IAWA LIST OF MJCROSCOPIC FEATURES FOR HARDWOOD IDENTIFICATION

byan

IAWA Comnilttee

4 .,

E. A. Wheeler, P. Baas & P. E. Gasson (editors)

IAWA LIST OF MICROSCOPIC FEATURES FOR HARDWOOD IDENTIFICATION with an Appendix on non-anatomical information

IAWA Conirnittee Veronica Angyalossy Alfonso - So Paulo, Brazil Leiden, The Netherlands Pieter Baas Sherwin Carlquist - Clareniont, California, USA So Paulo, Brazil Joao Peres Chirnelo Vera T. Rauber Coradin - Brasilia, Brazil Pierre Diienrte - Nogent-sur-Marne. France Peter E. Gasson -. Kew, UK Diciger Grosser Miinchen, FRG Jugo lhe - I-lighett, Victoria, Australia Keiko Kuroda Kyoto, Japan Regis B. Mifler Madison, Wisconsin, USA Ken Ogata - Tsukuba, Japan Hans Georg Richter - Hamhurg, FRG Ben L H. ter Welle - Utrecht, The Netherlands Elisabeth A. Wheelcr Raleigh, North Carolina. USA

edited by E. A. Wheeler, P. Baas and P.E.

Gasson

1989. IAWA Builetin n.s. 10 (3): 219-332 Puhljshed for the International Association ofWood Anatornisis at ihe Rijksherbariam, Leiclen, The Nerherlands

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PREFACE This list of rnicroscopic features for hardwood identification is lhe successorto lhe Standard List of Characters Suitable For Computerized Hardwood Jdentiflcation' published in 1981 (IAWA Bulietin n.s. 2: 99-145) with an explanation of lhe coding procedure by R.B. Milier. Thc 1981 publication greatly stimulated international exchange of information and experience on characters suitable for hardwood identiflcation, and inspired considerable debate on lhe most desirable coding procedures and ideruiflcation programs. Therefore, aL Lhe IAWA mceting during lhe XIV inLernational Botanic.aI Congress in Berlin, July 1987, it was decidcd to revise the 1981 standard list. Because oU Lhe continuing developments in computer technology and prograrnrning, ii was agreed to limit Lhe scope of lhe new list to dcfinitions, explanatory conirnentary, and iliustrations of wood anatomical descriptors, rather Lhan concentrate on codiiig procedures. A new Committee was appointed by lhe IAWA Council to work towards Lhe new list, and thanks to a substaruial grant from lhe USDA Competidve Research Granis - Wood Utilization Program (Grant No. 88-33541-4081), a workshop was held by lhe Committee from October 2-7, 1988, in lhe Department of Wood & Paper Science, North Carolina State University, Raleigh, NC. USA, under lhe joint auspices of IAWA and IUFRO Division 5. A preiiminary list was prepared during lhe workshop. IAWA rnembcrs were invited to cornment on this list, and these comrnents helped with lhe final preparation of Lhe new li st. Pie hst presented here was agreed to after review of subsequent drafts and extensive internal consuitation between conimotee members. Although ihis list has 163 anatomical and 58 misceilaneous features, it is not a complete li,[ encompassing ali Lhe structural patterns that one can encounter in hardwoods. lnstead it is intended to be a concise list of features usefui for identification purposes. Also, lhe numbers as signed to each feature in lhe present list are not meant to be codes for a computer program, but are intended to serve for easy reference, and to help translate data from one program/database tu another. Wood and wood celis are biological elements, formed in trees, shrubs, and climbcrs to fui! dl a physiological or niechanicai function. Aithough there is more discrete diversity in wood structure than in many other plant parts, there is also much continuous variation, and any auempt to ciassify this diversity into well-defined features has an artificial eleinent. Yet we are confident that in Lhe feature list presented here ambiguity of descriptors has heen iimited to a minimiitn. and we hope that ali present and future coileagues engaged in wood and decripti\ e wood anatomy will find this list a valuable guide and reference. The IAWA Committee: VERONICA ANGYALOSSY ALFONSO Diviso de Madeiras, I.P.T. Cidade Untversitaria, Sio Paulo. Brazil PIETER BAAS Rijksherbariuni, Leiden, 'lhe Netherlands
SHERWIN CARLQUIST

Rancho Santa Ana Bo:anic Garden, Clarculont, Ca!ifornia, U.S. A.

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JOAO PERES CHIMELO Diviso de Madeiras, I.P.T. Cidade Universitria, So Paulo, Brazil VERA T. RAUBER CORADIN Instituto Brasiliero de Desenvolvimento Florestal, Departmento de Pesquisa, Brasilia, Brazil PIERRE DTIENNE Division d'Anaomie des Bois, Centre Technique Forestier Tropical, Nogent-sur-Mame, France PETER E. GASSON Jodreil Laboratory, Royai Botanic Gardens, Kew, U. K. DIETGER GROSSER Institui fiir Holzforschung und Holztechnik der Universitiit Mnchen, Miinchen, F.R.G. JUGO ILIC CSIRO, Wood Scicnce & Technology, Highett, Victoria, Australia KEIKO KURODA Forestry & Forest Products Research Institute, Kansai Branch, Kyoto, Japan
REGIS B. MILLER

