11: Edwards, D. & A.O. Leistner. 1971. A 47-56. degree reference system for citing bio­ Rohwer, J.G. 1985. Der Ocotea-KompIex, logical records in southem Africa. Mitt. eine Synopsis. Dissertation, Fachbereich Bot. Stsarnml., München 10: 501-509. Biologie, Universität Hamburg. Hutehinson, J. 1964. The genera of fiower­ Rohwer, J.G. 1988. The genera Dicypel­ ing plants, 1 (24). Clarendon Press, Ox­ lium, Phyllostemonodaphne, Systemono­ ford. daphne and Urbanodendron (Lauraceae). IAWA Committee. 1989. IAWA List of Bot. Jahrb. Syst. 110: 157-171. microseopie features for hardwood iden­ Rohwer, J.G. & H.G. Richter. 1987. Aspi­ tification. IAWA Bull. n.s. 10: 219-332. dostemon, a new lauraceous genus from Kopp, L. 1966. Monograph ofthe American Madagascar. Bot. Jahrb. Syst. 109: 71- Persea. Mem. New York Bot. Gard. 14: 79. 31-121. Ross, J.H. 1973. Lauraceae: A new generic Kostermans, A.J.G.H. 1957a. Lauraceae. record for South Africa and a new spe­ Reinwardtia 4: 193-256. eies. Bothalia 11: 118. Kostermans, A.J.G.H. 1957b. Le genre Rupp, P. 1964. Polyglycol als Einbettungs­ Cryptocarya R. Br. (Lauracees) a Mada­ medium zum Schneiden botanischer Prä• gascar. BuH. Jard. Bot. Brux. 27: 173- parate. Microcosmos 53: 123-128. 188. Swart,J.P.J.&J.J.A. VanderWalt.1986. Mez, C. 1889. Lauraceae americanae, Mo­ Systematic wood anatomy of the southern nographie descripsit. J. König!. Bot. African Lauraceae. Proc. For. Prod. Res. Gart., Bot. Mus. Berlin; 5th reprint 1963. International- Achievements and the Fu­ Miller, R.B. 1981. Explanation of coding ture, Pretoria (1985), 16/9: 10 pp. procedure. IAWA Bull. n.s. 2: 111-145. Trockenbrodt, M. 1990. Survey and discus­ Miller, R.B. & P. Baas (coord.). 1981. Stan­ sion of the terminology used in bark an at­ dard list of characters suitable for com­ omy. IAWA Bull. n.s. 11: 141-166, this puterized hardwood identification. IAWA issue. BuH. n. s. 2: 99-45. Van der Merwe, J.J.M., A.E. Van Wyk & Pax, F. 1891. Lauraceae. In Engler & Prant!, P.D.F. Kok. 1988. Dahlgrenodendron, Die natürlichen Pfianzenfamilien 3 (2). W. a remarkable new genus from Natal and Engelmann, Leipzig. Pondoland. S. Afr. J. Bot. 54: 80-88. Raj, B. & H. Van der Werff. 1988. A con­ Van der Werff, H. 1986. A new species of tribution to the pollen morphology of neo­ Caryodaphnopsis (Lauraceae) from Peru. tropical Lauraceae. Ann. Missouri Bot. Syst. Botany 11: 415-418. Gard. 75: 130-167. Van der Werff, H. & H.G. Richter. 1985. Richter, H.G. 1981a. Anatomie des sekun­ Caryodaphnopsis Airy-Shaw (Lauraceae), dären Xylems und der Rinde der Laura­ a genus new to the Neotropics. Syst. Bot. ceae. Sonderbände des Naturw. Vereins 10: 166-173. Hamburg 5. Paul Parey, Hamburg. Van Wyk, A.E. 1985. The genus Eugenia Richter, H.G. 1981b. Wood and bark anat­ (Myrtaceae) in southern Africa: Structure omy of Lauraceae 1. Aniba Aubiet. IAWA and taxonomie value of bark. S. Afr. J. BuH. n.s. 2: 79-87. Bot. 51: 157-180.

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