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IAWA Journal, Vol. 15 (1), 1994: 3-45

SURVEY OF ENGLISH MACROSCOPIC TERMINOLOGY

by

Leo Junikka 1 Department of Botany, P.O. Box 7 (Unioninkatu 44), 00014 University of Helsinki, Finland

Summary Tenns of outer and inner bark characteris­ field characters of valuable timber with tics are critically surveyed. Different macro­ notes on barks. They were followed by Beard scopical tenns with their synonyms are listed (1944; Tobago), De Rosayro (1960; Ceylon), for a comparison of bark features. Sugges­ and Den Outer (1972; Ivory Coast). tions are given for a standardised usage of Whitmore (1962a, b, c), in his notable stud­ the tenns to stimulate a practice of pertinent ies on , described seven dis­ field notes and facilitate understanding of de­ tinct bark types and demonstrated that the scriptions. Preferred tenns are printed in bold bark may provide valuable taxonomic infor­ face and preferred definitions in italics. mation, and may shed light on a number of Key words: Bark morphology, tenninology. important taxonomic problems. Voorhoeve (1965) produced a comprehensive study of Introduction some Liberian high forest trees with a great Notes on the appearance of the bark or the deal of infonnation on bark characters. Roth fonn of a are often wanting or inade­ (1981) summarised studies on tropical barks quate in most tropical floras. Only a minority in Venezuelan Guyana with some characters of collectors include various field char­ and definitions worthwhile to comment. In acters on the labels of herbarium sheets. How­ recent years good examples of floras ever, careful descriptions of mature barks using different bark features are Tailfer's and other vegetative parts, especially in rain (1989) work on tropical and Polak's forest trees, may provide a valuable tool for (1992) work on Guyana. Thus good data are reliable and quick plant identification. Plant available, but only from a few areas and, vir­ collectors often cannot reach the crown can­ tually, only from a few plant groups such as opy of the rain forest emergents, or if they the (Francis 1951) or the obtain a branch sample, there may be no family Dipterocarpaceae (Whitmore 1962c). or fruits. Furthermore, the tedious In contrast to the strong standardisation in process of naming belonging to certain anatomical terms by the lAWA (1957, families or other taxonomic units which are 1964, 1989), bark anatomical terms were re­ notoriously difficult to identify in the herbar­ viewed to a certain extent by few authors ium might be greatly speeded up if good bark only, e.g. Esau (1969) and Martin & Crist characters were available. This, in turn, could (1970). Later a thorough discussion of vari­ prevent undue delay in publishing, for in­ ous tenns with references to relevant litera­ stance, the results of inventories of tropical ture and proposal of a list of bark anatomical forests. terms to be used was published by Trocken­ There have been several attempts to con­ brodt (1990). struct keys based on bark characters. Presum­ The present paper concentrates on bark ably, Lely (1925) from Northern Nigeria and morphology. As the anatomical composition Foxworthy (1927) from the Malay Peninsula of the bark highly contributes to the outer as­ were the first who carefully listed different pect and other characters of the bark, terms

1) Temporary: Herbarium, Department of Plant Ecology and Evolutionary Biology, University of Utrecht, The Netherlands.

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used in bark anatomy and those used in bark This paper is intended as a proposal to­ morphology overlap to a certain degree, par­ ward a consistent bark morphological termi­ ticularly those concerning bark tissues. nology. I have quoted 'representative' defi­ The first efforts to compile a descriptive nitions from the anatomical literature, from morphological terminology of barks were some field floras of different geographical made by Wood (1952), De Rosayro (1953), areas, and from common botanical textbooks and Wyatt-Smith (1954). These papers were written for tropical areas. Some well-known mainly concerned with bark features based terms have been omitted, although some syn­ on the authors' own experience from South­ onyms are accepted. I have restricted the east Asia. Also some terminological defini­ scope of this study to the English literature, tions of bark features are included in the but some common terms (e. g. rhytidome, glossaries of tree floras, such as Corner's phloeme) from French publications are in­ book 'Wayside trees of Malaya' (1940; 3rd cluded. ed. 1988). Various terminological notes can This tentative list of terms certainly does be found in textbooks of tropical botany. A not pretend to cover all the terms used in bot­ good example is Letouzey's (1986) book any and forestry. Hopefully, though, it will 'Manual of forest botany' in two volumes on bring more clarity in a rather confusing bark tropical Africa (translation from the original terminology, and thus contribute to a badly French, 1972). Among plant anatomical text­ needed standardization. books, Eames & MacDaniels (1947), Esau (1960, 1965, 1969, 1979), and Fahn (1990) have referred to macroscopical characteristics Structure, symbols, and abbreviations of barks. used in the list Rollet (1980, 1982) published a review about the variability of characters used in ma­ Structure croscopic bark analysis. Based on his survey This paper is divided into three main chap­ of field floras that contain notes on bark fea­ ters: 1) a list of accepted terms, 2) a survey of tures, and his own experience from the trop­ the terminology with the accepted definitions, ics, he described the potentiality of using synonyms and comments on relevant bark features, giving numerous examples. literature, and 3) a list of rejected terms with As Trockenbrodt (1990) has pointed out, many cross references. The accepted terms the large number of terms with partially or are printed in bold face, and the accepted def­ wholly overlapping meaning "reveals the lack initions in italics. The survey of the terminol­ of a concise and widely applicable terminol­ ogy (and the preceding list of accepted terms) ogy of bark structure." Not only are different is subdivided into five parts, dealing with: morphological terms applied to the same struc­ I bark and its component tissues ture but also, in some cases, a term is obscure­ II bark texture ly defined, thus allowing different meanings. III bark patterns The main difficulty with regard to bark mor­ a. bark patterns in tangential and cross phological descriptions is that features which sections are pure responses to the environment have b. fissuring to be distinguished from those which are in­ c. exfoliation herent to a particular species. One has to d. external markings know the variability of barks: between indi­ N exudation viduals of the same species variation is often V bark cutting considerable, and even in different parts of the same individual. Only efficient collecting Entries are printed in alphabetical order and of data from a large number of mature trees the attributes within an entry are arranged using accepted and standardised descriptive according to its meaning. Numbers between terms will enlarge the knowledge of bark parentheses, (1), (2), etc., indicate two or variation. A simple and well-defined termi­ more meanings of a particular term. The defi­ nology is, therefore, a prerequisite. nition of a term by its original author has been

Downloaded from Brill.com09/24/2021 07:42:43AM via free access Junikka - Macroscopic bark terminology 5 maintained, as far as possible. Sometimes an Ed Edlin (1976) author did not define a term explicitly, so that EM Eames & MacDaniels (1947) its meaning had to be deduced from the origi- Esl Esau (1960) nal context. Often the original definitions had Es2 Esau (1965) to be abridged to save space. For practical Es3 Esau (1969) reasons some definitions are placed under en- Es4 Esau (1979) tries in the chapter on rejected terms, but they Fh Fahn (1990) are commented upon in the list of accepted Fo Ford-Robertson (1971) terms. Fs Francis (1951) GE G6mez-Vazquez & Engleman (1984) Symbols GJ Ghouse & Jamal (1978) Hi Hightshoe (1989) is placed after a reference showing a * HJC Hall et al. (1970) direct quotation from the source. Ho Howard (1971) ~ refers to the term under which an ex- IAWA lAWA, Committee on Nomenclature planation is given, and which is usually (1964) preferred. Ir Irvine (1952) an alternative term which is also accept- Ja Jackson (1928) Ji ed. Jimenez-Saa (1973) Kr KrUssmann (1984) the cited author has explicitly used the Ku Kunkel (1965) synonym in the sense given here. Le Letouzey (1986) Cf. confer = compare with. MC Martin & Crist (1970) Me Metcalfe (1979) Inc!. the term is included in the definition MET Macdonald Encyclopedia of Trees proposed for this entry. (1982) See an explanation is given under the indi- MW Meijer & Wood (1964) cated entry, which is not synonymous. Nu Nultsch (1971) Oul Outer, den (1967) Syn. rejected synonym(s). 0u2 Outer, den (1972) Po Polak (1992) PW Penford & Willis (1961) Abbreviations PZ Panshin & De Zeeuw (1980) AHDE American Heritage Dictionary of RDMB Radford et al. (1974) English language (1980) REC Raven et al. (1976) Ba Bena (1960) Ro Rollet (1980, 1982) BCFTl British Commonwealth forest termi- Rt Roth (1981) nology 1 (1953) Ry Rosayro, de (1953) BCFT2 British Commonwealth forest termi- Sa Srivastava (1964) nology 2 (1957) SAF Society of American Foresters, Be Beard (1944) Committee on Forestry Terminology BKI Brooker & Kleinig (1990a) (1950) BK2 Brooker & Kleinig (1990b) Ss Storrs (1979) BM Bi.isgen & Mi.inch (1929) Th Thrower (1988) Bo Bor (1953) Ti Timberlake (1980) Br Brown (1971) Vo Voorhoeve (1965) Cg Chang (1954) Whl Whitmore (1962a) Cha Chattaway (1953) Wh2 Whitmore (1972) Ck Craddock (1932) Wo Wood (1952) Co Comer (1988) WS Wyatt-Smith (1954) Dr Dirr (1977) Wy Wyk, van (1985) Ed Edlin (1976) YYI Yunus et al. (1990)

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ACCEPTED TERMS: DEFINmON AND COMMENTS

I. Bark and its component tissues (Many terms commented upon here are indicated in the schematical drawing in Fig. 1)

bark bark, inner (1) all outside the vascular (1) = I: . regardless of its composition (Trocken­ (2) -t I: phloem, non-collapsed secondary. brodt 1990*). (3) the living part of the bark inside the inner- Syn.: rind (Ck, BM, Ja). most cambium (Raven et al. 1976*). (2) in dicotyledons the term bark in a popular Syn.: blaze (Be), live bark (Wo, Ba, sense is applied to all secondary tissues Ji), living bark (Po). external to the in stem and of (4) adherent rhytidome under the easily de­ woody plants (Metcalfe 1979*). tachable scales and the tissues below it (3) a dead outer protective tissue of woody (De Saint Aubin 1963). plants, derived from the ; varies (5) -t I: periderm (1). greatly in appearance and texture; often Comments: including all tissue from vascular cam­ Martin & Crist (1970) propose that anato­ bium outward (Dirr 1977*). mists refer to the inner bark as phloem. Trock­ (4) -t I: rhytidome (1), (2). enbrodt (1990), too, recommends a more ac­ Comments: curate expression instead of inner bark. He In plants with bark suggests the expression 'living secondary consists of the secondary phloem, the pri­ phloem' or 'secondary phloem up to the last­ mary tissues still existing outside the sec­ formed periderm'. Although the term phloem ondary phloem, the periderm, and the dead is mostly favoured by anatomists, the term tissue outside the periderm. Metcalfe (1979) inner bark is accepted here as an alternative, omits primary tissues from the definition of because many non-anatomists use it frequent­ bark. Dirr's (1977) definition has to be re­ ly. In this sense, inner bark or phloem is com­ jected, as it is self contradictory. plementary to outer bark or rhytidome. Other Although the etymological of the term meanings of the term should be avoided. bark is the same as the German 'Borke', mean­ ing rhytidome, there is no practical reason to change the well-established definition of the bark, outer bark (1). (1) = I: rhytidome (1). (2) -t I: rhytidome (2). II Ii " I I II I I (3) -t I: rhytidome (3). ~~'~'~~'~I~~~~I/ (4) used by Roth (1981) in the microscopical description of the outer layer of the phloem periderm [ where "the proper arrangement of cells rhytidome and especially the pattern produced by the hard bast, is disturbed by active dilatation growth." phellem Cf: I: phloem, collapsed secondary; bark, middle (1) (rejected term). phloem See comments under I: rhytidome. inner bark

\=~~~~::t----=.ca mbium bark fibre xylem fibre in the bark (Trockenbrodt 1990*). Syn.: (Esl). Fig. 1. Suggested terms for the tissue zones resulting from rhytidome formation. See comments under I: phloem fibre.

Downloaded from Brill.com09/24/2021 07:42:43AM via free access Junikka - Macroscopic bark tenninology 7 bast many species of Eugenia, it darkens purplish (1) any fibres of the outer part of the plant, on exposure (Wyatt-Smith 1954). A purplish though mostly obtainedfrom the second­ stain is, in fact, left on the knife when cutting ary phloem. the phloem. This is a useful distinctive feature Syn.: bass (EM!). (De Rosayro 1953). Rollet (1982) mentioned (2) --+ I: fibre, phloem. that in some cases rapid discoloration takes (3) --+ I: phloem, secondary phloem. place in after slashing. I (4) --+ 1: phloem. have not found any mention of the colour of phellogen. The fonning dilatation See comments in Trockenbrodt (1990). tissue commonly is not included in the defini­ tion of cambium. cambium a meristem with products of divisions ar­ cork ranged orderly in parallel files; consists (1) a trade product which is mainly extracted of one layer of initials and their undiffer­ from the cork tree (Quercus suber). entiated products, or derivatives (Esau (2) --+ I: phellem. 1960*). See comments in Trockenbrodt (1990). Syn.: lateral (REC). Applied only to the two lateral mcristems: cortex vascular cambium: the meristematic (1) the tissue of primary origin that belongs cell layer responsible for the development neither to the , the periderm, or of the wood and/or the phloem. the phloem; between epidermis (periderm) Syn.: wood cambium (Ed), main cam­ and the vascular system (Trockenbrodt bium (Ed). 1990*). pbellogen: the meristematic cell layer re­ Syn.: primary cortex (Me, BM), peri­ sponsible for the development of the peri­ blem (Ja!). derm (Trockenbrodt 1990*). (2) --+ I: rhytidome (1). Syn.: bark cambium (Ed), cork cam­ See comments in Trockenbrodt (1990). bium (lAWA!, Esl!, Ja!, Br!, BCFT2!, Oul!, Fh!, Cha!, Whl!, SAP!, BM, dilatation tissue REC!, Ry, PZ!, Cg!, Ck!, Nu!, EM!), all tissue affected by dilatation growth cork meristem (Ja!). (Trockenbrodt 1990*) (Fig. 2). Cf: I: peridenn; I: phellodenn; I: phel­ Syn.: intercalary proliferation tissue lem. (Whl), expansion tissue (Wy). There are also other meristems which fonn Incl.: radial inclusions (Ji), wedges additional tissues: (Po). dilatation meristem: a radially oriented C!: lILa: dilatation (growth); lILa: meristematic cell layer in the phloem ray streaks. of some plants; responsible for adistinct­ Comments: ly funnel-shaped phloem ray dilatation Different parts of the tissue may undergo a (Trockenbrodt 1990*). dilatation process caused by the tangential Comments: strain which is a result of an increase in diam­ Meijer (1974), Raven et al. (1976), and eter of stem. Dilatation growth patterns, often Fahn (1990), following Esau (1960), use the with distinct colour, should be described by term 'cambium' in both senses. Some authors drawings and short notes when they fonn dis­ (Jackson 1928, BCFTI 1953, Wyatt-Smith cernible patterns in the phloem. Voorhoeve 1954, IAWA 1964, Den Outer 1967,Panshin (1965) points out that dilatation tissue often & De Zeeuw 1980) have restricted it to vascu­ appears in different-coloured and -textured lar cambium only. Vascular cambium is fre­ vertical bands and stripes. These markings quently inconspicuous. If visible, it is usual­ are good characteristics, as Rollet (1982) has ly translucent or watery, but, for instance in demonstrated in the family Annonaceae.