ACKNOWLEDGEMENTS

The IAWA Cornrnittee is greatly indebted to the following institutions and individuais: The USDA Competitive Rescarch Grants-Wood Utilization Program (Grant No. 88-335414081) for financing the IAWA/1UFRO Workshop in Raleigh, North Carolina, and subsequent rneetings in London and Leiden by P. Baas, P.E. Gasson, and E. A. Wheeler. The Department of Wood and Paper Science, N. C. State University for offering hospitaiity and facilities during the IAWA/IUFRO Workshop in Raleigh; especially Dr. C.A. LaPasha and Ms. Vann Moore for help with preparation of the various drafts, and Ms. Milie Sullivan. The Forest Products Laboratory, Madison, Wisonsin, USA, for providing financial support towards the printing costs of this special issue, The Jodreli Laboratory, Royai Botanical Gardens Kew, UK, for supporting photographic work, and providing facilities and hospitality during a mecting in March 1989 for the selection of iliustrations. The BaiIey-Wctmore Laboratory of Plant Anatomy and Morphology, harvard University, and Dr. P. B. Tomhinson, Dr. D. Pfister, and Dr. A. Knoll for giving access to the Bailey negatives and darkroom facihities. The Rijksherbarium for various facilities; especially to Ms. Emma E. van Nieuwkoop for mounting the plates, and lay-out editing. All IAWA Members who have kindly given their comrnents on various drafts of this list: K.M. Bhat, India Lim Seng Choon, Kepong, Malaysia D. F. Cutler, Kew, UK W, C. Dickison, Chapel Hill, NC, USA T. Fujii, Tsukuba, Japan H. Gottwald, Hamburg, FRG Mary Gregory, Kew, UK Yvonne Hemberger, Harnburg, FRG Alberta M.W. Mennega, Utrecht, The Netherlands C.A. LaPasha, Raleigh, NC, USA A. Londono, Columbia Paula Rudail, Kew, UK M. Seth, india

Center for Wood Anatomy Research, Forest Products Laboratory, Madison,Wisconsjn, U.S.A. KEN OGATA Wood Technology Division, Forestry & Forest Products Research Institute, Tsukuha, Japan HANS GE0RG RICHTER Institui fr Holzbiologie und Holzschutz, Bundesforschungsanstalt fr Forst- und Holzwirtschaft, Hamburg, F.R.G. BEN J. H. TER WELLE Rijksuniversiteit Utrecht, Instiruut voor Systeniatische Plantkunde, Utrecht, Thc Netheriands ELISABETH A. WHEELER Department of Wood & Paper Science, North Carolina State University, Raleigh, North Carouna, U.S.A.

AcknowuedgemefltS for iflustrations Photograplis by courtesy of: I. W. Bailey, Bailey -Wetmore Laboratory of Plant Anatomy and Morphoiogy, Harvard University: 10, 11, 16, 18, 39, 57, 58, 64, 65, 148. Blumea: 38, 44, 73, 74 (Baas 1973), 174 (Van Vliet 1981). P. Dtienne: 129. P.E. Gasson: 2,4,7, 8, 12, 19, 21, 26, 28, 30-34, 36, 37, 40, 45-54, 63, 66, 75, 78-82, 84-86, 88, 90-93, 95-99, 102-106, III, 114-116, 118, 120, 122, 126-128, 130135, 137-144, 151, 153, 154, 156, 157, 159, 161, 163-168, 171, 172, 176, 178, 180182, 188. D. Grosser: 15, 27, 29, 55, 68, 71, 72, 112, 113, 146, 158, 170, 173, 177.

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1 AWA Buileun: 3 (Bricigwater & Baas 1982), 35 (Vidal Comes ex ai. 1988), 70 & 123 (Bridg water & Baas 1982), 155 (Topper & Koek-Noorman 1980), 175 Baas et ai. 1988), 184 (Gottwald 1983), 185 (Ter WelIe 1980). J. lhe: 56. C.A.LaPasha: 190. R. B. Milier: 160, 186, 187, 189. K. Ogata: 1, 5, 9, 13, 14, 20, 22, 24, 25, 41, 42, 61, 62, 76, 77, 83, 89, 94, 101, 107-109, 117, 119, 124, 125, 136, 145, 147, 149, 150, 152, 162, 179. E. A. Wheeler: 6, 17, 23, 43, 59, 60, 67, 87, 100, 110, 121, 169, 183. H. P. Wilkinson: 69.

EXI'LANATORY NOTES

Quantitative FeatureS - For quantitative features of general applicability (e.g., vessel frcquency, tangential vessel lumen diameter, vessel element hength, and fibre length), this list ineludes broad categories for easy use when identifying unknowns, as well as more precise quantitative descriptors (mean, range, standard deviation). When constnicting a database the nutnbers of samples as well as Lhe number of measurements or counis done per samplc should be recorded Different computer programs allow storage of different amounts of information (eg., ali measurements, orjust the means, ranges, and standard deviations), and use different algorithms for rnatching quantitative features. This publication does not recommend a particular program or a particular method for the storage and retrieval of quantitative data, but provides some guidance on how to obtain these data.

Variable Features and Relative Abundance - l3ecausc of wood's inherent variabihity, it is inevitable that some fe.atures will be well-defined in some samples while absent or iiidefined in other samples of the sarne species. Accommodatiflg such variability has always beco a problem in key constiUctiofl, and most keys (computerised or otherwise) have provisions for such situations. Describing relative abundance of some features, e.g., prismatic crystals, is also problematic, and textual cominents on reiative ftequency should be added to a description or database- In this list of descriptors, some features apply only when the characteristie is of commoo occurrence. For such features, the illustrations and examples are intended to help interpret 'common'. Although many keys have used these sarne features accompanied by the sarne qualifier 'common', there have been no extensive systernatic analyses to determine what per cent ocurrence constitutes 'common'. Therefore no quantitative criteria for 'common' have been
offered in this list.

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IAWA Bulietin n,s., Vol. lO (3), 1989 UST OF FEATURES

IAWA.Listof microscopie features for hardwood idcntification

Vessei-ray pirring - p. 253

227

Name
ANATOMICAL FEATI:RES

Growth rings - p.234 1. Growrh ring boundaries distinct 2. Growth ring boundaries indisiinct or absent Vesseis - p. 236 Porosity p. 236 3 Wood ring-poroiis 4. Wood semi-ring-porous 5. Wood diffuse-porous
Vessel arrangeinent - p. 238

30. Vessel-ray pits with distinci borders; similar to intervessel pits in size and shape throughout Lhe ray celi 31. Vessel-ray pits with much reduced borders to apparently siniple: pits rounded or angular 32. Vessel-ray pits with much reduced borders to apparently simple: pits horizontal (scalariforrn, gash-like) to vertical (palisade) 33. Vessel-ray pits of two distirict sizes or types in Lhe sarne ray cdl 34. Vesscl-ray pus unilaterally con'ipound and coarse (over 10 um) 35. Vessel-ray pits restricted to marginal rows Helica! thickenings - p. 256 36. Ilelica] t1'iickenings in vessel elernents present 37. Helical thickenings throughout body of vesse] elernent 38. Helical thickenings only in vessel element tails 39. Helical thickenings only in narrower vessel elernei:ts Tangential diameter of vessel lwnina p. 25 Mean tangential diameter of vessel luniina 40. 2^50um 41. 50-100nn 42. 100-200 .riii 43. ^200p.m 44. Mean, +1- Standard Deviation, Range, r - x 45. Vesseis of mo distinct diameter classes, wod no: nng-pero:s

6. Vesseis in tangential hands 7. Vessels in diagonal and/or radial partem 8. Vesseis in dendritic pattern Vessel groupings - p. 242 9. Vesseis exclusively solitary (90% or more) 10. Vessels iri radial multiples of4or more common 11. Vessel clusters common Solitary vessel outiine - p. 244 12. Solitary vessel outline angular Perforation plates p. 246 13. Simple perforatiori plates 14, Scalariform perforation plates 15. Scalariform perfor'ation plates with ^ 10 bars 16. Scalariforrn perforation plateswith 10-20bars 17. Scalariform perforarion plates with 20-40bars 18. Scalarit'onn perforation piates with ^ 40 bars 19. Reticulate, foraminate, and/or other types of multiple peiforatiori plaes lnteri.-essel pus: arrangernent and sire - p. 250 20. Intervessel pits scalariforrn 21. Inrervessel pus opposite 22. Intervessel pits alternate 23. Shape of alternate pits polygonal 24, Minute - 4-7 rn 25. SmaU 26. Medium - 7-10prn - ^:lOi.tm 27. Lrge 28. Range of iniervessel pit size (m) Vestured pits '- p. 252 29. Vestured pits

Vesseis per square rnilrnetre - p. 259

46, 5 5 vesseis per square millinieue 47. 5-20 vessels per square rnillimetrc 48. 20-40 vessels per square millirnetre 49, 40-100 vesseis per square rnillimetre 50. ^ 100 vcssels per square rnillimetre 51, Mean, +1- Standard Deviation, Range, n

Mean versei elernent iengrh - p. 259 52. ^350p.m 53. 350-800 piri 54. ^800sm 55, Mcan, +/- Standard Deviauon, Range. n = x

Tyloses and deposit.s ia vesseis -p. 259

56. Tyloses common 57. Tyloses sclerotic 58. Gums and other deposits in heartwoxl vessels
Wood vesselless - p.

262

59. Wood vesselless Tracheicis and fibres - p262 60. Vascular/vasicenflic tracheids present

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l(.) (i, 989

\\V.\ [_isL of nicroscpic caui Rays -p.282

tor Iidood

Ground tissue fibres - p. 264 61. Fibres with simple to minutely bordered pits 62. Fibres with clistinctly bordered pits 63. Fibre pits common in both radial and tangential walis 64. Helical thickenings m ground tissue fibres Septate fibres and parenchyma-like fibre band - p. 266 65. Septate fibres present 66. Non-septate fibres present 67. Parenchyma-like fibre bands alternating with ordinary flbres Fibre wall rhickness - p. 268 68. Fibres very thin-walled 69. Fibres thin- to thick-walled 70. Fibres vel'y thick-wallcd Mean fibre lengths - p. 269 71. 5900i.tm 72. 900-1600 jim 73. >1600j.tm 74. Mean, +/- Standard Deviation, Range, n = x Axial parenchyma - p. 270 75. Axial parenchyma absent or extremely rare Apotracheal axial parenchynia - p. 270 76. Axial parenchyrna diffuse 77. Axial parenchyma diffuse-in-aggregates
Pararracheal axial parenchyma - p 272

Ray width - p. 282 96. Rays exciusively uniseriate 97. Ray width 1 to 3 cells 98. Largerrays commonly 4- to lO-seriate 99. Larger rays comrnonly> lO-seriate 100. Rays with multiseriate portion(s) as wide as uniseriate portions Aggregare rays - p. 284 101. Aggregate rays Ray height p. 284 102. Ray height> 1 mm Rays of iwo distinci sizes - p. 286 103. Rays of mo distinct sizes Rays: celiular co,nposirion - p. 288 104. Ali ray celis procumbent 105. Ali ray celis upright and/or square 106. Body ray celis procumbent with one row of upright and/or squarc marginal celis 107. Body ray cells procumbent with mostly 2-4 rows of upright and/or square marginal cells 108. Body ray edis procunibent with over 4 rows of upright and/or squarc marginal celis 109. Rays with procumbent, squarc and upright cclls mixed throughout the ray Shearh celis - p. 292 110. Sheath celis Tile celLs - p. 292 111. Tile cells Pe,forared ray cells - p. 294 112. Perforated ray edis Disjunctive ray parerzchvma ccli wall. - p. 294 113. Disjunctive ray parenchyrna ceil walls Rays per ,nillimetre - p. 296 114, 54/mm 115. 4-12/mm 116. ^ 121mm Wood rayless - p. 297 117. Woodrayless Storied structure - p. 298 118. 119. 120. 121. 122. 123. All rays storied Low rays storied, high rays non-storied. Axial parenchyma and/or vessel elements sto: Fibres storied Rays and/or axial elements irregularly storie Number of ray tiers per axial mm

78. Axial parenchyma scanty paratracheai 79. Axial parenchyma vasicentric 80 Axial parenchyma aliform 81. Axial parenchyma lozenge-aliform 82. Axial parenchyma winged-aliform 83. Axial parenchyma confluent 84. Axial parenchyma unilateral paratracheal Banded parenchyma - p. 276 85. Axial parenchyma bands more than three celis wide 86. Axial parenchyma in narrow bands orlines up to three cells wide 87. Axial paxenchyma reticulate 88. Axial parenchyma scalariform 89. Axial parenchyma in marginal or in seemingly marginal bands Axial parenchyma ceil type/srrand length 90. Fusiform parenchyma cells 91. Two celis per parenchyma sirand 92. Four (3-4) cdlis per parenchyma strand 93. Eight (5-8) cells per parenchyma strand 94. Over eight cdlls per parenchyma sirand 95. Unlignified parenchyma p. 280