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periderm persistent bark, even though deep cracks may (1) the secondarily developed protective bark develop (cf. examples like Betula, Robinia, tissue replacing the epidermis, or built Sequoia; Esau 1960). According to Eames & during rhytidome formation; consists of MacDaniels (1947), generally several to many phellem, phellogen, and phelloderm times more phellem cells are produced than (Trockenbrodt 1990*). phelloderm cells, although there is a great Syn.: secondary cork (Oul !), corky deal of variation in different plants. Trocken­ layer (Ja), inner bark (Kr), occasion­ brodt (1990) restricts the term 'cork' to the ally used in the meaning of phellem trade product obtained from cork oak. (lAW A 1964), laminae in dead bark ('laminated dead bark') (Ji). phelloderm Cf: I: phellem; I: phellogen; I: phel­ a secondarily formed living tissue devel­ loderm. oped inwardfrom the phellogen,forming (2) the outer bark or epiphloem, ... cork cam­ a part ofperiderm and resembling cortical bium and its products (Jackson 1928*). . Syn.: middle bark (Ry!, WS, Wh2), Comments: secondary cortex (Whl!, Ck), cork The first-formed periderm originates most skin (RECl), outer edge (Wo!). commonly in the subepidermal layer. In older Cf: I: periderm; I: phelloderm; I: phel­ trees last periderm separates dead outer bark logen. from living inner bark which consists of sec­ ondary phloem (Esau 1960). Comments: The arrangement of periderm layers may The phelloderm may be thick- or thin­ vary with the species and/ or different parts of walled, and probably gives some protection the stem, when seen in cross section. Meijer against pathogens, heat and desiccation (Mar­ & Wood (1964) mention that some species of tin & Crist 1970). The phelloderm is usually Shorea may have differently coloured peri­ a thin layer and generally indistinguishable derms in the same tree, but generally they from the inner bark, except when it has a show a uniformly coloured periderm which distinctive colour (De Rosayro 1953). Wood is characteristic for the species. See discus­ (1952), Wyatt-Smith (1954), and Rollet sion concerning periderm in Trockenbrodt (1980) consider its colour generally a valu­ (1990). able feature for diagnostic purposes. Accord­ ing to Whitmore (1962a), it usually occurs as phellem a distinct, up to I mm thick, red-brown layer a secondarily formed protective tissue in of soft powdery tissue in Dipterocarpaceae. stems and roots consisting of dead cells This coloured layer, usually found under thin, with chiefly suberised walls; developed easily chipping rhytidome of some species, outward from the phellogen and forming may be streaky and sometimes the colour of a part ofperiderm. the just peeled piece may be different from Syn.: cork (EsI!, Ro, Rt, Ja!, IAWA!, the piece after longer exposure. This is a high­ Oul!, Fh!, MC!, Whl!, Cha!, Me!, ly diagnostic feature in such genera as Ko­ Ho!, SAF!, REC!, BM, Cg!, Ck!, koona and Lophopetalum (Whitmore 1972). EM!, Br!, BCFT2!), cork cortex (Ja), However, usually one finds under a thin rhy­ inner surface of the dead bark (Ji)? tidome a green layer, which is too common to Cf: I: periderm; I: phelloderm; I: phel­ be of diagnostic value (Rollet 1980). logen. Comments: phellogen, see under I: cambium. Suberised walls of phellem have waxy or fatty material resistant to the passage of gases phloem and water vapour, isolating the outer parts of = I: inner bark the bark. When developed, phellem frequent­ a principal assimilate conducting tissue ly forms a strong tissue and may result in a composed of different tissues and usually

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located outward of the xylem and inward scIerenchymatic elements: fibres, of the periderm. , and their intermediates forming Syn.: inner bark (WS, lAWA, MC, various structures in the secondary phlo­ Oul!, Ho, Cg, Ry!, REC, GJ, Br, em. Patterns thus formed may be seen in Ro, SAF!, PZ, Wo, Ed!, GE!, Wh2), cross and tangential section. bast (Ro!, SAF!, MET, Ed!, Ja!, BM, Syn.: hard bast (BM, Ro, Rt), hard Ry), inner rind (BM), live bark (Ro i), bark (IAWA), stereome (BM!), inclu­ liber (Ja, MET), bass (Fo). sions (fibrous -, grid-fibrous -, and Cf: I: bark, inner (3); bark, middle sandy inclusions) (Ji). (1) (rejected term). Note! Sieve elements, companion cells and phloem parenchyma are called soft Phloem can be divided into: bast, soft bark or leptome respectively primary phloem: phloem of primary by some authors. origin (Trockenbrodt 1990*). Cf: bast (2) (in rejected terms). Syn.: primary bast (Ja!), primary cor­ tex (Rt). phloem rays: rays traversing the phlo­ secondary phloem: all bark tissues de­ em (Trockenbrodt 1990*) rived from the vascular cambium during Inc!.: striated inner bark (WS). secondary growth (Trockenbrodt 1990*). Comments: Syn.: secondary bast (Ja), secondary The general arrangement of the tissues of liber (Ja), secondary cortex (Rt, Ja), the phloem is governed by the interaction be­ bast (Oul!, Ou2). tween the products of the vascular cambium Secondary phloem can be subdivided in­ and the dilatation tissue. According to Whit­ to two layers that are sometimes visible more (1972), phloem usually plays a role of to an unaided eye: little value for tree identification. However, non-collapsed secondary phloem: there are some aspects of morphology, col­ part of the secondary phloem with open our, texture, exudate, and smell which are and non-collapsed sieve-elements (Trock­ shown to be worthwhile. enbrodt 1990*). Wyatt-Smith (1954) suggests subdivision Syn.: functional phloem (Esl, Sa, of living phloem into 'outer inner bark' and REC), functioning phloem (Es3, Es4, 'inner inner bark' in some cases, correspond­ Fh), conducting phloem (Es2, Es4, ing to Wood's (1952) terms 'outer layer' and Wh 1, Rt), conducting zone (GJ), in­ 'inner layer', although he does not always ner inner bark (WS), inner bark (Ro, consider it feasible. The distinction is made Rt), inner layer (of the live bark) (Ji). on grounds of disparity in texture and colour. collapsed secondary phloem: part of Equally, Rollet (1980) and Roth (1981) dis­ secondary phloem where sieve elements, tinguish outer bark, inner bark, and middle companion or Strasburger cells, and bark within the phloem, although Rollet pre­ sometimes axial phloem parenchyma cells fers to avoid the term middle bark and, in­ are collapsed (Trockenbrodt 1990*). stead, speaks of outer bark with or without Syn.: non-functional phloem (EsI, the dilatation zone. However, Wyatt-Smith Sa, REC), non-functioning phloem (1954) and Whitmore (1972) adopted the (Es3, Es4, Fh), non-conducting phlo­ term middle bark in a completely different em (Es2, Es4, Whl, Rt), non-con­ sense, referring to an often coloured phello­ ducting zone (GJ), non-active phloem derm in some species. (Fh), outer inner bark (WS), outer Rather than adopting a highly confusing bark (Ro), outer layer (of the live bark) terminology as just discussed, a simple sub­ (Ji). division of phloem into non-collapsed and collapsed phloem may suffice. These terms Other structures in the secondary phloem correspond well enough to macroscopical that can sometimes be distinguished with aspects like colour, texture, and occurrence an unaided eye: of dilatation tissue. The term middle bark

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should be avoided, as Trockenbrodt (1990) a hard, tough bark (Carya), the fibres make proposed. up the greater part of the secondary phloem According to IAWA (1964) terminology, and provide considerable mechanical support hard bark sometimes occurs in concentric for the stem (Eames & MacDaniels 1947). See layers that alternate within a growth ring with Trockenbrodt (1990) concerning the terms layers of soft bark, or are more irregularly 'bast', 'bast fibre', and 'hard bast'. distributed in a of soft bark. Here such structures in the phloem are called rhytidome 'phloem with laminated texture'. (1) = I: outer bark In macromorphological studies it is some­ the dead outer part of bark including the times possible to distinguish between scleren­ last-formed periderm, cut off by this peri­ chymatic tissues and parenchymatic or dila­ dermfrom still living secondary phloem. tation tissues, mainly because of differences Syn.: outer bark (Esl!, Fh!, REC, Br, in colour. Therefore, the term sclerenchym­ Cg, Ed, MC!, Whl), bark (MET, Ja), atic elements is suggested for all sclerified cortex (MET), rind (Ja), outer rind structures observed in a cross section. Reli­ (BM), primary rind (BM), true bark able identification of different layers can be (Ed!, Ir), scale bark (EM!), shell bark obtained only in microscopy. (EM!), secondary bark (GE). The phloem rays are frequently much af­ Cf.: I: phloem. fected by the collapse of sieve elements and Note! In German literature, rhytidome associated cells, and by stem growth. The is referred to as 'Borke', which is lin­ wide phloem rays commonly become dila­ guistically close to bark. tated by cell division and tangential extension (2) all tissues outside the innermost phello- of cells. They remain radial, but they flare to­ gen. ward the periphery of the stem. The narrow Syn.: outer bark (Fh!, REC, WS, rays commonly become curved (Esau 1979). Oul!, Nu!, IAWA!, Le!, BCFT2!, Wyatt-Smith (1954) described striated in­ Wo, Th, Ry, Fo!, Wh2), bark (Ck, ner bark in which radial stripes of a different BM, Ro, Cha), true bark (PW!), colour, often white, are visible, for instance dead bark (Wo, Ro, Le!, Ba, Po, Ji). in Annonaceae. Furthermore, they are con­ (3) dead portion of the bark excluding last­ nected with the fissures in the rhytidome. formed periderm. Apparently these radial stripes are rays dila­ Syn.: outer bark (Ho!). tated in a different way. (4) superficial layer of the dead bark which detaches spontaneously (Tailfer 1989*). phloem fibre Comments: bark fibre of the phloem (Trockenbrodt Definition (1) follows more or less closely 1990*). Esau (1960), Martin & Crist (1970), and Fahn Syn.: bast (Ja, EM, BCFT2, Fo), (1990). Definition (2) is more or less similar bass (EM, BCFT2 !), bast fibre to that given by Chattaway (1953), BCFT2 (IAWA, PZ, Ja, Nu!, Oul, Es1), (1957), Penford & Willis (1961), De Saint hard bast (Nu, Ja), liber fibre (Jat). Aubin (1963), IAWA (1964), Den Outer Cf: II: texture fibrous. (1967), Ford-Robertson (1971), Nultsch Comments: (1971), Ghouse & Jamal (1978), and Letou­ Fibres are taxonomically important. When zey (1986). Definition (3) follows more or they occur in the secondary phloem, they are less that of SAF (1950), Bena (1960), Esau conspicuous elements because of their thick (1960), Howard (1971), and Trockenbrodt walls and characteristic distribution patterns (1990), and definition (4) Troupin (1978). (Esau 1979). They occur in a variety of form Rhytidome is composed principally of and arrangement, often in definite tangential dead masses of cortical and phloem tissue in bands (Liriodendron, ) or solitary the young stems, and secondary phloem in (Cephalanthus). In some woody plants with the old stems. Thickness of the rhytidome

Downloaded from Brill.com09/24/2021 07:42:43AM via free access Junikka -- Macroscopic bark tenninology 11 ------may be genetically controlled, or affected by Bena (1960) applies a French tenn \~corce the exposure to the sun or frequent fires. In morte' in the sense of the definition (2), and addition, structure and proportion of tissues restricts the term 'rhytidome' to the bark out­ may vary during tree growth, and thus affect side phellem. De Saint Aubin (1963) divides thickness of the bark. scaly or flaky rhytidome into two layers: Diagnostic characters of the rhytidome are outer rhytidome ('rhytidome externe') is a a rcsult of the weathering process, the tangen­ part which detaches easily when hit by a tial strain, the way of growing of the peridenn, machete. The inner rhytidome ('rhytidome the arrangement of the phellem, and the make­ interne') consists of thin layers adherent to up and amount of tissues cut off by the peri­ the tissues below. He adds that these rhyti­ denn from the bark. Typically, periderms are dome layers usually differ in colour. Srivas­ only thin layers in the former inner bark tis­ tava (1964) remarks that rhytidome origin­ sue, thus fonning only a small component of ates from the secondary phloem and, in such the whole outer bark. Occasionally periderm cases, may be considered merely a special tissue dominates the bark (Quercus suber), or tissue of the secondary phloem. there is only just one or occasionally two su­ Trockenbrodt (1990) proposes the replace­ perficial peridenns and therefore outer bark is ment of the tenn outer bark by the unambigu­ absent or 1-2 mm thick (Fagus sylvatica; ous tenn rhytidome. However, also the tenn Whitmore 1963). Wood (1952) mentions the rhytidome is used in different senses. I accept alternation of colour in bands formed by the the equivalent term outer bark for rhytidome, usually darker peridenn layers and the lighter because both terms are common in present coloured secondary phloem, which sometimes literature. Last-formed periderm should be occurs in species fonning successive peri­ noted, if there is a discernible colour differ­ derms. Mostly, however, colour is unifonn ence. or nearly so.

II. Bark texture (consistency) texture The texture may be described as fol­ composition of the bark largely deter­ .lows: mined by the character of the cells com­ corky: outer bark with the texture and posing the tissue, but also affected by the quality of cork. progress of decay of the rhytidome. Syn.: corky outer bark (WS). Comments: fibrous: outer and/or inner bark mainly The term texture is frequently used in composed offibres. wood, referring to its relative unifonnity to Syn.: fibrous bark (Es3, Ss, Ro), the touch or in the reaction to cutting tools. shreddy bark (RDMB), fibrous flakes Here it is applied to bark consistency. Tex­ (Le). ture is often complicated by the presence and IneZ.: fibrous inner bark (WS). distribution of sclereids and fibres set in a pa­ brittle: outer and/or inner bark is hard, renchymatous matrix (Wood 1952). A rough breakable. Outer bark may emit a metallic distinction between the texture of outer bark sound on cutting, inner bark may be fi­ and inner bark should be made when the brous but still brittle (Wyatt-Smith 1954, blaze is examined. In addition, the quality of Jimenez-Saa 1973). scales or flakes may be detennined. There is Syn.: crustaceous bark (Bo). much confusion in tenninology and different IneZ.: brittle outer bark (WS), fibrous­ categories of texture used by authors. It is breakable live bark (Ji), glass-break­ next to impossible to present a list of tenns able live bark (Ii), living bark with that are mutually exclusive. brittle fibres (Po).