1\V,\ Bulk'tin r., 'Vol. li) Secretory elements and cambial

OU and mucilage celis - p. 300
variants

). 1)')

1 .\\V1\ List

ol

nhicroscopic icatures fnr i irdocd dcnti:cr

-p300

124. Oil and/or mucilage cdlis associated with ray parenchyma 125. 011 andj or rnucilage cells associated with axial parenchyrna 126. Oil and/or mucilage cells presem among fibres Inrerceiluiar canais - p. 302 127. Axial canais in long tangential limes 128, Axial canais in short tangential lines 129. Axial canais cliffuse 130. Radial canais 131. Intercellular canais of traumatic origin Tubes / tubuies - p. 306 132. Laticifers or tanniniferous tubes Cambial varians - p. 308 133. Included phloem, concentric 134. Included phloem, diffuse 135. Other cambial variants Mineral inclusions - p. 310 Pris,naric crystals -p. 310 136. Prisrnatic crystals present 137. Prisrnatic crystals in upright and/or square ray crus 138. Prismatic crystals in procumbcnt ray celis 139. Prismatic crystals in radial alignment in procumbent ray cells 140. Prismatic erystais in chambered upnght and/or square ray celis 141. Prismatic crystals in non-charnbered axial parenchyma cells 142. Prismaiic crystals in chanibered axial parenchymacells 143. Prismatic crystals in fibres Druses - p. 313 144 Druses present 145. Druses in ray parenchyrna celis 146. Druses in axial parenchyrna cells 147. Druses in fibres 148. Druses in chambered cells Olher crystai types -- p. 313 149. Raphides 150. Acicular crystals 151. Styloids and/or elongate crystals 152. Crystals of other shapes (mostly small) 153. Crystal sand Orher dia gnoszic crystal features - p. 315 154. More than one crystal of about the sarne size per celi or chamber 155. Two distinct sizes of crystals perceil or chamber 156. Crystals in enlarged cells 157. Ciystals in tyloses 158. Cystoliths

P. 318 Silica 159. Silica bodies preseni 160. Silica bodies in rav colis 161. Silica bodies in axial j cnchvnm c cis 162. Silica boclies iii fibres 163. Vitreous silica
APPENDIX - Non-anatornical information Geographical distribution - p. 321

p. 2 1

164. Europe and temperate Asia (Brazier and Franklin region 74) 165. Europe, excluding Mediterranean 166. Mediterranean including Northern Africa and Middle East 167, Temperate Asia (China), Japan, IJSSR 168. Central South Asia (Brazier and Franklin region 7) 169. India, Pakistan, Sri Lanka 170. Bumaa 171. Southeast Asia and the Pacific (Brazier and Franklin region 76) 172. Thailand, Laos, Vietnam, Cambodja (Indochina) 173. Indomalcsia: Indonesia, Philippines, Malaysia, Brunei, Papua New Guinca, and Solomon Jslands 174. Pacific Islands (including New Caledonia, Samoa, Hawaii. and Fiji) 175. Ausu'alia and New Zcaland (Bra7.ier and Franklin region 77 176. Australia 177. NewZealand 178. Tropical mainland Africa and adjaceni isla,ids (Brazier and Franklin region 78) 179. Tropical Africa 180. Madagascar & Mauritius, Runion & Cornores 181. Southem Africa (south of the Tropic of Capricorn) (BraLier and 1-ranklin region 70) 182. North America, north of Mexico (Braaler and Franklin region $0) 183. Neotropics and temperate Brazil (Brazier and Franklin rcgion 8 II 184. Mexico and Central America 185. Carrubbean 186. Tropical South America 187. Southern Brazii 188. Temperate South America including Argentina. Chile, Uruguav. and S. Paraguav Brazicr and Franklin region 82) Habit - p. 321 189. Tree 190. Shrub 191. Vine/liana Wood of co,nmerciai importance - p. 322 192. Wood of commercial irnportance Specfl c gravity . p. 322 193. Basic spccific gravity iow, S 0.40 194. Basic specific gravity medium, 0.40-0,75 195. Basic spccific gravity high, ^ 0.75

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Hearwood colour - p. 323 196. Heartwood colour darker than sapwood colour 197. Heartwood basically brown or shades of brown 198. Heartwood basicaily redor shades ofred 199. Heartwood basically yeilow or shades of yeliow 200. Heartwood basically white to grey 201. Heartwood with streaks 202. Heartwood not as above Odour - p. 325 203. Distinct odour Heartivood fluorescence - p. 325 204. Heartwood fluorescent Warer & ethanol extracrs: fluorescence & colour - p. 326 205. Water extract fluorescent 206. Water cxtract basically colour!ess to brown or shades of brown 207. Water extract basically redor shades ofred 208. Water extract basically yeilow or shades of yellow 209. Water exuact not as above 210. Ethanol extract fluorescent 211. Ethanol extract basically colour!ess to brown or shades of browri 212. Ethano extract basically red or shades of red 213. Ethanol extract basically yellow or shades of yellow 214. Ethanoi extract not as above Frorh rest p.327 215. Froth test positive Chrorne Azurol-S test - p. 328 216. Chrome Azurol-S test positive Burning splinter test - p. 328 217. Splinter burns to charcoal 218. Splinter burns to a fuli ash: Colour of ash bright white 219. Spiinter burns to a fui! ash: Colour of ash yellow-brown 220. Splinter bums to a fuli ash: Colour of ash other than above 221. Splinter burns to a partial ash