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loose: outer and/or inner bark breaks up long strips in such trees as Aquilaria malaccen­ on cutting into coarse or fine grains (vs. sis and Artocarpus scortechinii. Esau (1969) firm). mentions that the presence of fibres "induces Syn.: crumbly bark (Ro). the bark to split in a reticulate, or netted, pat­ Incl.: granular flakes (Le), granular tern" (, Juglans regia). outer bark (WS), crumbly outer bark Brittle texture may include both sclereids (WS), chunky inner bark (Wh2!). and fibres. Bor (1953), who used the term granular: usually referring to inner bark 'crustaceous' for the black, hard, gritty bark, which is mainly composed of sclereids and Wyatt-Smith (1954) found brittle bark (Whitmore 1972). common in Diospyros. Cf: m.c: bark, stringy. In describing the loose texture authors mealy: outer bark falls off like powder have used many terms to describe the detach­ (Rollet 1980). ing of portions of the bark. Letouzey (1986) Syn.: mealy bark (Ro, Rt). applies the term 'granular' to flakes or scales Cf: m.c: bark, powdery. which break off easily. He gives examples like Milida excelsa (== Chlorophora) and Al­ homogeneous: either fibres or sclereids stonia boonei. Wyatt-Smith (1954) mentions occur (vs. heterogeneous) (Whitmore longitudinally broken, coarse granules, i. e. 1972). 'granular outer bark' in Shorea spp. He also Note! Do not confuse homogeneous limits the term 'crumbly' to outer bark which texture with homogeneous bark. breaks in small particles on cutting, and gives soft: outer and/or inner bark is soft and the family as an example. Rollet easy to cut (vs. hard) (Wyatt-Smith (1982) applies the term 'crumbly' also to 1954). similar qualities of the phloem, as distinct Syn.: soft bark (Ro) (vs. hard bark from fibrous texture (for example most of (Ro, Rt), compact bark (BKl), stone the family Chrysobalanaeeae). However, he bark (BM». shows that the distinction between fibrous Incl.: soft outer bark (WS), soft inner and crumbly inner bark is somewhat arbitrary bark (WS) (vs. hard inner bark (WS». because closer examination reveals merely laminate: concentric, cylindrical or inter­ sclereids and fibres in varying quantity. Roth rupted layers in the phloem formed by (1981) characterises brittle and crumbly con­ sclerenchymatic elements. sistency, mentioning that it consists of "stone Syn.: laminated bark (MW, Ro), lam­ cell groups which separate easily from the inated inner bark (WS), laminated live surrounding parenchymatous tissue." She bark (Ii), long strings (of live bark) also stresses that the more advanced the pro­ (Ji), several layers (Ji), layered pattern gress of decay is in the rhytidome, the more (in living bark) (Po). crumbly and mealy it will be. Mealy texture Incl.: superposed plates (Rt), tangen­ can be detected by rubbing between fingers, tial bands (Rt), concentric rings (Rt). for example in Chaetocarpus schomburgkia­ Cf: lILa. ripple marks; L dilatation nus (Rollet 1982). tissue. Homogeneous layers may be found in the Comments: phloem, although barks may be chiefly het­ In addition to rhytidome, the development erogeneous in texture. This should be noted of phellem is stimulated by forest fires. Rollet from the blaze. Whitmore (l962a) found that (1982) observed very spectacular ridges, over smooth barks are fibrous inwards and granu­ 10 cm thick, in Caesalpinia coriaria, a species lar outwards, and described the cause of this from Llanos. Sometimes corky outer bark in terms of the great amount of tangential may be easily detached by hand (Rollet 1982). strain the smooth barks have to accommodate. Wyatt-Smith (1954) found corky texture in The term 'soft' (vs. hard) texture should outer bark of Pithecellobium confertum. be restricted to an evaluation of cutting with According to Wyatt-Smith (1954), barks reference to outer and! or inner bark, without with fibrous texture can often be peeled off in regard to anatomical structure. Inner bark has

Downloaded from Brill.com09/24/2021 07:42:43AM via free access Junikka - Macroscopic bark terminology 13 ------usually a yielding texture, often moist, for bark: Anisoptera spp. Jimenez-Saa (1973) example in most Annonaceae. Wyatt-Smith distinguished three types of laminated tex­ (1954) mentions examples of soft outer bark, ture, which are hardly applicable (see rejected for instance certain Litsea spp., and Campno­ terms). Roth (1981) elaborated a classifica­ sperma auriculatum. tion according to distribution of sclerenchym­ Trees with hard texture in the bark occur atic tissue, where sclerenchymatic tissue may in open and semideciduous forests affected be visible as superposed plates, tangential by regular fires, but also in closed forests. bands or continuous and concentric rings in The hardness of the rhytidome is variable, the phloem structure. She also called these and depends on the amount of phellem and patterns 'hard bast types'. sclereids (Rollet 1982). Also special inclu­ Concentric laminate structures are useful in sions, such as Ca-oxalate crystals, in a large describing a blaze in the field. For instance, number may result in difficulty of cutting Meijer & Wood (1964) identified layers of (Roth 1981). Bark may be so hard and reso­ stone cells alternating with other layers of nant, for instance in resiniferum, bark which they consider "a good character to that a clear ringing sound may be heard if distinguish some species of Dipterocarpa­ struck with a smart blow with the back of a ceae." The term laminate texture is related to dao (= a large heavy knife used by the Bur­ the ripple marks, often meaning the same, but mese) (Bor 1953). Inner bark is hard and separated here, because in ripple marks rays then usually dry, for example in lxonanthes are sometimes involved. Moreover, the term icosandra (Wyatt-Smith 1954). ripple marks is primarily used in de­ Laminate texture refers to phloem where scribing tangential sections. laminae are formed through dilatation tissue Whitmore (1962a) distinguishes a laminat­ which contains sclereids in tangential layers. ed bark type in Dipterocarpaceae because of Wyatt-Smith (1954) and Whitmore (1962c) the laminated structure in the inner bark. See mention one example of such laminated inner the definition in the list of rejected terms.

III. Bark patterns

III.a: Bark patterns in cross and tangential flame-marks section a pattern resembling flames formed by phloem rays seen in the cross section of corrugations phloem (Rollet 1982). the inner sUrface of the bark is corrugated Cf: lILa: phloem, scalariform. matching the similar pattern on sapwood. Comments: Syn.: corrugated sapwood (Fs, Ro). This is a variant of scalariform phloem. Comments: Rollet (1982) gives, among others, Anacar­ De Rosayro (1953) and Jimenez-Saa (1973) dium giganteum and Scaphium macropodum used the term in this sense. When the bark is as examples. removed, one may see a wrinkled or corru­ gated sapwood surface like corduroy or the phloem, mottled ridged formation of a washing board. It was phloem which is marked with spots of found that large aggregate rays of the wood various colours or shades in tangential terminate in the indentations (Francis 1951). section. Syn.: mottled inner bark (WS), mot­ dilatation (growth) tled living bark (Po). the process ensuring the tangential wid­ Comments: ening of the bark during growth (Trock­ Wyatt-Smith (1954) gives examples, such as enbrodt 1990*) (Fig. 2). Endospermum malaccense, and Polak (1992) See comments under I: dilatation tissue. Aspidosperma cruentum. No distinction is

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Fig. 2. Close-up of the blaze where dilation growth seen in the blaze of Catostemma fragrans, Guyana (photo A.M. Polak). - Fig. 3. Ripple marks in Tabebuia insignis var. monophylla, Guyana (photo A. M. Polak).

made between tissues involved in mottled pat­ ripple marks terns. The term mottled is restricted to spots fine, parallel, horizontal lines in the tan­ as seen in the blaze, and the term patchy to gential section of wood or bark, caused bark surface patterns. by the storied structure of all the wood and bark elements, or by the distribution in horizontal layers of the wood rays phloem, scalariform and bark rays only (Voorhoeve 1965*) a pattern formed by phloem rays in (Fig. 3). the cross section of phloem resembling Cf: II: texture laminate; ill.a: streaks. ladder-like structure with radial 'rungs' (Rollet 1982). Comments: Cf: lILa: flame-marks. BCFT2 (1957), IAWA (1964), Meijer Comments: (1974), and Panshin & De Zeeuw (1980) re­ This is the structure formed by phloem strict the term to tangential and longitudinal rays which separate other secondary phloem surfaces of certain . Symington (1943) parts. Rollet (1982) gives a number of ex­ gave examples of diagnostic ripple marks in amples, for instance Scottellia coriacea and sapwood, viz. ferrea, Balanocarpus Khaya ivorensis. heimii (= Neobalanocarpus). Kunkel (1965)

Downloaded from Brill.com09/24/2021 07:42:43AM via free access Junikka - Macroscopic bark tenninology 15 ~------~~------~------uses it to describe horizontal lines in cambium (2) longitudinal grooves with width of the (Stockwerkbau). Rollet (1982) compares it in ridges between the grooves at least three his glossary with "markings similar to small times that of the grooves or fissures ridges on the sand of the seashore." The tenn (Wyatt-Smith 1954*). is close to the term laminate structure, but (3) ridges separating fissures are twice or kept separate here because the tenn ripple more times as wide as fissure (Jimenez­ marks has a broader definition which may be Saa 1973*). useful in describing wood characteristics in (4) longitudinal grooves with width less than the blaze as well. flat-topped ridges separating them (Tail­ fer 1989*). streaks Comments: striations on the suiface of the blaze usu­ In barks with successive peridenn fonna­ ally formed by phloem rays and scleren­ tion, the expansion growth of the stem causes chymatic tissues. cracking of the rhytidome. More or less lon­ Streaks may be: gitudinal ridges and grooves are fonned. Fis­ longitudinal: longitudinal strzatlOns sure formation depends on certain internal against different-coloured background. properties such as a fibrousness, ray dilata­ Syn.: striated inner bark (WS), streak­ tion, peridenn formation, etc. Irregular fis­ ed inner bark (Ry), striped structure suring is often the result of fonnation of (Ro). scales which shed only partly (Roth 1981). reticulate: regular or wavy, net-like stri­ The bark should be studied in full-grown ations against a different-coloured back­ trees, otherwise results are not reliable. ground. Definitions (2), (3), and (4) are very nar­ Syn.: gritty inner bark (Ry), grid­ row. In addition to these, some other types of fibrous inclusions (Ji). fissuration are described like 'fluted bark' Comments: (Wyatt-Smith 1954; followed by Den Outer De Rosayro (1953) applies 'streaks' only 1972), and 'ridged bark' (see rejected tenns). to longitudinal structures. He gives Wormia The definition of ridged bark does not differ triquetra (= Dillenia) as an example, which much from the term fluted bark which more has an appearance similar to coconut wood. likely seems to be just a variety of ridged Wyatt-Smith (1954) uses the tenn for radial bark. In literature the tenn 'fluted' is also ap­ stripes of a different colour, often white (An­ plied to a stem, its characteristic shape is nonaceae). Apparently these are identical fonned by unequal secondary growth. with phloem rays. Rollet (1982) mentions De RosayrQ (1953) classified fissured bark numerous examples, grouping them accord­ into five types and some sub-types according ing to dominant colour of the tangential sec­ to the closeness of fissures, ridges or the tion. shape of fissures, and also the mode of Reticulate patterns are fonned by phloem cracking (with some examples). He does not rays and tangential layers of phloem fibres. name these types. Furthennore, Rollet (1980) Rollet (1982) mentions, for example, Esch­ distinguishes 'fine-fissured', 'coarse-fissur­ weilera corrugata (= Lecythis) and Terminalia ed', 'deep-fissured', 'shallow-fissured' and superba. 'wavy-fissured', but does not give any defi­ nitions. He suggests that a more refined clas­ sification of fissured barks should consider III. b: Fissuring the width of ridges and grooves. The patterns of the outer surface of the bark, fissured bark are often extremely difficult to describe. (1) bark cracked lengthwise into fissures sep­ Here, perhaps even more than with other fea­ arated by ridges. tures, authors use descriptive terms without Syn.: striated bark (Ja, Ss, Ku), plated defining them properly. Therefore much con­ bark (RDMB). fusion and overlap exist. I recommend to de-

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7 8

Figs. 4-9. Schematic drawings of different fissure and ridge types as seen in cross section. - 4: Fissures are deep and ridges rounded. - 5: Fissures are shallow and irregular. - 6: Square­ shaped fissures. - 7: Compound fissures. - 8: Flattened ridges. - 9: Hollow ridges.

scribe fissures and ridges separately with boat-shaped: oval or elliptical fissures measures of their average width. If used in which are not continuous (Fig. lO). the given sense, the descriptive terms accept­ Syn.: boat-shaped fissured bark (WS, ed in the present paper should be sufficient. MW), vertical crevices (Le), boat­ shaped cracks (Ry). fissures: more or less longitudinal short: less than 15 cm long (Wyatt­ grooves between ridges in the rhytidome. Smith 1954). They may be classified according to depth long, elongated: more than 15 cm long and length as follows: (Wyatt-Smith 1954). deep: at least as deep as half of the total thickness of the bark (Fig. 4). In addition, fissures can be classified into Syn.: deep-fissured bark (Ro, GJ, different morphological types (see Whit­ YYI), fissured bark (Po), furrowed more 1962a, Yunus et al. 1990): bark (Ro!), deeply furrowed bark (Hi, REC), deeply cracked bark (Th), V -shaped: more or less acute fissures ridged bark (Ro!), rugged bark (Ro!), sometimes penetrating inner bark, lined slitted fissuring (Ro), slotted fissur­ by special associated periderm( s) (See ing (Ro). Fig. 4). shallow: less than half as deep as the Syn.: acute fissures (WS), V section total thickness of the bark (Fig. 5, see fissures (Wh 1). also Figs. 6, 8, and 9). round: the outer surface is concave in Syn.: shallow-fissured bark (GJ, cross section (Wyatt-Smith 1954). YYI), shallow-furrowed bark (Hi), square-shaped: flat bottom fissures, al­ fine-fissured bark (Ra), scaly-fissur­ ways restricted to the dead outer bark ed bole (Whl), cracked bark (Ro, (Fig. 6). Po, Ji), vertically cracked bark (WS), Syn.: square fissures (WS), square shallow-cracked bark (Th). section fissures (Whl).

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irregular: different-sized gaps or fur­ Comments: rows in the bark surface (Fig. 5). Syn.: irregular section fissures (Whl). The definitions of 'deep fissures' and compound: anastomosing shallow fis­ 'shallow fissures' are followed more or less sures which are formed in the bottoms of closely by Yunus et al. (1990). Wyatt-Smith existing fissures (Fig. 7). (1954) and Rollet (1980) use the tenn 'crack­ Syn.: compound section fissures ed bark' which describes the bark that just (Whl). begins to open. They divide cracked bark wavy: longitudinal grooves that are into 'grid-cracked' (~IILc: bark, tessellated) coarse with wavy, more or less irregular and 'vertically cracked bark'. These cracks, faces difficult to define clearly (derived less than 1 cm wide, are continuous or non­ from De Rosayro [1953] and Wyatt­ continuous. Whitmore (1962a) used the tenn Smith [1954]) (Fig. 11). 'scaly-fissured bole' to indicate the intenne­ Syn.: wavy fissured bark (WS, Ro), diate fonn between 'smooth' and 'shaggy­ coarse fissured bark (Ro). scaly bole'. Moreover, he described bark types like 'deep fissured bark', and 'shallow­ Fissures may be: fissured bark' (in Dipterocarpaceae), which parallel: grooves are parallel, long, usu­ are bark types as a whole. See the definitions ally regular. The resulting, long ridges in the list of rejected terms. may crack transversally (De Rosayro Boat-shaped fissures are generally a sign 1953) (Fig. 12). of young barks, where cracks or fissures are oblique: grooves are short or long, anas­ just opening because of the tangential expan­ tomosing but not so regular and distinct sion growth of a stem. However, boat-shaped as in reticulate fissured barks. The result­ fissures can be deep. De Rosayro (1953) and ing ridges may crack transversally. Wyatt-Smith (1954) give Calophyllum spp. reticulate: grooves join each other and as examples. divide again irregularly leaving non-con­ Wyatt-Smith (1954) classified grooves or tinuous and sometimes obscure ridges. fissures according to shape in cross section: Cf: m.b: ridges reticulate; m.c: bark, shallow or deep, round (surface in form of tessellated. hemisphere), acute (with converging faces), or square (with parallel faces). The terms ridges: more or less continuous raised 'coarse-fissured' and 'pitted bark' were used parts of the rhytidome between fissures. by RolIet (1982) in his glossary, but without Ridges may be: a definition, however. flattened: the outer surface is plane in cross section (Fig. 8). Ridges may be described by their width, hollow: the outer surface is concave in length, height or depth, shape, or the way cross section (Wyatt-Smith 1954) (Fig. 9). they are sloughed. They may also fonn dis­ Syn.: hollow-ridged bark (WS). tinctive patterns, such as reticulation. Besides rounded: the outer surface is convex in the use of shape and regularity, De Rosayro cross section (Fig. 4). (1953) proposed to describe the distance V -shaped: the outer surface is sharp in between ridges (widely vs. closely spaced cross section (de Rosayro 1953). ridges). Hightshoe (1989) mentions, within reticulate: ridges join each other and the context of 'shallow-furrowed bark', that irregularly divide again enclosing non­ narrow ridges occur, for instance, in Lirio­ continuous fissures (Wyatt-Smith 1954) dendron tulipifera, and wide ridges in iu­ (Fig. 14). glans cinerea. Ridges may also become Syn.: reticulate-ridged bark (WS), cracked across into elongated, rectangular, bark like stretched net (Le), interlaced platy segments coming close to tessellated strands (Ro), diamond-shaped pattern bark. Wyatt-Smith (1954) gives an example (Po). of 'hollow-ridged bark', viz. Shorea lepro­ Cf: IILb: fissures reticulate. sula.