NAME Family, genus, species, authority. When creating a database iL is essential to record the fuli taxonomic information on the speci. mens, i.e., record farni!y, genus, species, authority. For authorities foilow curnmonly used abbreviations (listed in Mabberley 1987). Reference to Willis's Dictionary of Fiowering P!ants and Fems(Willis 1973) and Mabberley (1987) is helpful in determining the farnilia) affinities of various genera, and preferred fami!y names. When preparing a database, a!so indicate which particular classification scheme, with respect to fami!y delimitation, is heing used (e. o., Takhtajan 1980, 1987; Cronquist 1981, 1988; Thorne 1976). li can be usefui to retrieve information on the wood anatomy of particular fami!ies or to restrict the search for the identity of an unknown wood to a certain farni!y or farni!ies. Consequently, it is advisable to record the family as a feature. For the family name it is not critical what method of coding is employed so long as it is cleariy explained in notes acconlpanying the database. Por instance, the family can he indicated as 3-1etter acron yms (Weber 1982) or as numerical codes (see pp. 127 and 144-145 in Milier 1981).

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GROWTH RTNGS

1. Growth ring boundaries distinct 2. Growth ring boundaries iridistinct or absent Deflnitions: Growth ring boundaries distinct = growth rings with an abrupt structural change at thc boundaries between them, usually including a change in fibre wall thickness and/or fibre radial diameter, figs. 1, 2. Growth ring boundaries indistinct or absent = growth rings vague and marked by more or less gradual structural changes ai their poorly defined boundarics, or not visible, fig. 3. Comments: Growth ring boundanes can be marked by one ar more of the following structural changes: a. Thick-walled and radially llattened latewood fibres ar tracheids versus thin-walled earlywood fibres ar tracheids, fig. 1, e. g., Weinmannia trichosperma (Cunoriiaeae), Laurus noblis (Lauraceae). b. Marked differcnces in vesse] diarneter between latewood and earlywood of the following ring as in semi-riiig-porous and ring-parous woods, figs. 5-8, e.g., Juglans regia (Jug]andaceae), Ulmus procera (til niaceae). c. Marginal parenchyma (terminal or initial), fig. 2, e.g.,Xylopia ntida ( Annonaceae), Brachystegia Iauren:ii (Caesalpiniaceae), Juglans regia (Juglandaceae), Liriodendron tuliptfera (Magnoliaceae). lrregularly zonate, tangential parenchyma bands witfioui associated abrupi changes in fibre diameter or wall thickness are not considered marginal and do not represeni discinct growth ring boundaries, e.g., Eschwei lera subglandulosa (Lecythidaceae), Irvingia excelsa (S imaroubaeeae). d. Vascular tracheids and very narrow vessel elements very numerous or forming the ground tissue of the larewood, and absent from the earlywood, e.g., Sanbucus nigra (Caprifolia. ceae). . Decreasing frequency of parenchynia bands towards the latewood resulting in distinct fibre zones, e. g.. Lecythis pisonis (Lecythidaceae), Doneila pruniformis (Sapotaceae). t. Distended rays, e. g., Fagus spp. (Fagaceae). See Carlquist (1980, 1988) for other types of growth ring boundaries and for commonly occurring combinations of several of the above fearures. .Although absence of growth ring boundaries is a clear enough descriptor, the differences been 'indistinct' and 'distinct' boundaries are somewhat arbitrary, and there are intermediates :g. 4). Growth nngs may appear distinct when observed macroscopically, yet have indistinci boundaries ai the light microscopic leveI; distinctness of the ring boundaries should be judged with a microscope. Indistinct growth ring boundaries are very cornmon in tropical trees (fig. 3. g., Spondias mombin - Anacardiaceae, Parkia ntida -- Min,osaceae, Coelocarvon preuasii \tvristicaceae; Xanrhophvllum philippinen.ve - Polygalaceae). Nonperiodical, sporadic occurrence of ring boundaries (due to unusual clitnatic extrenies or niaic cvcrlts) should he recorded as rings ahsent or bouridaries indist inct.

Figs. 1 & 2. Growth ring boundaries distinct (feacure 1). - 1: Weinmunnia trichosperma, boundary niarked by differences in fibre and vessel diniensions, x 80. - 2: Xylopia ntida, boundary rnarked by thick-wallcd latewood fibres and marginal parenchyma band, x 48. - Fig. 3. Growib ring boundaries indistinct or absent (feanire 2), Xanthophyllum philippinense x 22. Fig. 4. Growch ring boundaries intermediate between distinct and indisunct (features 1 and 2 variable), Jacaranda copaia, x 48.