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I1I.c: Scaling ring bark and scaly bark. Whcn flakes are de­ tached, a pattern of differently coloured areas bark, flaky; flakes may result. See III.c: patchy bark. (1) bark which has large and variable patches De Rosayro (1953) uses 'flakes' for large or flakes ofrhytidome, more than 7.5 cm rectangular pieces ( zeylanicus), long, which become detached (Wyatt­ and 'scales' for small pieces like scales of Smith 1954*). fish (D. hispidus), but does not indicate the Syn.: scale (Le!, Co!), sheet bark borderline between both terms. Letouzey (BM), shaggy bark (Ro!), exfoliating (1986), and Corner (1988) use flakes inter­ bark (RDMB, Hi). changeably with scales. Polak (1992) origi­ IneZ.: ringed bark (RDMB). nally follows De Rosayro's concept distin­ Cf: IILc: bark, scaly; III.c: bark, guishing flakes and scales, but he relates patchy; IILc: ring bark. flakes to thickness of the stem. Hence a (2) a bark with numerous very thin bark scales flake, with a size of 10 x 15 cm from a tree (Voorhoeve 1965*). 0.2 m thick is regarded merely a scale on a (3) Meijer (1974) follows definition (2), but tree I m thick. Relating the size of flakes to calls barks also flaky when "scales flake the thickness of stem may lead to confusion, off". though, since the thickness of the trunk will (4) ___ IILc: bark, scaly. increase during the life-span of the tree. (5) --- lILc: bark, patchy. Wyatt-Smith (1954), Jimenez-Saa (1973), and Rollet (1980) recognise papery bark, Flakes may be (see also Rollet 1980, with very thin pieces peeling off like paper. Letouzey 1986): They do not indicate the size of the pieces of rectangular. bark falling off. However, their examples, irregular. among others, Melaleuca leucadendron and circular. certain Eugenia spp., suggest that in their pub­ papery: thin and very flexible flakes lication, in fact, flakes are described. (vs. slab-like flakes). Hightshoe (1989) uses the term 'exfoliat­ scrolled: long, thin flakes which are roll­ ing bark' to describe thin or coarse strips, or ed up by their edges (Fig. 16). sheets peeling either vertically (such as Thuja Syn.: scrolled bark (WS, Ro, Wh2), occidentalis), horizontally (Betula lutea = B. scroll-like dehiscence (Ry), peeling alleghaniensis). Brooker & Kleinig (1990b) bark (Co, Kr, REC), horizontally I describe 'minniritchi bark', which is a special vertically exfoliating bark (Hi), min­ type of eucalypt bark that is split both longi­ niritchi bark (BK2). tudinally and horizontally, the free edges of shaggy: loosened, usually slightly curv­ the flakes rolling back while still adherent ed rectangular or irregular flakes which to the trunk to uncover new green bark be­ may hang for a time on the stem (Whit­ neath. more 1962a) (Cf. Fig. 12). Rollet (1980) used the term 'shaggy bark' Syn.: shaggy bark (REC, Ro, Kr), for Chlorophora excelsa (= Milicia) and Dip­ flaky bark (Ro!), shaggy-scaly bole terocarpus sublamellatus. (Whl). Comments: bark, heterogeneous Flakes are produced when bark develops more than one type of bark is encoun­ large sheets of periderm that are intermediate tered in the same stem (Fig. 13). in size between the parts shed in the cases of Cf: II: texture heterogeneous.

Fig. 10. Bark with boat-shaped fissures in Duguetia uniflora, Guyana (photo A.M. Polak). - Fig. 11. Bark with developing wavy fissures in Bridelia micrantha, . - Fig. 12. Bark with parallel fissures in Terminalia amazonia, Guyana (photo A.M. Polak). - Fig. 13. Hetero­ geneous bark in boehmii, Malawi.

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Comments: (2) named after stem appearance when seen Surface patterns of the stem may depend from a distance; fissures are square­ on the developmental stage of the bark. The shaped or irregular and scattered or close appearance of bark from the base of the tree (seldom); the width of fissures is lO (25) upwards may change considerably. I found a mm, but they do not penetrate into the typical example in an African species, Brachy­ inner bark; sloughing of the bark is con­ stegia bussei, where scaly zones alternated tinual, usually in large scales up to c. 50 with fairly smooth ones. cm long, which are not formed in scol­ lops: scales are elongate or isodiametric, bark, patchy layer-thick, loose or adherent, and occur lighter blotches on the outer surface of scattered or in zones (abridged from Whit­ the rhytidome resulting from the irregular more 1962a). dehiscence of old rhytidome plates, usu­ (3) thin papery flakes separated by a mesh ally two colours dominate on the bark pattern of fine shallow vertical and hori­ (Rollet 1980, Letouzey 1986) (Fig. 18). zontal fissures (Hightshoe 1989*). Syn.: mottled bark (Ro), marble-like Scales: blotches (Le), blotched bark (Ja), spot­ According to persistence one may rough­ ted bark (Le), flaky bark (Kr!), patchy ly divide them as follows (see also Whit­ exfoliating bark (Hi). more 1972): Comments: adherent: retained some time on the Kriissmann (1984) gives examples like stem. Platanus, Acer, Stewartia, and Parrotia. The loose-hanged: shed seasonally (Fig. colour of patches where flakes or scales were 19, see also Fig. 13). recently shed, may give some additional in­ formation and should be reported with the Scales may be roughly divided according use of colour charts. However, the differen­ to density as follows (see also Whitmore ces in colour may be caused by , espe­ 1972): cially on smooth barks. close. distant. bark, powdery Scales may be roughly divided according bark covered with a fine powder-like to shape as follows (Whitmore 1962a, crust which usually rubs off easily. It is 1972, Rollet 1980, Letouzey 1986): mostly associated with smooth barks rectangular. (Wyatt-Smith 1954). irregular. circular. Comments: papery: thin and very flexible. This is a result of a usually long process flat-sided: one or several layers thick. of decay caused by weathering. Bor (1953) chunky: usually with irregular rough gives some characteristic examples, viz. Apo­ faces and irregularly chunky shaped. rusa spp. and Bischofiajavanica. scollop-shaped: thickest in the middle, tapering to the edges, and leaving a scal­ bark, scaly; scales lop-shaped depression on the bole on (1) bark which has small more or less irreg­ sloughing (ef. Fig. 22). ular patches or scales of rhytidome, less Syn.: plates (Ji). than 7.5 cm long, which become detach­ scrolled: thin scales which are rolled up ed (Wyatt-Smith 1954*). by their edges. Syn.: scaly peeling bark (Th), flaky shaggy: loose and usually slightly curv­ bark (Ti), alligator bark (Ir). ed, rectangular or irregular scales which Inc!.: ringed bark (RDMB). may hang for a time on the stem (Fig. 19). Cf: m.c: bark, t1aky; III.c: ring bark; Syn.: shaggy-scaly bole (Whl), shag­ IILc: bark, tessellated. gy bark (REe, Ro, Kr).

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Fig. 14. Bark with reticulate ridges in . - Fig. 15. Tesselated bark in Dio­ spyros kaki. (Figs. 14 & 15 are redrawn from Vaucher 1990). - Fig. 16. Bark with scrolled flakes in Eucalyptus citriodora (redrawn from Thrower 1988). - Fig. 17. Bark with mussel­ shell markings in Afzelia bracteata (redrawn from Voorhoeve 1965).

Comments: Voorhoeve (1965) defines scales simply Definition (2) applies to a bark type of as "thin pieces of bark". According to Meijer Dipterocarpaceae. See more details and draw­ (1974), scaly bark "can be dippled-scaly with ings in Whitmore (1962a). For thc third defi­ oblong scales, or tessellate, like some spe­ nition Hightshoe (1989) gives Ostrya virgini­ cies of Barringtonia." However, he calls bark ana as an "indicator" species. 'flaky' when "scales flake off". Irvine (1952) Scaly barks vary much in detail. Many has described very thick bark with large scales trees have fissured bark with scaly ridges, with the term 'alligator bark', for example in and in many cases one cannot distinguish Butyrospermum parkii (= Vitellaria para­ between scaly barks and fissured barks, or doxa). Roth (1981) does not distinguish scales scales are visible on ridges. Therefore it is and flakes, but instead uses the terms small practical to describe shapes of scales and fis­ and large scales. sures separately.

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Downloaded from Brill.com09/24/2021 07:42:43AM via free access Junikka - Macroscopic bark terminology 23 bark, stringy op as localised wound responses caused by thick, loose, long-fibred bark, never de­ insect of attack. Bark has an appear­ ciduous (Penford & Willis 1961 *). ance like rotting from the surface inwards, Syn.: stripping bark (Co), fibrous and there is a lot of variation in surface fea­ peeling bark (Th), fibrous strips (Le). ture between trees and even parts of a tree. Cf: II: texture fibrous. Whitmore (1962c) mentioned some Shorea species having this type of bark. Comments: This definition is followed more or less closely by Rollet (1980). Beard (1944) noted bark, tessellated "ropy strings" in such species as Vitex diva­ surface marked by more or less regular, ricata and Citharexylum spinosum. The term square or oblong plates or blocks re­ has been applied, moreover, to certain rough­ maining for a long time on the stem (Fig. barked eucalypts. This texture arises, accord­ 15). ing to Chattaway (1953), through the expan­ Syn.: cracked bark (Co), reticulately sion of the phloem parenchyma, separating scaly bark (Ss), grid-cracked bark bundles of fibres from one another in the (WS, Wh2, Ro, Ji), crocodile-skin rhytidome. Brooker & Kleinig (1990a) add bark (Bo), gritty bark (Po), blocky that stringy bark is thick, spongy, deeply and bark (Hi). widely furrowed, interlaced beneath, with Cf: III.c: bark, scaly; lII.c: bark, outer layers weathered to grey or grey brown. flaky. This is a highly diagnostic feature. Comments: The definition follows more or less close­ ly De Rosayro (1953), Hall et al. (1970) and bark, surface rotten Brooker & Kleinig (1990a). This is a special bark is shortly fissured (with fissures type of bark, where rhytidome blocks are usu­ varying in depth and cross section), scaly, ally retained. The rhytidome is short-fibred, rugose or smooth; sloughing is very vari­ breaking up into small plates. According to able scales being small, adherent, chunky Brooker & Kleinig (1990a), plates may be or flat-sided; in a transverse section the hard and woody (in Eucalyptus, viz., paper­ inner edge of the outer bark is following fruited bloodwoods and some boxes), firm or surface configurations, not parallel to spongy (in woody-fruited bloodwoods), and cambium (Whitmore 1962a). flaky (in yellow bloodwoods). The scales Cf: III.b: bark, fissured (1); m.c: are usually thick and adherent. Wyatt-Smith bark, scaly (1), (2); lII.c: bark, het­ (1954) applied the term grid-cracked to a erogeneous. special case of cracked bark with fissures Comments: which are just beginning to split, of which A special type of the bark, which has a Rollet (1980) gives examples like achroma complex surface pattern. According to Whit­ lagopus and Nauclea diderichii. I consider all more (1962a), the thickness of outer bark these terms as synonyms for tessellated bark. and undulations of its inner edge are govern­ Hightshoe (1989) mentions Diospyros virgi­ ed by the amount of rotting at the bark sur­ niana as an "indicator" species of 'blocky face. Moreover, the periderms (if any) devel- bark'.

Fig. 18. Patchy bark of Brachystegia bussei, Malawi. - Fig. 19. Bark with loose-hanged, shaggy scales in Pterocarpus rotundifolius var. polyanthus, Malawi. - Fig. 20. Rugose bark of Brachystegia glaucescens, Malawi. - Fig. 21. Smooth bark with in vertical lines (Oco­ tea tomentella), Guyana (photo AM. Polak).

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Downloaded from Brill.com09/24/2021 07:42:43AM via free access Junikka - Macroscopic bark tenninology 25 ring bark is 'ring rhytidome' or 'annular rhytidome'. (1) a type of rhytidome in which periderms Nevertheless, ring bark is a well-established may be formed parallel to the first one. tenn, so the change of the tenn is not follow­ The concentric cylinders of bark thus ed here. formed result in a rhytidome which de­ taches annually with large sheets (Esau III.d: External markings 1960, Rollet 1980). Syn.: very smooth bark (Ro), smooth bark, dippled bark (BKl). (1) bark covered with shallow, usually cir­ Cf: scale bark, bark, peeling (1) (both cular depressions, more than 1 cm in in rejected tenns). diameter. These are scars of the scaled­ (2) the outer bark when disengaged in strips off old bark (Wyatt-Smith 1954*) (Fig. or layers (Jackson 1928*). 22). Syn.: jigsaw puzzle (like -) (Ro!,MW), Comments: dippled-scaly bark (Co). The rhytidome is shed annually, either IneZ.: oyster shell markings (Ro). once in a season or in several phases, and the Cf: III.c: scales scollop-shaped; III.d: resulting dead bark accumulates at the base of bark, pock-marked; HI.d: scroll marks. the trunk in certain trees (Brooker & Kleinig (2) named after stem appearance seen from a 1990a). The bark falls off in one piece or in distance when the bark surface is dippled large pieces or strips rolled up by their longi­ or smooth with scattered scollop-shaped tudinal edges (Rollet 1980). Srivastava (1964) scales and scroll marks; sloughing occurs has shown that if the first periderm was deep­ as scollops, small ones being entire, large seated, the subsequent periderms follow the ones in pieces (abridged from Whitmore outlines of the external periderm, i.e., they 1962a). encircle the axis and, consequently, form ring bark. Some barks are intennediate be­ Comments: tween ring bark and scaly bark, so outer lay­ Definition (1) is followed more or less ers of rhytidome are detached as relatively closely by De Rosayro (1953) and Rollet large flakes (for example in Platanus, Arbu­ (1980). Yunus et al. (1990) use the tenn when tus) (Esau 1965, Fahn 1990). Esau (1969) shallow depressions may be visible on the mentions that the bark detaches as flakes surface of the bark when slashed. Jimenez­ (Betula pendula), or as large plates (Platanus Saa (1973) and Polak (1992) apply the tenn acerifolia) when sclereids are numerous. Usu­ 'dippled' to more or less round depressions ally fibres are then lacking or only scarce. without distinguishing pock-marked bark Most barks are of the scale bark type with from dip pled bark. Rollet (1980) employs the fissuring different from the rare ring bark term 'oyster shell markings' describing de­ type. The number of peridenns developed in pressions deeper than covered by the term the rhytidome, and their arrangement (paral­ dippled bark in the sense of Wyatt-Smith lel, or ramified and anastomosing) are deci­ (1954), but here 'oyster shell markings' is sive for scale formation; size, thickness, tex­ included in dippled bark. Whitmore (1962a, ture, shape, and outlines of the scales. Wood 1972) uses the term in a slightly different (1952) used the term 'peeling bark' for all meaning. According to him, the surface of a kinds of barks. dippled bark is patchy in colour, sloughing Roth (1981) proposes a new term for ring off as small, round, usually close-set, scol­ bark which is a literal translation of the lop-shaped scales like in Pometia and Swin­ Gennan word 'Ringelborke'. Her suggestion tonia. The term 'dippled' (or more precisely

Fig. 22. Dippled bark of sp., Malawi. - Fig. 23. Bark with eye marks (Maca­ ranga capensis), Malawi. - Fig. 24. Bark with ring-grooves (Myrica salicifolia), Malawi. - Fig. 25. Prickles on the bark (Bombax malabaricum) (redrawn from Thrower 1988).

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'dippled-scaly') was originally introduced by Comments: Comer. According to him, 'dippled-scaly Anatomically lenticels are structurally dif­ bark' has small, round, thin, and very nu­ ferentiated portions of the periderm which are merous flakes looking like irregular discs characterised by a relatively loose arrangement which leave closely set scars on the bark sur­ of cells, where gaseous exchange takes place face (Melanorrhoea malayana = Gluta). (Esau 1965). Their width varies from I mm The size and thickness of the scales, and to 10 mm (Comer 1988). Lenticels may per­ colour of old and newly exposed surface may sist for many years in barks with superficial give some diagnostic information. It seems periderm layers, like in Betula and practicable to describe barks by the term dip­ avium, widening due to the thickening of the pled in combination with scroll-marks and stem. On barks with successive periderm pock-marked bark. layers, however, the lenticels are difficult to notice in the fissures. On thick, homogene­ bark, lenticellate; lenticels ous dead barks they tend to be lost, or at least barks which have on the surface more or are inconspicuous (Eames & MacDaniels less raised, often somewhat corky spots 1947, Wood 1952). According to Busgen & or lenticels (Fig. 21). Munch (1929), fissures of the bark usually Syn.: pimpled (pimply) bark (Ro), initiate from the lenticels, as can be seen in pustulate bark (Vo, Ku, Ja). Populus. Here a combination of characters is pre­ Lenticels are good field characters for iden­ sented which authors, as Wood (1952), tification. Their size, colour, and density Wyatt-Smith (1954), Rollet (1980), and should be noted. A rough estimate, like nu­ Roth (1981), considered useful. merous/scarce, is enough. However, they may occur relatively more abundantly on Lenticels: some parts of the bark than on others. U sual­ The form of lenticels may be according to Iy they are most conspicuous on young the greatest diameter: twigs. linear: usually horizontally elongated like buttonholes (rarely vertical). round: (usually small). bark, pock-marked stellate: star-shaped. bark covered with small shallow circular diamond-shaped: rhomboid. depressions, less than I cm in diameter The size of lenticels may be divided ac­ (Wyatt-Smith 1954*). cording to the greatest diameter (Jimenez­ Syn.: pock-marking (Wo), dimpled Saa 1973): bark (Ss), eye-marked bark (Ro!), large: more than 5 mm in diameter. pock-pitted bark (Ro!), pitted bark medium: 3-5 mm in diameter. (Vo, Dr), pitting (Wo!), spotted bark small: less than 3 mm in diameter. (Le). Cf: III.d: bark, dippled. Lenticels may occur on bark: numerous (vs. scarce) (only when Comments: present in large numbers or only few). De Rosayro (1953) reported this pattern solitary (vs. compound). (together with dippled bark) from a variety of in vertical lines (Fig. 21). Syzygium species. Pock-markings occur on in horizontal lines. the outer edge of the live bark (~ I: phello­ in oblique lines. derm). They affect the surface of the bark as According to their consistency they may seen on separation of the dead bark (Wood be: 1952). Meijer & Wood (1964) draw attention soft. to this diagnostic feature describing it as a powdery. "bark with small irregular flakes and a rough compact. appearance."

Downloaded from Brill.com09/24/2021 07:42:43AM via free access Junikka - Macroscopic bark terminology 27 ------=------bark, rough Comments: bark which has an uneven surface; the This condition may also be seen on the term includes various scaly, flaky and smooth upper trunk of a tree that has a rough fissured barks. bark on lower parts. The feature is diagnostic Cf: III.d: bark, rugose; III.d: bark, in some cases. smooth.

Comments: bark, smooth Wyatt-Smith (1954) has a similar defini­ (1) bark is even, thin, unbroken, though it tion. Rough bark patterns can be seen from a may be bumpy or pimply from the lenti­ distance without closer inspection. eels (Corner 1988*) (Fig. 21). Syn.: close bark (Dr). bark, rugose Cf: III.d: bark, lenticellate; m.d: bark, (I) bark which is covered by wrinkles, de­ rugose. pressions, small irregular scales, shallow (2) named after stem appearance seen from fissures; bark appears smooth from a a distance; bark is thin (1-2 mm), usual­ distance (Fig. 20). ly without rhytidome; bark surface is en­ Syn.: wrinkled bark (Vo!, Rt), sculp­ tire, often with raised hoop and eye tured bark (Rt!), little rough bark (Le), marks; sloughing is absent or inconspicu­ smooth bark (WS, Ou2, Ry, Wh2), ous with powdery appearance or tiny smoothish bark (Rt). (1-2 mm) scales (abridged from Whit­ Cf: IILd: bark, smooth; III.d: bark, more 1962a). rough. (3) epidermis persistent, no outer layer of (2) small, medium or large, close or scatter­ dead cells, surface smooth, no cracks or ed lenticels or minute cracks (Whitmore peeling (Thrower 1988*). 1972*). (4) -7 IILd: bark, rugose (1). Syn.: firm bark (Dr). (5) -7 III.c: ring bark. (3) covered with wrinkles, small depressions, furrows or ridges (Den Outer 1972*). Comments: Comments: The definition (1) is more or less simi­ The definition (I) is comparable with that lar to that by Radford et al. (1974), RoBet by Roth (1981). Most rugose barks are thin. (1980), Kriissmann (1984), Letouzey (1986), Whitmore's (1972) definition of rugose Hightshoe (1989), and Yunus et al. (1990). barks falls within his own circumscription of Smooth barks may have hoops, eye-marks, smooth barks. Roth (1981) also distinguishes prickles, and other types of excrescences, but fairly smooth or smoothish barks which not scales or fissures. Certain species may "arise through exfoliating of the cork or have the same appearance throughout the through continuous desquamation of very whole life of the tree, with lenticels and a small scales", but here they are considered unique periderm. A thin phellem layer may rugose barks. Meijer & Wood's (1964) and then cover the surface of the stem (Rollet Meijer's (1974) definitions of 'non-fissured 1980). bark' (see rejected terms) are close to rugose The definition (2) is a description of a bark. bark type of Dipterocarpaceae which was Rugose appearance is a common feature studied in detail by Whitmore (1962a). This in young developing barks, but the terminol­ corresponds well with the first definition, ogy is often obscure and difficult to interpret whereas Thrower (1988) in definition (3) ap­ precisely from the context. plies the term only to the epidermis, for ex­ ample in species of Ficus. Thus he does not bark, scribbly apply the term to bark in the pure sense. smooth bark carries characteristic 'scrib­ Diagnostic characteristics, e.g. eye marks, bles' caused by insect larvae (Brooker & prickles, etc. should be noted, although the Kleinig 1990a*). bark is regarded otherwise as smooth.

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burls distribution is too irregular for scars, but hard woody excrescences on a tree, more they may sometimes be formed from branch or less rounded inform, usually resulting scars. from the entwined growth of a cluster of adventitious buds (SAF 1950*). hoops Syn.: burr (Ja), gnaur (Ja). (1) raised transverse rings which partially Cf: IILd: warts. or completely encircle the stem (Cf. Fig. Comments: 24). The definition is more or less similar to Syn.: hooped bark (WS, Ro, Ry), that by Jackson (1928) and Sutton & Tinus horizontal rings (Po), hoop marks (1983). Panshin & De Zeeuw (1980) describe (Whl), ring-like marks (Ro). it as "a bulge formed on the trunk or branches Inc!.: hoop-rings (Th), hooped bark of a tree by abnormal wood formation." (Ji), ring-hooped bark (Ji), horizontal As some forest dictionaries give a wider folds (Le). definition for burr (see rejected terms), the Cf: IILd: ring-grooves. term burl is preferred here. (2) ~ III.d: eye marks. Comments: cankers Jimenez-Saa (1973) applies hooped bark (1) definite, relatively localised, necrotic le­ to partial hoops and ring-hooped bark to sions primarily of the bark and cambium complete hoops. Rollet (1980) does not dis­ (SAF 1950*). tinguish hooped and ring-grooved bark in the (2) ~ IILd: warts. text, although in his glossary he mentions two separate terms, viz. hooped and ring-like Comments: marks. Wyatt-Smith (1954) gives Sindora The definition follows Jackson (1928) and spp. as an example of hooped bark. BCFTl (1953) more or less closely. The These hoops usually occur on smooth­ term is confined to injuries or wounds of the barked trees. Their origin is obscure, but, ac­ bark without diagnostic significance. cording to Thrower (1988), they are associ­ ated with eye-marks. Jimenez-Saa (1973) eye marks regards them as scars left by fused deciduous stipules. Whitmore (1962a) considers their eye-shaped marks on trunks and branches distribution too irregular for leaf scars. of smooth-barked trees (Thrower 1988*) (Fig. 23). Syn.: hoops (Rt). mussel-shell markings Cf: III.d: hoops. parallel, sinuous, concentric ridges on the bark left after shedding of the rhyti­ Comments: dome (Fig. 17). Roth (1981), referring to eye marks as Syn.: parallel, sinuous ridges (Le). hoops, mentions that they mainly correspond Cf: III.d: bark, dippled. to leaf scars or scars of lateral branches. As a result of expanding dilatation growth they Comments: partly or completely surround the stem. She This is a very conspicuous and diagnostic considers that complete "hoops", i.e. eye feature in some barks. Voorhoeve (1965) marks, originate from decussate phyllotaxis, published a fine photograph of mussel-shell or from very broad leaf bases. Thrower markings on the stem of Afzelia bracteata. (1988) follows this explanation. Whitmore Polak (1992) has found the character in Inga (1962a) has a definition comparable with that alba, and mentions it in the identification key by Thrower, but he suggests that the origin on bark and slash characters, but he uses an­ of eye marks is by the distortion of hoop other term ('oyster shell markings') in the marks formed on the stem. He adds that their text, which is incorrect.

Downloaded from Brill.com09/24/2021 07:42:43AM via free access Junikka - Macroscopic bark tenninology 29 ------prickles Syn.: thorn (Le!, Ro, Ja, Dr, RDMB, (1) sharp outgrowths from the bark, detach­ REC, Ti), prickle (WS). able without tearing the wood (Voorhoeve Cf.: m.d: prickle (1); excrescence, 1965*) (Fig. 25). pointed (rejected tenn). Syn.: cork prickle (Rt). Spines may be (Letouzey 1986): Cf.: m.d: spine. simple (vs. branched). (2) ~ IILd: spine. straight (vs. curved). See comments under spines. hard (vs. supple). tapering. ring-grooves conical. transverse grooves which partially or Comments: completely encircle the stem (Fig. 24). The distinction between non-woody 'prick­ Syn.: ring-grooved bark (WS), rill les' and woody 'spincs' is followed by many (Ku). authors. Defined in this way, they may have Cf.: IILd: hoops. diagnostic value. Prickles occur in such spe­ Comments: cies as pentandra, Fagara macrophylla The definition is similar to those of Kunkel (= Zanthoxylum), Hura crepitans, and Jaca­ (1965) and Radford et al. (1974). ratia. Spines can be found for instance in Ba­ In one African tree, Myrica salicifolia, lanites wilsoniana, Bridelia grandis, Flacour­ about 8 m tall, I found curious-looking ring­ tia rukam, and formosum (Rollet grooves, the bark exfoliating by small, irreg­ 1980). Some authors do not make any differ­ ular scales. Ring-grooves were densely dis­ ence between prickles and spines (Bor 1953). tributed from the tree base up to the crown Roth (1981) describes pure cork fonna­ and branches. These grooves were fonned in tions or cork prickles originating from phel­ the phloem, as the blaze did not show any logen. Their appearance is variable, i. e. a grooves in the sapwood. I have not found roundish ring-like base with sharp-pointed any reports of this feature in literature about end, or flattened. They may occur on the Myrica species. Wyatt-Smith (1954) mentions base of stem, on the upper side of twigs, Shorea parvifolia as an example of ring-groov­ sometimes densely crowded (Hura crepitans). ed bark. Roth regards these as an excellent diagnostic character. However, De Rosayro (1953) does scroll marks not rate occurrence of prickles or spines, al­ raised sinuous marks between depres­ though a notable character in trees of Ceylon, sions which are formed when scollop­ very highly, because this is mainly a juvenile shaped scales are detached (Whitmore feature. 1962a) (Fig. 22). Wyatt-Smith (1954) defines prickles as Syn.: scroll-marked bark (Wh2). similar to spines (stiff, sharp, woody, narrow, Cf.: m.d: bark, dippled. long processes), but thicker and wide-based. Besides 'thorn' Timberlake (1980) uses the Comments: tenn 'knob' for raised woody protuberances The term 'scroll marks' is close to the on a trunk, often with a thorn on top (Acacia). tenn dippled bark. Whitmore (1962a) uses 'Pointed excrescences' (Letouzey 1986) the tenn scroll-marked (bark) or scroll marks resemble structures known as root spines! when speaking of smooth barks. This bark spine roots, which gradually develop into type is close to dippled bark. aerial roots with a sharply pointed apex, for instance in Bridelia micrantha and some spines palms (Jenik & Harris 1969). These spine sharp hardened outgrowths of the bark formations are uncommon in dicotyledons, and wood, usually modified branches or providing useful diagnostic infonnation, how­ ; tearing the wood, if detached. ever, in certain cases.

Downloaded from Brill.com09/24/2021 07:42:43AM via free access 30 IAWA Journal, Vo1.15 (1),1994 warts hard or firm excrescences other than len­ Comments: ticels, spines, prickles, or burls which The definition is more or less similar to that are formed on the stem. by Jackson (1928) and Voorhoeve (1965). Syn.: bumpy excrescence (Le), can­ Warts may be, for instance, adventitious root kered bark (WS), verrucose bark (Ja!, buds which are just appearing on the main HJC!, Vol). stem. Warts are distinctly smaller than burls Cf: III.d: burls; III.d: bark, ­ which are usually formed by a cluster of ad­ late. ventitious buds.

W. Exudation blaze, dry sticky: adhesive and sticks to the fingers the surface of the blaze is not exuding, (vs. non-sticky). feels dry when touched. Example (sticky): Hevea brasiliensis Syn.: dry live bark (Ji). (Wyatt-Smith 1954*). odorous: smells pleasantly, for instance blaze, wet like incense or wintergreen (Rollet 1982). the surface of the blaze slightly exuding, smelly: smells more or less unpleasantly, feels moistened when touched (Rollet for instance like garlic, pepper, sulphur, 1980). excrement (Rollet 1982, Letouzey 1986). Syn.: succulent live bark (Ji). abundant: exudation is profuse for a Comments: while (vs. scanty). When a blaze is wet, the exudate is usually Cf: IV. blaze, wet. a watery ooze from sieve elements. Note! The Comments: degree of wetness of the blaze depends partly The definition more or less closely fol­ on the season in which the bark is slashed. lows Wood (1952), Wyatt-Smith (1954), Den Outer (1972), Jimenez-Saa (1973), Rollet exudate (1980), and Letouzey (1986). moisture or liquid of the living tissue, Certain families are characterised by their which seep or flows after slashing, usu­ exudates (such as in Burseraceae). In ally from the inner bark. other families different species may have Purely physically an exudate may be: their own type of exudate, or even certain clear: transparent (vs. opaque) (Wyatt­ species may show more than one type of ex­ Smith 1954). udate (Wood 1952). De Rosayro (1953) re­ Examples (clear): Myristicaceae, certain gards the nature and type of exudation "the Calophyllum spp. (Wyatt-Smith 1954). most important single character of diagnostic Examples (opaque): Artocarpus elasticus, value in identification." Some authors, e. g. certain Anacardiaceae (Wyatt-Smith 1954). Wood (1952), Rollet (1980, 1982), Letouzey coloured: may be white, yellow, gold­ (1986), have recommended that such diag­ en, brownish, red, blackish. nostic characters as a colour, discoloration, discoloured: the colour changes within coagulation (both after few minutes expo­ a few minutes. sure), smell, taste, speed with which it is frothy: forms foam when rubbed be­ oozed (in a continuous stream or in droplets), tween fingers (Rollet 1980). and how it is distributed in layers, should be liquid: flows readily, often transparent. noted in the description of exudates. How­ Syn.: sappy exudate (Ku, Le, REC, ever, Rollet (1980) values the colour of exu­ Ja), watery exudate (WS,Ji). dation most. Discoloration may also give viscous: flows slowly, not necessarily useful hints for identification. sticky. De Rosayro (1953) has reserved the term Syn.: thick exudate (Ji). 'watery exudate' for thin, clear or coloured