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POROSITY 3. Wood ring-porous 4. Wood semi-ring-porous 5. Wood diffuse-porous Definizions: Wood ring-porous = wood in which the vesseis in the earlywood are distinctly larger than those in the Iatewood of the prcvious and of the sarne growth ring, and foi-rn a well defined zone or ring, and in which there is an abrupt transition to the latewood of the sarne growth ring, fig. 5, e. g., Quercus robur (Fagaceae), Frw(nus exce(sjor (Oleaceae), Phellodendron a,nurense (Rutaceae), Buineija lanuginosa (Sapotaceae), Ul,nu.s americana (Ulmaceae). Wood semi-ring-porous = 1) wood in which the vesseis in the earlywood are distinctly larger than those in the Iatewood of the previous growth ring, but in which there is a gradual change to narrower vesseis in Lhe intermediate and latewood of the sarne growth ring; or 2) wood with a distinct ring of closely spaced earlywood vesscls that are not markedly larger than the Iatewood vessels of the preceding ring or the sarne growth ring. Alternative definition: intermediate condition between ring-porous and diffuse-porous wood, figs. 6, 7, e.g., Cordia trichozona (Boraginaceae), Juglans nigra (J ugi andaceae). Lagersrrocmial f (Lythraceae), Cedrela odorara (Meliaceae), Prerocarpus indicas ( Papilionaceae), oribunda Prunus amygdalus (Rosaceae), Paulownja romentosa (Scrophujarjaceae) Wood diffuse-porous = wood in which the vesseis have more or less the sarne diarneter throughout the growth ring, figs. 9, 10, e. g., Acer spp. (Aceraceae), RJ2ododefldron wadanum (Encaceae), CercidipJiyllu,njapjcum (Cercidiphyllaceae), Swjetenia spp. (Meliaceac), Entero. lobium spp. (Mirnosaceae); the vast rnajority of tropical species and most temperate species. Comments: The three features for porosity form an intergrading continuum and many species range from diffuse-porous to semi-ring . porous, or from ring-porous to semi-ring-porous. Porosity (feaiures 3-5) is coded independently of vessel arrangenient (features 6-8).This iniplies that woods with a distinct vessel arrangernent (features 6-8), as well as those with evenly disUibuted vesseis, may be diffuse-porous. In some temperate diffuse-porous woods (e- g, Pagas spp. Fagaceae, Platanu.s spp. Plalanaceae) the Iatest formed vesse]s in the latewood may be considerahly smaller than those of the earlywood of the next ring, but vessel diarneter is more or less uniforrn throughout rnost of the growth ring (fig. 10). In a description, characteristjcs of the earlywood ring of ring-porous woods should be noted, i.e., describe how rnany vessels wide the ring is. Sudo's (1959) key used the features 'pore ring: l-seriate' and 'pore ring: multiseriate'. Such characteristics can be useful in distinguishing between species, e.g., Ulmus americana typically has an earlywood zone that is one vessel deep, Ulrnus rubra has ai earlywood zone that is more than two vesseis deep. Caurion: Slow grown ring-porous woods have narrow growth rings with very little latewood (fig. 8). Be careful not to confuse the closely spaced earlywood zones of slow-grown ringporous woods with a tangential partem, or to interprel such woods as diffuse-porous.

Fig. 5. Wood ring-porous (feature 3), Phellodendron amurense, x 28. - Figs. 6 & 7. Wood semi-ring-poroti s (feature 4). 6: Prunus sp., x 25. - 7: Paulownia tomenzosa, x 18. - Fig. 8. Wood ring-porous (feature 3), bui with narrow rin g s givino false irnprcssion of di ffuc- nr semi_nngporsj. Caralpa binon [odes. x 30.

--

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Figs. 9 & 10. Wood diffuse-porous (feature 5), - 9: Rhododendron wadanum, x 75. - 10: Cercidiphyllwn japonicum, classified as diffusc-porous despite di fference s in vcssel diarneter of Iatest forrned latewood and earlywood. x 30.

'.4

VESSEL ARRANGEMENT 6. Vesseis in tangential bands 7. Vesseis in diagonal and for radial pattern 8. Vesseis in dendritic pattern Definirions: Vesseis in tangential bands = vesseis arranged perpendicular to the rays and forrning g shori or lon o tangential bands; these bands can be straight or wavy; includes ulrniforrn and festooned, figs. 11-13, e.g., Kalopanax piclus (Araliaceae), Patagonulu americana (Boraginaceae), Enkianthus cornuus (Ericaceae), Madura pom (fera (Moraceae), Pirtosporusn tobira (Pinosporaceae), Cardwellia sublirnis (Proteaceae). Vesseis in diagonal and/or radial pattern = vesseis arranged radialiy or intermediate between tangential and radial (i. e., oblique), figs. 14, 17, 20. e. g., Lithocarpus edulis (Fagaceae), Calophyllum brasiliense, C. papuanwn, Mesuaferrea (Guttiferae), Eucalyptus diversicolar, E. obliqua (Myrtaceae), Amyris sylvatica (Rutaceae), Chloraluma gonocarpa (Sapotaceae). Synonym for diagonal: in echelon'. Vesseis in dendritic pattern = vesseis arranged in a branching patiem, forining distinct tracEs, separated hy arcas devoid of vesseis, figs. 15, 16, e. g, Rhus aromarica (Anacardiaceae), Castanea dentara (Fagaceae), Chionanihus retusus (Oleaceae), Rhamnu.s carhartica (Rhamnaceae), Bumelia lanuginosa (Sapotaceae). Synonym: ti ame-like.

Figs. 11-13. Vesseis in tangential bands (feature 6). 11: Latewood vesseis in tangential bands (note aiso feature 3, wood ring-porous), Kalopanax picrus, x 80. - 12: Vesseis and parenchyma 'festooned', Cardwellja sublimis, x 30, - 13: Ali vesseis in tangential bands, Enkianrhus cornuus, x 75. Fig. 14. Vessels in a diagonal patterri (feature 7), Calophyllum papuanuin, x 29.

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..

Fig. 19. Vesseis in a diagonal to dcndritic pattcrn (fcatures 7 and 8), Burnelia ohwifo1ia (n(Ite also featiire 5, wood difuse-porous), x 45. - Fig. 20. Vesseis in a diagonal tu radial pattcrn (feature 7), Lirhocarpus edulis, x 29.

:. k
a

. ti
'1

Procedure: Vessel disnibution patterns (tangential, diagonal /radial, dendritic) are determined from the cross section at a low magnilication, and are recorded only where there is a distinct pattern. In ring-porous woods, only the internied iate- wood and latewood are earnined. The ring ofvessels at the beginning of lhe growth ring of ring-porous woods is not considered when determining vessel distribtnion pattems.

&q4.

18

Figs. 15 & 16. Vesseis in a dendritic pattern (feature 8). - 15: Rhamnus cathartjca (note also feature 5, wood diffuse-porous), x 60. - 16: Rhus arornatica, clendritic pauern restrictcd to Iatewood vesseis (note also feature 3, wood ring-porous), x 35. --- Fig. 17. Vessels ir] a radial pattern (feature 7), Amyris sy/vazica, x 18. - Fig. 19. Narrow vesseis in a tangential io diagonal pattern (features 6 and 7), Kalopanax picrus (note also feature 3, wood ring-porous), x 80.