Downloaded from Brill.com09/24/2021 07:42:43AM via free access Junikka ~ Macroscopic bark terminology 31 ------~ exudates (see rejected terms). He gives ex­ refer to kino as reddish resin. However, Hall amples of the red or reddish type, for instance et al. (1970) include it in gums which develop Myristicaceae, and of the colourless type, for in the cambial region of Eucalyptus, often as instance Campnosperma zeylanicum and Man­ a result of injury. The terms 'gum' and 'kino' gifera zeylanica. Wyatt-Smith (1954) defined are used indiscriminately in the literature as the same term as an exudate resembling water Hillis (1987) has pointed out. According to (Antiaris toxicaria, Melanorrhoea = Cluta). Hillis (1987), kino-bearing species are, for I am inclined to refer to the exudate by its example, Butea frondosa (= B. monosper­ physical properties, e. g. colour, quality, etc., ma), Pterocarpus marsupium, and some instead of using "traditional" terms like latex, Eucalyptus species. He also mentions that resin, ctc. For instance latex is not always appearance of kino may be characteristic of whitish, or resin brownish, and both include the species, and remarks that during spring colourless types as well as gums. The chemi­ and early summer it changes increasingly to­ cal nature of exudates does not correspond ward red. well with the visual characterisation of tradi­ tional terms (cf. Hillis 1987). However, I latex accept some "traditional terms" because they (1) latex is a colloidal suspension or emul­ are widely used, and, if defined properly, are sion of water-insoluble substances, sus­ descriptive to a certain degree. pended in an aqueous phase. It is typi­ cally white (milky), but may be yellow to gum red, or colourless (Hillis 1987). (I) viscid secretions, exuding naturally or on Syn.: lactiferous exudate (Ry). incision or infection, generally colourless, (2) the milky juice or exudate of certain plants: nontoxic, odourless, and usually tasteless; it is always opaque and of a white or yel­ and on desiccation, or exposure to air, low colour (Wyatt-Smith 1954*). they become hard, clear, glassy masses Cf: exudate creamy (rejected term). (Hillis 1987*). Syn.: mucilaginous exudate (Ss, Ja), Comments: gelatinous exudate (Ja). Definition (1) is more or less similar to (2) a comprehensive term for non-volatile, those given by Eames & MacDaniels (1947), viscous exudates from many plants and Esau (1960), Jimenez-Saa (1973), and Comer trees, which either dissolve or swell up (1988). Latex is found in a considerable num­ in contact with water; of complex chemi­ ber of angiosperm families, and is a source of cal structure, they may be considered to many economic products. The suspensions be highly polymerised carbohydrates may be of teipenoids (e.g. rubber), proteins, (BCFT2 1957*). essential oils, mucilage, and other compo­ nents (Hillis 1987, Eames & Mac Daniels Comments: 1947, Esau 1960). Comer (1988) regards it The definition is similar to those given by as very useful in the identification of many Jackson (1928), Voorhoeve (1965), Den plants; it may change colour on exposure, its Outer (1972) and Rollet (1980). Hillis (1987) consistency being generally sticky, rubbery, gives examples like Acacia and Sterculia. resinous or gummy, but in some cases it is watery and even brown in colour. De Rosayro kino (1953) employs the term lactiferous exudate newly formed kino is usually of a thick for an opaque and milky consistency. He consistency similar to that of honey, distinguishes three categories of colour: ranging from straw to blood-red in col­ (i) creamy or dirty white on exposure (for our; ... and dries to a brittle, semi-trans­ example Artocarpus spp.), (ii) pure white parent solid (Hillis 1987*). (majority of ), or (iii) coloured Comments: (Carcinia). AHDE (1980) and BCFT2 (1957), when Definition (2) is more or less similar to dealing with several Old World tropical trees, that used by Jackson (1928), SAF (1950),

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BCFT2 (1957), .Kunkel (1965), Voorhoeve era in Burseraceae), mastics (Pistacia len­ (1965), Den Outer (1967), Rollet (1980), tiscus var. chios), and lacquer (Rhus Thrower (1988), and Tailfer (1989). Accord­ vernicifera =R. verniciflua). ing to Wyatt-Smith (1954), 'creamy exudate' (2) an exudate which is adhesive, clear and (see rejected terms) occurs in Alstonia spp. colourless, or pale yellow to dark brown and Sapotaceae. or black in colour (dammar) (Wyatt-Smith 1954*). Syn.: resinous exudate (WS). resin IneZ.: resinous exudation (Ry), tarry (1) colourless exudate or light brown glossy exudate (Ry) (see rejected terms). solids, which are stable, inert and amor­ (3) a thick, syrupy or gummy exudate of the phous and become sticky when heated bark, generally clear or slightly coloured: (Hillis 1987). (i) yellow (Calophyllum), (ii) whitish or Hillis (1987) divides broadly into colourless (Canarium zeylanicum); both two main types: types are always accompanied by a strong oleoresins: a solution of resin acids in aromatic scent (De Rosayro 1953). turpentine; oleoresin is a pale yellow, Cf: exudate tarry (rejected term). clear and sticky mass, which upon eva­ (4) very sticky sap, usually solidifying on poration of volatile compounds becomes exposure (Kunkel 1965*). brittle. Oleoresins exudes from the resin canals Comments: of wood and bark when exposed in coni­ The definition of oleoresin is more or less fers and Canarium luzonicum. similar to that used by Jackson (1928) and 'natural resins': they can be colour­ BCFT2 (1957). Definition (2) is followed less, as with dammars, or light brown more or less closely by Den Outer (1972) and glossy solids; they are stable, inert and Rollet (1980), definition (3) by Iimenez-Saa amorphous, become sticky when heated (1973), and definition (4) by Storrs (1979) and are fusible (above lOODC) with no and Tailfer (1989). sharp melting points. The main difference between these defini­ Resins or resin exudates that are stored tions is found in colour. De Rosayro (1953) on the hark or in cavities. They arc found distinguished also a 'tarry exudate' which in several genera of Dipterocarpaceae is related to the definition (2) (see rejected and Leguminosae/Caesalpiniaceae. Well­ terms). It is characteristic of the genus Seme­ known resins are dammars, copals (hard, cmpus. Wyatt-Smith (1954) includes the tarry from different genera, for example Copai­ exudate in resin and gives Dipterocarpus spp. fera spp.), elemis (soft, a numher of gen- as an example.

V. Bark cutting blaze Comments: (I) = slash The term has heen used in the first sense a shallow Cllt removing a portion of'the by SAF (1950), BCFTI (1953). De Rosayro bark and wood so as to aid ill species (1953), and Rollet (1982). Originally 'slash' identification (Figs. 2 & 3). was used for "the bark slashed off by a cut", Syn.: slash (Vo. Ou2. Ku. Wh I, Wh2!, and the wound on a trunk, and especially the Le, WS, Woo Ry!, Ro!, BCFTI!. light coloured exposed sapwood was known Bo!, Po!), cut (WS', Wh2', Ry', Ro', as 'hlaze' (Wood 1952). Bo!. Po'). A slash or hlaze reveals the eolour of the q: IV: hlaze, dry/wet. inner layers of hark, sapwood (hardwood) (2) -'> I: hark, inner (3). and exudate, a possihle discoloration when

Downloaded from Brill.com09/24/2021 07:42:43AM via free access Junikka - Macroscopic bark terminology 33 exposed to the air, texture, smell, taste, etc. slash (Letouzey 1986). De Rosayro (1953), evalu­ (1) = blaze. ating the nature of exudation as the most im­ (2) residual logging debris (SAF 1950, portant and specific character of the blaze, HeFTI 1953). points out that a blaze should be made on the stem and not on a buttress because of tissue Comments: differences. Letouzey (1986) mentions some Slash is accepted here as an alternative other precautions to be observed when noting term for blaze. The term slash is commonly the appearance of bark: age of the tree, habi­ used also in this second sense. However, there tat of the tree, exposure of the bark, and the is no danger of confusion. height at which the sample is taken.

List of rejected terms

bark, alligator -7 IILc: bark, scaly. bark, dimpled -7 m.d: bark, pock-marked. bark, blocky -7 IILb: bark, tessellated. bark, dippled-scaly -7 IILd: bark, dippled. bark, blotched -7 IILc: bark, patchy. bark, exfoliating bark, cankered -7 m.d: warts. - horizontal -7 lILc: bark, flaky. - vertical -7 lILc: bark, flaky. -7 bark, chunky inner II: texture granular. - patchy -7 lILc: bark, patchy. bark, close -7 m.d: bark, smooth (1). bark, eye-marked -7 IILd: bark, pock­ bark, coarse fissured -7 III. b: bark, fissured, marked. fissures wavy. bark, fibrous inner -7 II: texture fibrous. bark, compact -7 II: texture hard. bark, fibrous peeling -7 m.c: bark, stringy. bark, cracked -7 IILb: bark fissured, fissures bark, fine fissured -7 III. b: bark, fissured, shallow; -7 m.c: bark, tessellated. fissures shallow. bark, crocodile-skin -7 IILc: bark, tessel­ lated. bark, firm -7 IILd: bark, rugose (2). bark, crumbly -7 II: texture granular. bark, fluted - deep, continuous, round-bot­ tomed longitudinal grooves with rounded bark, crustaceous -7 II: texture brittle. ridges between them; width of ridge and bark, dead -7 I: rhytidome (2). width of groove approximately equal and - laminated, see I: periderm. total width more than 2.5 cm (Wyatt-Smith 1954*). See m.b: bark, fissured. bark, deep fissured - fissures are coarse and close; the size and section of fissures are bark, furrowed -7 IILb: bark, fissured, fis­ very variable, but fissures do not pen­ sures deep. etrate into inner bark; ridges are flat-top­ bark, granular outer -7 II: texture granular. ped; sloughing occurs continually; chun­

ky small scales (few cm) are adherent bark, grid-cracked -7 IILc: bark, tessellat- (abridged from Whitmore 1962a). See ed. IILb: bark, fissured. bark, gritty -7 lILc: bark, tessellated. bark, deeply cracked -7 IILb: bark, fissured, fissures deep. bark, gritty inner -7 lILa: streaks reticulate. bark, deeply furrowed -7 III.b: bark, fissured, bark, hard -7 II: texture, hard; -7 I: phloem fissures deep. (with) sc1erenchymatic elements.

Downloaded from Brill.com09/24/2021 07:42:43AM via free access 34 IAWA Journal, Vol. 15 (1),1994 bark, hard inner --t II: texture hard. bark, living - (1) the part of the bark outside the vascular cambium up to and including bark, hooped --t m.d: hoops. the last-formed periderm (Chan 1985*). See I: bark, inner. bark, inner - (2) -7 I: bark, inner (3). - inner --t I: phloem, non-collapsed sec­ ondary phloem. bark, mealy -7 II: texture mealy. - outer --t I: phloem, collapsed second­ ary phloem. bark, middle - (1) in the phloem between in­ ncr and outer bark, sometimes visible; it is bark, laminate - named after slash appear­ the place where the division and differen­ ance; bark is scaly fissured; ridges are 2-6 tiation of cells occur, and its relative thick­ cm wide with several unevenly overlap­ ness varies in the live bark depending on a ping flat layers; fissures are usually irregu­ species (Roth 1981 *). lar in section, short; bark looks as though - (2) -7 I: phelloderm. rotting from surface inwards; tissues loose, bark, minniritchi --t III.c: bark, flaky, flakes soft; in this type a sclerosis plays a role scrolled. in which tangential sheets of sclerified tis­ sues (close compound laminae) are formed bark, mottled --t III.c: bark, patchy. to varying extent in the phloem prolifera­ tion tissue; layers can be seen in cross bark, non-fissured - (1) bark is not fissured, section (abridged from Whitmore 1962a). but may still be cracked, flaky or pock­ See II: texture laminate. marked and not really smooth (Meijer & Wood 1964*). See m.d: bark, rugose. bark, little rough -7 m.d: bark, rugose (1). - (2) Totally smooth or scaly, or covered with lenticels (Meijer 1974*). Sce III.d: bark, live bark, rugose. - dry --t IV: blaze, dry. - fibrous-breakable -7 II: texture brittle. bark, peeling - (1) all species with successive - glass-breakable -7 II: texture brittle. periderm layers show more or less peeling, - inner layer (of the live bark) -7 I: phloem, from time to timc, of one or more cork non-collapsed secondary phloem. layers; this may be haphazard, scasonal or - laminated --t II: texture laminate. controlled by external conditions such as phloem contains very thin, flexible lami­ frost, fire or insolation; ... form of peeling nae of less than 1 mm; one can separate is directly related to mode of laying down wide and and long laminae with fingers of periderms (Wood 1952*). See scale (Jimenez-Saa 1973). bark (rejected terms); IILc: ring bark. - (with) long strings -7 II: texture lami­ - (2) -7 IILc: bark, flaky, flakes scrolled. nate. refers to laminae of live bark, which do bark, pimpled (pimply) -7 m.d: bark, lenti­ not break easily and which can be pulled cell ate. out as long strings, for instance in some species of Annonaceae and Lecythidaceae bark, pitted -7 IILd: bark, pock-marked. (Jimenez-Saa 1973). bark, plated -7 IILh: bark, fissured (1). - (with) several layers --t II: texture lami­ nate. bark, pock-pitted -) IILd: bark, pock­ laminae, which are not easily separable, marked. and are in some cases about 2 mm thick (Jimenez-Saa 1973). bark, pustulate -) m.d: bark, lenticellate. - outer layer (of the live bark) -7 I: phloem, collapsed secondary phloem. hark, reticulately scaly -) IILc: bark, tessel­ - succulent --t IV: blaze, wet. lated.

Downloaded from Brill.com09/24/2021 07:42:43AM via free access Junikka - Macroscopic bark terminology 35 ------bark, ridged - (I) deep continuous, longi­ bark, spotted -) IILc: bark, patchy; -) IILd: tudinal grooves often with flat-topped bark, pock-marked. ridges between them: width of the ridge and width of the furrow approximately bark, stone -) II: texture hard. equal, and total width more than 2.5 cm; e.g. Shorea curtisii (Wyatt-Smith 1954*). bark, streaked inner -) lILa: streaks longitu­ See m.b: bark, fissured. dinal. - (2) -) IILb: bark, fissured, fissures deep. bark, striated -) m.b: bark, fissured (1). bark, ringed -) IILc: bark, flaky; -) IILc: bark, striated inner -) I: phloem, phloem bark, scaly rays; -) III.a: streaks longitudinal. split or cracked bark with circular fissures (Radford et al. 1974*). bark, stripping -) m.c: bark, stringy. bark, rugged -) IILb: bark, fissured, fissures, bark, true -) I: rhytidome (1), (2). deep. bark, verrucose -) III.d: warts. bark, sculptured -) III.d: bark, rugose (1). bark, winged - bark with one or more thin, bark, secondary -) I: rhytidome (1). flat longitudinal expansions or elongate plates (Radford et al. 1974*). See m.c: bark, shaggy -) IILc: bark, flaky; -) IILc: bark, scaly. bark, flaky, flakes shaggy; -) IILc: bark, scaly, scales shaggy. bark, wrinkled -) m.d: bark, rugose (1). bark, shallow-cracked -) lII.b: bark, fissured, bass -) I: bast (1); -) I: phloem fibre; -) I: fissures shallow. phloem. bark, shallow-fissured - named after bole ap­ bast -) I: phloem fibre. pearance from a distance; outer bark is a - hard -) I: phloem, sclerenchymatic coherent mass united by periderms; V-sec­ elements; -) I: phloem fibre. tion fissures are long and often penetrat­ - primary -) I: phloem, primary phloem. ing into the inner bark; ridges are 25-50 - secondary -) I: phloem, secondary mm wide and fissures about one-third of phloem. this; sloughing is infrequent, scales are to - soft, see I: phloem (note under sc1eren­ 5 x 15 cm in size, ridge-wide, layer-thick chymatic elements). or chunky, and adherent (abridged from Whitmore 1962a). See IILb: bark, fis­ bast fibre -) I: bark fibre; -) I: phloem fi­ sured. bre. bark, shallow-furrowed -) m.b: bark, fissur­ blaze -) bark, inner (3). ed, fissures shallow. bole bark, sheet -) IILc: bark, flaky. - scaly-fissured -) IILb: bark, fissured, fissures shallow. bark, shell -) I: rhytidome (1). - shaggy-scaly -) IILc: bark, flaky, flakes shaggy; -) m.c: bark, scaly, scales bark, shreddy -) II: texture fibrous. shaggy. bark, soft -) II: texture soft; see I: phloem burrs - (1) lar$e excrescences on a tree trunk (note under sc1erenchymatic elements). or major bnanch, and to the enlarged root