Commenrs: These features often occur iri cornbination, Vesse] arrangement in some woods intergrades between tangential and diagonal (fig. 18). Diagonal and dendriric oftcn iritergrade (fig. 19). AlI applilable features shou]d be recorded. The arrangement of pore clusters seen in most species of Ulmus (Ulmaceae) lias been called ulmiform; thjs describes woods where the latewood clusters are predorninantly in wavy tangential hands (feature 6) and sornetimes temi to a diagonal patern (feature 7). Tangential ares o vesseis, typical of the Proteaceae (fig. 12), have heen cal]ed festooned. Since, in ring-porous temperate spccics, these patterns (features 6-8) may be resuicied tu the latcwood, their expression depends on ring width, and when rings are narrow these patterns are not obvious.

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VESSEL GROUPINGS 9. Vessels exelusively solitary (90% or more) 10. Vesseis in radial multiples of 4 or more common 11. Vessel clusters common DeJinitions: Vessels exclusively solitary = 90% or more of the vessels are completcly surrounded by other elernents, i. e., 90% or more appear not to contact another vessei, as viewed in cross section, fig. 21, e. g., Aspidosperina quebracho (Apocynaceae), Caraipa spp. (Bonnctiaceae), Eucalyptus regnans (Myrtaceae), Malus sylvestris (Rosaceae), Schirna wallichii (Theaceae). Radial multiples o! 4 or more cornmon = radial files of 4 or more adjacent vesseis of common occurrence, fig. 22, e.g., Cerberafloribunda (Apocynaceae), Ilex aquifliwn (Aquifoi iaceae), Brachylaena hutchin.sii (Compositae), Elaeocarpus hookerianus (Elaeocarpaceae), Strychnos nux-vomica (Loganiaceae), Arnyris balsamjfera (Rutaceae), Gambeya excelsa (Sapotaceae). Clusters common = groups of 3 or more vesseis having both radial and tangential contacts, and of common occurrence, fig. 23, e.g., Polyscias elegans (Araliaceae), Pirtosporurn ferrugineum (Pittosporaceae), latewood of Glediisia triacanihos, Gymnocladus dioica (Caesaipiniaceae), Morus alba (Moraceae), and Ailanrhus altissima (Sirnaroubaceae). Com.'nenzs.' Feature 10 'radial multiples of 4 or more common' should be used only when radial muitiples of 4 or more are an obvious feature of the transverse section. Feature 11 'clusters common' applies only when clusters are frequent enough that they are easily observed during a quick scan of a cross section. Clusters and radial rnultiples of 4 or more are not rnutually exclusive and can occur in combination. Woods with vesseis in tangential bands (feature 6) ofren have clusters. The most common vessel grouping is radial multiples of 2 to 4 wirh a variable proportion of solitary vessels (fig. 24). The absence of features 9-11 automatically irnplies this condition, When describing a wood, ao index of vessel grouping can be calcuiated in the manner recomrnended by Carlquist (1988): count the total number of vesseis in a minimum of 25 vessel 'groups' (i.e., count both solitary vesseis and vessel multiples as a 'group'), divide the total number of vesseis by 25 (the number of groups counted). An index of 1.00 indicates exclusively solitary vessels, and the higher the index, the greater the degree of vessel grouping. Caution: Care is needed to recognise rhe foliowing as not being multiples: (i) solitary vesseis composed of vessel elements with oblique overlapping end walis giving the appearance ofvessel pairs on the cross section as in Cercidiphyllurn (Cercidiphyllaceae) and Illicium (llhiciaceae), and (ii) closely associated solitary vessels, as in some species of Eucalyptus (Myrtaceae) and Calo- phyllurn (Guttiferae) (Brazier & Franklin 1961). Fig. 21. Vesseis 'cxclusivcly solitary' (feature 9), Aspidosperrrza quebracho, x 45. - Fig. 22. Radial rnultiples of 4 or more cominon (feature 10), Elaeocurpus hookerianus, x 29. Fig. 23. Clusters common (feature 11), Gymnocladus dioica, latewood, x 140. - Fig. 24. Vesseis partly solitary, partiy in radial multiples of 2-4, or very small c1usters (features 9, 10, and 11 absens), Drypetes gerrardii, x 75.

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SOL1TARY VESSEL OUTLINE 12. Solitary vessel outline angular Definition.

Solitary vessel outline angular = shape oU so]iary vessel outline is angular as viewed in cross section, fig.25, e.g., Aexroxkonpuncratum (Aextoxicaceae), Cercidiphyllwnjaponicum (Cercidiphyllaceae). Iatewood vesseis of white oaks (e. g., Quercus alba, Q. roburFagaceae), Stemonurus luzoniensis (Icacinaceae), Hortonia spp. and Mol1ini spp. (Monimiaceae).

Procedure: In ring-porous woods, examine the latewood because in these woods the earlywood vesseis are almost always circular to oval in outlije. Use Lhe outline of the solitary vesseis because the cornrnon walis of vesseis in inultiples can be flattened giving pari oU the vesseis ao angular oulune. Cornmen:s: Absence of feature 12 implies chai Lhe vessel outlinc is circular to oval (fig. 26) as in Banara regia (Flacourtiaceae) and the latcwood oU red oaks (e. g,, Que rcusfcdcara - Fagaceae). Caution: For fossil or archaeological samples, use this tature only when there obviously is no distortion from shlinkagc or post-depositional events. Distortion and 'folding' ofthe rays mdicates that the wood has been compressed during burial and that vessel outline probably has heen aliered. Fig. 25. Solitary vessel outline angular (feature 12), Stemonurus luzoniensis, x 75. -- Fig. 26. Outline of vessels rounded (feature 12 absent), Banara regia, x 45.