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stock found in certain trees; the grain is expansion tissue ~ I: dilatation tissue. highly contorted and presents a character­ istic type of figure (SAF 1950, BeFTI exudate 1953, and Ford-Robertson 1971). See - creamy - latex which is thick in consis­ m.d: burl. tency and opaque but not sticky (Wyatt­ - (2) ~ III.d: burl. Smith 1954*). See IV: latex (1), (2). - gelatinous ~ IV: gum (1). cambium - lactiferous ~ IV: latex (1). - bark ~ I: phellogen. - mucilaginous ~ IV: gum (1). - cork ~ I: phellogen. - resinous ~ IV: resin (2). - main ~ I: cambium, vascular cam- - sappy ~ IV: exudate, liquid. bium. - tarry: black, tarry, acrid exudation, :... wood ~ I: cambium, vascular cam­ which is usually slow and appears ini­ bium. tially as black dots or streaks which later coagulate; the stem often shows black, cork tar-like blotches on its exterior (De Ro­ - secondary ~ I: periderm (1). sayro 1953). See IV: resin (3). - spongy ~ I: phellem. - thick ~ IV: exudate viscous. - watery: cork prickle ~ m.d: prickles (1). (1) thin, usually clean or resembling a so­ lution of colouring matter in water which cork skin ~ I: phelloderm. is subdivided into (i) red or reddish exu­ date, (ii) colourless exudation (De Rosayro cortex 1953). See IV: exudate. - cork ~ I: phellem. (2) ~ IV: exudate liquid. - primary ~ I: cortex (1); ~ I: phloem, primary phloem. fissures - secondary ~ I: phloem, secondary - acute ~ m.b: bark, fissured, fissures phloem; ~ I: phelloderm. V-shaped.

cracks, boat-shaped ~ III.b: bark, fissured, fissuring fissures boat-shaped. - slitted ~ m.b: bark, fissured, fissures deep. crevices, vertical ~ III.b: bark, fissured, fis­ - slotted ~ III. b: bark, fissured, fissures sures boat-shaped. deep. cut ~ V: blaze = slash. flakes - fibrous ~ II: texture fibrous. edge - granular ~ II: texture loose. - inner: thin and often inconspicuous lay­ er, which is the very young phloem and cambium of the current growing season folds, horizontal ~ III.d: hoops. (Wood 1952*). See I: cambium; I: phlo­ em, non-collapsed secondary phlo­ gnaur ~ m.d: burl; ~ burr (rejected terms). em. - outer ~ I: phelloderm. hoop marks ~ m.d: hoops. excrescences hoop-rings ~ m.d: hoops. - bumpy ~ m.d: warts. - pointed: long, thick, woody outgrowth inclusions from the trunk (Letouzey 1986). See - fibrous ~ I: phloem, scIerenchymatic m.d: spines. elements.

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- grid-fibrous --7 III. a: streaks reticu­ - conducting --7 I: phloem, non-collaps­ late; --7 I: phloem, sclerenchymatic ed secondary phloem. elements. - non-active --7 I: phloem, collapsed - radial --7 I: dilatation tissue. secondary phloem. - sandy --7 I: phloem, sclerenchymatic - non-functional --7 I: phloem, collapsed elements. secondary phloem. - non-functioning --7 I: phloem, collaps­ knob, see III.d: spines. ed secondary phloem. - non-conducting --7 I: phloem, collaps­ layer ed secondary phloem. - corky --7 I: periderm (1). - inner: corresponds to the younger phlo- plates --7 IILc: bark, scaly, scales scollop­ em and rays without youngest phloem, shaped. which is a thin and often inconspicuous - superposed --7 II: texture laminate. layer (Wood 1952). See I: phloem, non­ collapsed secondary phloem. proliferation tissue, intercalary --7 I: dilata­ - outer: corresponds to the older phloem tion tissue. and rays with remnants of cortex and peri­ cycle according to the age and species ridges, parallel, sinuous, concentrical --7 m.d: (Wood 1952). See I: phloem, collapsed mussel-shell-markings. secondary phloem. rill --7 m.d: ring-grooves. leptome, see I: phloem (note under scleren­ chymatic elements). rind --7 I: bark (1); --7 I: rhytidome (1). - inner --7 I: phloem. liber --7 I: phloem. - outer --7 J: rhytidome (1) - secondary --7 I: phloem, secondary - primary --7 I: rhytidome (1). phloem. rings, horizontal --7 m.d: hoops. liber fibre --7 I: phloem fibre. scale bark markings, oyster-shell --7 m.d: bark, dip­ (1) a type of rhytidome in which the sub­ pled. sequent periderms develop as restricted overlapping strata, each cutting out a marks, ring-like --7 II!.d: hoops. scale-like mass of tissue (Esau 1960*). See IILc: ring bark; II!.c: bark, flaky; meristem IILc: bark, scaly; bark, peeling (rejected - cork --7 I: phellogen. terms). - lateral --7 I: cambium. (2) --7 I: rhytidome (1). periblem --7 I: cortex (1). stereome --7 I: phloem, sclerenchymatic ele­ ments. pitting --7 m.d: bark, pock-marked. structure, striped --7 lILa: streaks longitudi­ phloem nal. - functional --7 I: phloem, non-collaps­ ed secondary phloem. thorn --7 m.d: spines. - functioning --7 I: phloem, non-collaps­ ed secondary phloem. wedges --7 I: dilatation tissue.

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GLOSSARY OF THE SUGGESTED MACROSCOPICAL BARK TERMS

I. Bark and its component tissues (Fig. 1)

bark - all tissue outside the vascular cam­ ph ell em - a secondarily formed protective bium regardless of its composition. tissue in sterns and roots consisting of dead cells with chiefly suberised walls; bark, inner = phloem. developed outward from the phellogen and forming a part ofperiderm. bark, outer = rhytidome. phelloderm - a secondarily formed living bark fibre - fibre in the bark. tissfle developed inward from the phello­ gen, forming a part of periderm and re­ bast - any fibres of the outer part of the sembling cortical parenchyma. plant, though mostly obtained from the secondary phloem. phellogen, see under cambium. cambium - a meristem with products of di­ visions arranged orderly in parallel files; phloem = inner bark - a principal assimi­ consists of one layer of initials and their late conducting tissue· composed of dif­ undifferentiated products, or derivatives. ferent tissues and usually located outward of the xylem and inward of the periderm. Applied only to the two lateral meristems: - vascular cambium: the meristematic Phloem can be divided into: cell layer responsible for the development - primary phloem: phloem of primary of the wood and/or the phloem. origin. - phellogen: the meristematic cell layer - secondary phloem: all bark tissues responsible for the development of the derived from the vascular cambium during periderm. secondary growth. There are also other meristems which form Secondary phloem can be subdivided into additional tissues: two layers that are sometimes visible to - dilatation meristem: a radially orient­ the unaided eye: ed meristematic cell layer in the phloem ray - non-collapsed secondary phloem: of some plants; responsible for a distinctly part of the secondary phloem with open funnel-shaped phloem ray dilatation. and non-collapsed sieve-elements. - collapsed secondary phloem: part cork - a trade product which is mainly ex­ of secondary phloem where sieve ele­ tracted from the cork oak tree (Quercus ments, companion or Strasburger cells, suber). and sometimes axial phloem parenchyma cells are collapsed cortex - the tissue ofprimary origin that be­ longs neither to the epidermis, the peri­ Other structures in the secondary phloem derm, or the phloem; between epidermis that can sometimes be distinguished with (periderm) and the vascular system. the unaided eye: - sclerenchymatic elements: fibres, dilatation tissue - all tissue affected by di­ sclereids, and their intermediates forming latation growth (Fig. 2). various structures in the secondary phlo­ em. Patterns thus formed may be seen in periderm - the secondarily developed pro­ cross and tangential section. tective bark tissue replacing the epidermis, - phloem rays: rays traversing the phlo­ or built during rhytidome formation; con­ em. sists of phellem, phellogen, and phello­ derm. phloem fibre - bark fibre of the phloem.

Downloaded from Brill.com09/24/2021 07:42:43AM via free access Junikka - Macroscopic bark tenninology 39 rhytidome = outer bark - the dead outer periderm, cut off by this periderm from part of bark including the last-formed still living secondary phloem.

II. Bark texture (consistency) texture - composition of the bark largely -loose: outer and/or inner bark breaks determined by the character of the cells up on cutting into coarse or fine grains composing the tissue, but also affected by (vs. firm). the progress of decay of the rhytidome. - granular: usually referring to inner bark which is mainly composed of scle­ The texture may be described as follows: reids. - corky: outer bark with the texture and - mealy: outer bark falls off like powder. quality of cork. - homogeneous: either fibres or sclere- - fibrous: outer and/or inner bark main­ ids occur (vs. heterogeneous). ly composed offibres. - soft: outer and/or inner bark is soft - brittle: outer and/or inner bark is hard, and easy to cut (vs. hard). breakable. Outer bark may emit a metallic - laminate: concentric, cylindrical or in­ sound on cutting, inner bark may be fi­ terrupted layers in the phloem formed by brous but still brittle. sclerenchymatic elements.

III. Bark patterns lILa: Bark patterns in cross and tangential streaks - striations on the sUrface of the section blaze usually formed by phloem rays and sclerenchymatic tissues. corrugations - the inner surface of the bark Streaks may be: is corrugated matching the similar pattern - longitudinal: longitudinal strzatlOns on sapwood. against different-coloured background. - reticulate: regular or wavy, net-like dilatation (growth) - the process ensuring striations against different-coloured back­ the tangential widening of the bark during ground. growth (Fig. 2). flame-marks - a pattern resembling flames IILb: Fissuring formed by phloem rays seen in the cross section ofphloem. bark, fissured - bark cracked lengthwise into fissures separated by ridges. phloem, mottled - phloem which is marked with spots of various colours or shades in fissures: more or less longitudinal grooves tangential section. between ridges in the rhytidome. phloem, scalariform - a pattern formed by They may be classified according to depth phloem rays in the cross section of phlo­ and length as follows: em resembling ladder-like structure with - deep: at least as deep as half of the total radial 'rungs'. thickness of the bark (Fig. 4). - shallow: less than half as deep as the ripple marks - fine, parallel, horizontal total bark thickness (Fig. 5, see also Figs. lines in the tangential section of wood or 6,8, and 9). bark, caused by the storied structure of all - boat-shaped: oval or elliptical fissures the wood and bark elements, or by the which are not continuous (Fig. 10). distribution in horizontal layers of the - short: less than 15 cm long. wood rays and bark rays only (Fig. 3). -long, elongated: more than 15 em long.

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In addition, fissures can be classified into Flakes may be: different morphological types: - rectangular. - V-shaped: more or less acute fissures - irregular. sometimes penetrating inner bark, lined by - circular. special associated periderm( s) (See Fig. 4). - papery: thin and very flexible flakes - round: the outer surface is concave in (vs. slab-like flakes). cross section. - scrolled: long, thin flakes which are - square-shaped: flat bottom fissures, rolled up by their edges (Fig. 16). always restricted to the dead outer bark - shaggy: loosened, usually slightly curv­ (Fig. 6). ed rectangular or irregular flakes which - irregular: different-sized gaps or fur­ may hang for a time on the stem (Cf. Fig. rows in the bark surface (Fig. 5). 12). - compound: anastomosing shallow fis­ sures which are formed in the bottoms of bark, heterogeneous - more than one type existing fissures (Fig. 7). of bark is encountered in the same stem - wavy: longitudinal grooves that are (Fig. 13). coarse with wavy, more or less irregular faces difficult to define clearly (Fig. 11). bark, patchy -lighter blotches on the outer Fissures may be: surface of the rhytidome resulting from - parallel: grooves are parallel, long, usu­ the irregular dehiscence of old rhytidome ally regular. The resulting, long ridges may plates, usually two colours dominate on crack transversally (Fig. 12). the bark (Fig. 18). - oblique: grooves are short or long, anastomosing but not so regular and dis­ bark, powdery - bark covered with a fine tinct as in reticulate fissured barks. The powder-like crust which usually rubs off resulting ridges may crack transversally. easily. It is mostly associated with smooth - reticulate: grooves join each other and barks. divide again irregularly leaving non-con­ tinuous and sometimes obscure ridges. bark, scaly; scales - bark which has small more or less irregular patches or scales of ridges: more or less continuous raised parts rhytidome, less than 7.5 cm long, which of the rhytidome between fissures. become detached. Ridges may be: Scales: - flattened: the outer surface is plane in According to persistence one may roughly cross section (Fig. 8). divide them as follows: - hollow: the outer surface is concave in - adherent: retained some time on the cross section (Fig. 9). stem. - rounded: the outer sUrface is convex in -loose-hanged: shed seasonally (Fig. cross section (Fig. 4). 19, see also Fig. 13). - V-shaped: the outer surface is sharp in Scales may be roughly divided according cross section. to density as follows: - reticulate: ridges join each other and - close. irregularly divide again enclosing non­ - distant. continuous fissures (Fig. 14). Scales may be roughly divided according IIl.c: Scaling to shape as follows: - rectangular. bark, flaky; flakes - bark which has large - irregular. and variable patches or flakes of rhyti­ - circular. dome, more than 7.5 cm long, which be­ - papery: thin and very flexible. come detached. - flat-sided: one or several layers thick.

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- chunky: usually with irregular rough - round: (usually small). faces and irregularly chunky shaped. - stellate: star-shaped. - scollop-shaped: thickest in the mid­ - diamond-shaped: rhomboid. dle, tapering to the edges, and leaving a The size of lenticels may be divided ac­ scollop-shaped depression on the bole on cording to the greatest diameter: sloughing (Cf. Fig. 22). - large: more than 5 mm in diameter. - scrolled: thin scales which are rolled - medium: 3-5 mm in diameter. up by their edges. - small: less than 3 mm in diameter. - shaggy: loose and usually slightly curv­ ed, rectangular or irregular scales which Lenticels may occur on bark: may hang for a time on the stem (Fig. 19). - numerous (vs. scarce) (only when present in large numbers or only few). bark, stringy - thick, loose, long-fibred - solitary (vs. compound). bark, never deciduous. - in vertical lines (Fig. 21). - in horizontal lines. bark, surface rotten - bark is shortly fis­ - in oblique lines. sured (with fissures varying in depth and According to their consistency they may be: cross section), scaly, rugose or smooth; - soft. sloughing is very variable scales being - powdery. small, adherent, chunky or flat-sided; in - compacted. a transverse section the inner edge of the outer bark is following sUrface configura­ bark, pock-marked - bark covered with tions, not parallel to cambium. small shallow circular depressions, less than 1 em in diameter. bark, tessellated - surface marked by more or less regular, square or oblong plates or bark, rough - bark which has an uneven blocks remaining for a long time on the sUrface; the term includes various scaly, stem (Fig. 15). flaky and fissured barks. ring bark - a type of rhytidome in which bark, rugose - bark which is covered by periderms may be formed parallel to the wrinkles, depressions, small irregularly first one. The concentric cylinders of bark desquamating scales, shallow fissures; thus formed result in a rhytidome which bark appears smooth from a distance (Fig. detaches annually with large sheets. 20).

III.d: External markings bark, scribbly - smooth bark carries char­ acteristic 'scribbles' caused by insect lar­ bark, dippled - bark covered with shallow, vae. usually circular depressions, more than 1 em in diameter. These are scars of the bark, smooth - bark is even, thin, unbro­ scaled-off old bark (Fig. 22). ken, though it may be bumpy or pimply from the lenticels (Fig. 21). bark, lenticellate; lenticels - barks which have on the surface more or less raised, burls - hard woody excrescences on a tree, often somewhat corky spots or lenticels more or less rounded in form, usually (Fig. 21). resulting from the entwined growth of a Lenticels: cluster of adventitious buds. The form of lenticels may be according to the greatest diameter: cankers - definite, relatively localised, ne­ -linear: usually horizontally elongated crotic lesions primarily of the bark and like buttonholes (rarely vertical). cambium.

Downloaded from Brill.com09/24/2021 07:42:43AM via free access 42 IAWA Journal, Vol. 15 (l), 1994 ------eye marks - eye-shaped marks on trunks scroll marks - raised sinuous marks be­ and branches of smooth-barked trees tween depressions which are formed when (Fig. 23). scollop-shaped scales are detached (Fig. 22). hoops - raised transverse rings which parti­ ally or completely encircle the stem (Cf. spines - sharp hardened outgrowths of the Fig. 24). bark and wood, usually modified branches or leaves; tearing the wood, if detached. mussel-shell markings - parallel, sinu­ ous, concentric ridges on the bark left Spines may be: after shedding of the rhytidome (Fig. 17). - simple (vs. branched). - straight (vs. curved). prickles -- sharp outgrowths from the bark, - hard (vs. supple). detachable without tearing the woolf (Fig. - tapering. 25). - conical. ring-grooves - transverse grooves which warts - hard or firm excrescences other than partially or completely encircle the stem lenticels, spines, prickles, or burls which (Fig. 24). are formed on the stem.

IV. Exudation blaze dry - the surface of the blaze is not - abundant: exudation is profuse for a exuding, feels dry when touched. while (vs. scanty).

gum - viscid secretions, exuding naturally or blaze, wet - the surface of the blaze slightly on incision or infection, generally col­ exuding, feels moistened when touched. ourless, nontoxic, odourless, and usually tasteless; and on desiccation, or exposure exudate - moisture or liquid of the living to air, they become hard, clear, glassy tissue, which seep or flows after slashing, masses. usually from the inner bark. kino - newly formed kino is usually of a Purely physically an exudate may be: thick consistency similar to that of honey, ranging from straw to blood-red in colour; - clear: transparent (vs. opaque). . .. and dries to a brittle, semi-transparent - coloured: may be white, yellow, gold- solid. en, brownish, red, blackish. - discoloured: the colour changes with­ latex - latex is a colloidal suspension or in a few minutes. emulsion of water-insoluble substances, - frothy: forms foam when rubbed be­ suspended in an aqueous phase. It is typi­ tween fingers. cally white (milky), but may be yellow to -liquid: flows readily, often transparent. red, or colourless. - viscous: flows slowly, not necessarily sticky. resin - colourless exudate or light brown - sticky: adhesive and sticks to the fin­ glossy solids, which are stable, inert gers (vs. non-sticky). and amorphous and become sticky when - odorous: smells pleasantly, for instance heated. like incense or wintergreen. - oleoresins: a solution of resin acids in - smelly: smells more or less unpleasant­ turpentine; oleoresin is a pale yellow, clear ly, for instance like garlic, pepper, sul­ and sticky mass, which upon evaporation phur, excrement. of volatile compounds becomes brittle.

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-'natural resins': they can be colour­ British Commonwealth forest terminology 2 less, as with dammars, or light brown (1957). Empire Forestry Assoc., London. glossy solids; they are stable, inert and Brooker, M.I.H. & D.A K1einig. 1990a. amorphous, become sticky when heated Field guide to Eucalypts. Vol. 1: South­ and are fusible (above JOO °C) with no eastern . Revised edition. Inkata sharp melting points. Press, Melbourne & Sydney. Brooker, M.I.H. & D.A Kleinig. 1990b. Field guide to Eucalypts. Vol. 2: South­ V. Bark cutting western and Southern Australia. Inkata Press, Melbourne & Sydney. blaze = slash - a shallow cut removing a Brown, C.L. 1971. Secondary growth. In: portion of the bark and wood so as to aid Trees: structure and function (eds. M.H. in species identification (Figs. 2 & 3). Zimmermann & C.L. Brown): 67-123. Springer Verlag, Berlin. slash = blaze. Biisgen, M. & E. Miinch. 1929. The structure and life of forest trees. 3rd ed. (transl. by T. Thompson). Chapman & Hall, London. Acknowledgements Chan, L.-L. 1985. The anatomy of the bark The author is greatly indebted to Jifke of Libocedrus in New Zealand. IAWA Koek-Noorman for the worthwhile discus­ Bull. n. s. 6: 23-34. sions and her valuable advice concerning Chang, Y.-P. 1954. Bark structure of North the structure of this article. I also thank Tim American Conifers. Techn. Bull. U. S. Whitmore, Pieter Baas, and an anonymous D.A 1095: 1-86. referee for their careful reading of the manu­ Chattaway, M.M. 1953. The anatomy of script, and Lubbert Westra for correction of bark. I. The genus Eucalyptus. Austral. J. the English text. Bot. 1: 402-433. Thanks are due to Marcel Polak in Utrecht Corner, E.I.H. 1988. Wayside trees of Ma­ for permission to use his slides (Figs. 2, laya. 3rded. Vol. 1. United Selangor Press, 3, lO, 12, 21). The author is also grateful to Kuala Lumpur. Henrik Rypkema for a schematic illustration Craddock, O. 1932. The rind of the Po do­ of Fig. 1 and to Asko Nerg for the rest of the carps, with special reference to the bark. illustrations (Figs. 4-9, 14-17,25). New Zealand I. Forest. 3: 61-65. Societas Biologica Fennica Vanamo and Dirr, M.A 1977. Manual of woody land­ the Finnish Cultural Foundation provided scape plants: their identification, ornamen­ financial support. tal characteristics, culture, propagation and uses. Stipes Publishing, Champaign. Eames, AI. & L.H. MacDaniels. 1947. An References introduction to . 2nd ed. Mc­ American heritage dictionary of the English Graw-Hill, New York. language (1980). Houghton Mifflin, Bos­ Edlin, H.L. 1976. Trees and man. Columbia ton. University Press, New York. Beard, F. S. 1944. Key for the identification Esau, K. 1960. Anatomy of seed plants. John of the more important trees of Tobago on Wiley & Sons, New York. characters of bark and blaze. Empire For­ Esau, K. 1965. Plant anatomy. 2nd ed. John est. J. 23: 34-36. Wiley & Sons, New York. Bena, P. 1960. Essences forestieres de Guy­ Esau, K. 1969. The phloem. Handbuch der ane. Bureau Agricole et Forestier Guya­ Planzenanatomie VI2. Gebr. Borntraeger, nais, Paris. Berlin. . Bor, N.L. 1953. Manual of Indian forest Esau, K. 1979. Phloem. In: Anatomy of the botany. Oxford University Press, London. Dicotyledons. 2nd ed. Vol. 1. (eds. C.R. British Commonwealth forest terminology 1 Metcalfe & L. Chalk): 181-189. Claren­ (1953). Empire Forestry Assoc., London. don Press, Oxford.

Downloaded from Brill.com09/24/2021 07:42:43AM via free access 44 IAWA Journal, Vol. 15 (1),1994

Fahn, A. 1990. Plant anatomy. 4th ed. Per­ Jimenez-Saa, J. H. 1973. Forestry develop­ gamon Press, Oxford. ment in Surinam. Forest botany. Project Ford-Robertson, F.C. (ed.). 1971. Terminol­ working document 4. FO: SF/SURl7l1 ogy of forest science, technology practice 506. FAO, Paramaribo. and products. English language version. Kriissmann, G. 1984. Manual of cultivated The multilingual forestry terminology broad-leaved trees & . Vol. I, A-D series 1. Society of American Foresters, (transl. Michael E. Epp). B T. Batsford, Washington D.C. London. Foxworthy, F.W. 1927. Commercial timber Kunkel, G. 1965. The trees of Liberia. BLV trees of the Malay Penisula. Malayan For­ Bayerischer Landwirtschaftsverlag, Mlin­ est Records 3. Federated Malay States chen. Governement, . Lely, H.V. 1925. The useful trees of North­ Francis, W.D. 1951. Australian rain-forest ern Nigeria. Crown Agents, London. trees. Forestry and Timber Bureau, Com­ Letouzey, R. 1986. Manual of forest botany. monwealth of Australia. Tropical Africa. 1. General botany (transl. Ghouse, A.K.M. & A. Jamal. 1978. Studies R Hugget). Centre Technique Forestier on the bark anatomy of some Cassia spe­ Tropical, Nogent-sur-Marne. cies. Indian J. Bot. 1 (1 & 2): 91-97. Macdonald encyclopedia of trees. 1982. Mac­ G6mez-Vazquez, B.G. & E.M. Engleman. donald & Co. Ltd., London & Sydney. 1984. Bark anatomy of Bursera longipes Martin, RE. & J.B. Crist. 1970. Elements (Rose) Standley and Bursera copallifera of bark structure and terminology. Wood (Sesse & Moc.) Bullock. lAWA Bull. Fiber 2: 269-279. n.s. 5: 335-340. Meijer, W. 1974. Field guide for trees of Hall, N., RD. Johnston & G.M. Chippen­ West Malesia. University of Kentucky dale. 1970. Forest trees of Australia. De­ Book Store, Lexington, USA. partment of National Development For­ Meijer, W. & G.H.S. Wood. 1964. Diptero­ estry and Timber Bureau & Australian carps of Sabah (North Borneo). Sabah Governement Publ. Service, Canberra. Forest Records 5, Forest Department, San­ Hightshoe, G.L 1989. Native trees, shrubs, dakan. and for urban and rural America. Metcalfe, C.R. 1979. The stem. In: Anatomy Van Nostrand Reinhold, New York. of the Dicotyledons. 2nd ed. Vol. 1 (eds. Hillis, W.E. 1987. Heartwood and tree exu­ C. R. Metcalfe & L. Chalk): 166-180. dates. Springer Verlag, Berlin. Clarendon Press, Oxford. Howard, E. T. 1971. Bark structure of the Nultsch, W. 1971. General botany. Academic Southern . Wood Sci. 3: 134-148. Press, London. IAWA, Committee on Nomenclature. 1957. Outer, R.W. den. 1967. Histological investi­ International glossary of terms used in gations of the secondary phloem of Gym­ wood anatomy. Trop. Woods 107: 1-36. nosperms. Meded. Landbouwhogeschool. IAWA, Committee on Nomenclature. 1964. 67-7: 1-119. Multilingual glossary of terms used in Outer, R.W. den. 1972. Tentative determina­ wood anatomy. Konkordia, Winterthur. tion key to 600 trees, shrubs and climbers IAWA Committee. 1989. IAWA list of micro­ from the Ivory Coast, Africa, mainly based scopic features for hardwood identifica­ on characters of the living bark, besides tion. IAWA Bull. n.s. 10: 219-332. the rhytidome and the leaf. 1. Large trees. Irvine, F. R. 1952. West African botany. Meded. Landbouwhogeschool. 72-18: 1- Oxford University Press, London. 73. Jackson, B. D. 1928. A glossary of botanic Panshin, A.J. & c. de Zeeuw. 1980. Text­ terms, 4th ed. Gerald Duckworth, London. book of wood technology. McGraw Hill, Jenik, J. & B.l. Harris. 1969. Root-spines New York. and spine-roots in dicotyledonous trees of Penford, A.R. & J.L. Willis. 1961. The tropical Africa. Oesterr. Bot. Z. 117: 128- Eucalypts: botany, cultivation, chemistry, 138. and utilization. Leonard Hill, London.

Downloaded from Brill.com09/24/2021 07:42:43AM via free access Junikka - Macroscopic bark terminology 45

Polak, AM. 1992. Major timber trees of Thrower, S.L. 1988. Hong Kong trees. Om­ Guyana. A field guide. Tropenbos Series nibus volume. The Urban Council, Hong 2. Tropenbos Foundation, Wageningen. Kong. Radford, AE., W.C. Dickison, J.R. Massey Timberlake, J. 1980. Handbook of & e.R. Bell. 1974. system­ acacias. Division of Land Utilisation, Min­ atics. Harper & Row, New York. istry of Agriculture, Gaborone, Botswana. Raven, P.H., R.F. Evert & H. Curtis. 1976. Trockenbrodt, M. 1990. Survey and discus­ Biology of plants. 2nd ed. Worth Pub­ sion of the terminology used in bark anat­ lishers, New York. omy. lAW A Bull. n.s. 11: 141-166. Rollet, B. 1980, 1982. Interet de l'etude des Troupin, G. 1978. Flore du Rwanda. Sper­ ecorces dans la determination des arbres matophytes. Tome 1. Musee Royal de tropicaux sur pied. Bois Forets Trop. 194 l' Afrique Centrale, Tervuren, Belgique; (novembre-decembre): 3-28; 195 (ler Vaucher, H. 1990. Baumrinden. Ferdinand trimestre): 31-50. Enke, Stuttgart. Rosayro, R.A de. 1953. Field characters in Voorhoeve, AG. 1965. Liberian high forest the identification of tropical forest trees. trees. Centre for Agricultural Publications Empire Forest. Rev. 32: 124-141; Corri­ and Documentation, Wageningen. genda and Addenda in 32: 205. Whitmore, T.e. 1962a. Studies in systematic Rosayro, R.A de. 1960. Field key to the bark morphology. 1. Bark morphology in identification of wet evergreen (rain forest) Dipterocarpaceae. New Phytol. 61: 191- tree species. Ceylon Forest. N. Ser. 4 (3): 207. 287-294. Whitmore, T.e. 1962b. Studies in system­ Roth, 1. 1981. Structural patterns of tropical atic bark morphology. 2. General features barks. Handbuch der Pflanzenanatomie. of bark construction in Dipterocarpaceae. Band IX, Teil 3. Gebr. Borntraeger, Ber­ New Phytol. 61: 208-220. lin. Whitmore, T.e. 1962c. Studies in systematic Saint Aubin, G. de. 1963. La foret du Gabon. bark morphology. 3. Bark in Centre Technique Forestier Tropical, No­ Dipterocarpaceae. Gard. Bull. Singapore gent-sur-Marne. 19 (2): 321-371. Society of American Foresters, Committee Whitmore, T.C. 1963. Studies in systematic on forestry terminology. 1950. Forestry bark morphology. 4. The bark of beech, terminology. A glossary of technical terms oak and sweet . New Phytol. 62: used in forestry. Revised edition. Society 161-169. of American Foresters, Washington. Whitmore, T.e. (ed.). 1972. Tree flora of Srivastava, L.M. 1964. Anatomy, chemistry, Malaya. A manual for foresters. Vol. 1. and physiology of bark. Int. Rev. Forest. Longman, London. Res. 1: 203-277. Wood, G.H. S. 1952. Bark as a means of Storrs, A E. G. 1979. Know your trees. tree identification. J. Oxford Univ. Forest Some of the common trees found in Zam­ Soc. 6: 15-27. bia. The Forest Department, Ndola, Zam­ Wyatt-Smith, J. 1954. Suggested definitions bia. of field characters (for use in the identifi­ Sutton, R.F. & R.W. Tinus. 1983. Root and cation of tropical forest trees in Malaya). root system terminology. Forest Sci. Mon­ Malayan Forester 17: 170-183. ograph. 24. Society of American Forest­ Wyk, AE. van. 1985. The genus Eugenia ers, Washington. (Myrtaceae) in southern Africa: structure Symington, e.F. 1943. Foresters' manual of and taxonomic value of bark. S. African Dipterocarps. Malayan Forest Records 16. J. Bot. 51: 157-180. Syonan Hakubutukan, Kuala Lumpur. Yunus, M., D. Yunus & M. Iqbal. 1990. Sys­ Tailfer, Y. 1989. La foret dense d' Afrique tematic bark morphology of some tropi­ tropic ale. Identification pratique des cal trees. Bot. J. Linn. Soc. 103: 367- principaux arbres. Tome 1. AC.C.T., 377. Paris.

Downloaded from Brill.com09/24/2021 07:42:43AM via free access