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13. Siniple perforation plates 14. Scalarifor'm perforation plates 15. Scalariform perforation plates with 10 bars 16. Scalariform perforation plates with 10-20 bars 17. Scalariform perforation plates with 20-40 bars 18. Scalariform perforation plates with ^ 40 bars 19. Reticulate, forarninate, and/or other types of multiple perforation plates

Definizions: Simple perforation plate = a perforation plate with a single circular or elliptical opening, g. 27, e. g., Aesculus hippocasranwn (Hippocastanaceae), Entandrophragrna spp. (Meliaceae), Prerocarpus spp. (Papilionaceae), Zellwva spp. (Ulmaceae). Scalariform perforation plate = a perforation plate with elongated and parailel openings separated by one to many mainly unbranched bars, lig. 29, examples follow. Scalariforrn perforation plates with :^ 10 bars, e. g., Corytus aveilana (Corylaceae), Goupia spp. (Goupiaccae), Liriodendron tulipifera (Magnoliaccae), Coula edulis (Olacaceae), Rhizophora mangle (Rhizophoraceae). Scalariform perforation plates with 10-20 bars, e.g., Betula verrucosa (Betulaceae), Altingia excelsa, Liquida,nbar styracflua (Harnamelidaceae), Saco glortis gabonensis (1 lumiriaceae), Schi,na wallichii (Theaceae). Scalariform perforation plates with 20-40 bars, fig.29, e.g.,Cercidiphyllum japonicum (Cercidiphyllaceae), Dicoryphe sripulacea (l-Iamamelidaceae), Nyssa ogeche (Nyssaceae), Staphylea pinn.ara (Staphyleaceae).
Scalariform perforation plates with ^ 40 bars, e.g., Aexroxicon puncrazum (Aextoxicaceae), Hedyosmwn spp. (Chloranth aceae), Dillenia iriquerra (Dileniaceae). Reticuate perforation plate = a plate with closely spaced openings separated hy wall portions that are much narrower than the spaces between them, or with a profuse and irregular branching of wall portions resulting in a netlike appearance, fig. 30, e. g., Didymopanax morotoroni (Araliaceae), Iryant hera juruensis (Myristicaceae). Forarninate perforation plate = a plate with circulas or elliptical openings like a sieve: the rernaining wall portions can be thicker than n the reticulate type, fig. 3 1, e. g., Oroxylum indicum (Bignoniaceae).

Other types = for instance, complex or radiate perforation plates, see comments and figs. 32-35. Co,nments: Determine the type(s) of perforation plate fron radial sections or maccrations, preferahly examine at least 25 vessel elements. For scalariform perforation plates, record ali the fcature categories thar encompass the range of the number of bars. Feature 14 'scalariforrn perforation plates' is a general category included to accommodate infortnation from cxisting literature that indicates whether scalariform perforation platcs are prescnt, but not the number of bars. Feature 14 is to be recorded wi th other appropriate features for bar number (15 18).

Fig. 27. Simple perforation plates (fcature 13), Aesculus hippocastanum, x 105. - Fig. 28. Simple perforation plate and scala.riform perforation plate with 2 bars (features 13, 14, 15), Didynopanax morotoronj, x 115. - Fig. 29. Scalariforrn perforation plate with 20-40 bars (features 14 and 17), Staphylea pinnata, x 220. Fig. 30. Reticulate perforation plate (feature 19), Didymopanax ,norototonj x 115. - Fig. 31. Foraminate perforation platc (feature 19), Oro'.hvn indu ,iun, 115.

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v.

Simple perforations are the most common Lype of perforation plate, and occur in over 80% of the world's woods (Wheeler et ai. 1986). Most woods have exclusively sirnple perforations, some have simple perforations together with scalariforin and/or other types of multiple perforatiori plates, and still others have exclusively scaiariform perforalion plates. When more tban one type of perforation plate is present (fig. 28), ai] types should be recorded and may be uscd to identify ao unknown (e.g., Didymopanax mororotoni Araiiaceae, Oxydendron arboreumEricaceae, Fa,us sylvatica - Fagaceae, and Piaranus occidenralis - Platanaceae have both simpie and scalariform plates). In those woods with both simple and scalariforrn perforalion plates, the narrower vessel elcmcnts and the latewood vessel elements are more likely to have scalariform perforatioii piares. Scalariforrn, reticulate, and foraminate piares forni a continuum, and the larter mo are often confused in the literature. Reticuiate and forarninate plates are restricted to re]ative]y ew taxononiic groups and are cornbined here. Reticulate perforations frequent]y occur itt combinatiori with scalariform piares and are an elaboration of that type. !ryanthera (Myristicaceae), Dendropanax and Didymopanax (Araliaceae) have scaiariforrn, reticulate, and varied intermediates plus simpie perforations; Myrceugenia estreilensis (Myrtaceae) has simple and niuldplc plates and the larter can be variousiy described as irregular-scaiariforni, forarninate, or even reticulate; Markhainia and Oro.xylu.rn (Bignoniaceae) have simple and foraminate platcs. !ryanthera (Myristicaceae) also lias cornpound scaiariform piares with few coarse bars with scts of fine secondary bars hetween them, which are often branchcd. Similar examples occur in Didymopanax marototoni (Araliaceae) and Ternsrroemia serram Theaceae). In these cases, botli feature 14 (scalariform perforation piares present) and feature 19 appiy. As pointed out by Cariquist (1988), the temi 'ephedroid' shouid not be used for foraminate perforations in dicotyledons, Radiate perforation piates (alto feature 19) with a central wall and radiating simp]e and branched bars extending to the lateral vessel wall are found in Cyiharexylurn myrianthwn (Verbenaceae) (Vidal Gomes ex ai, 1989) and Caryocar microcarpum (Caryocaraceae). Other types of multiple perforations may be found in the future and should be recorded as feature 19, 'reticulate, forarninate, and/or other types of rnultiple perforations'.

34 Fig. 32. Perforation plate obliquely compound scalariforrn with anasiornosing bars (features 14 and 19), Jryanthera paraensis, x 290. Fig. 33. Perforation plate regularly reticulate (feature 19, often occurring together with feature 14, scalariform perforarion p]ates), IryantherajuruensLs, x 290. - Fig. 34. Perforation plate cornpiex scalariforni and reticu]ale (features 14 and 19), Iryanthera elliptica, x 290, - Fig. 35. Radiate perforation piatc (feature 19), Cyrharexyiwn myrianrhum, x 300 (SEM).

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INTERVESSEL PITS: ARRANGEMENT AND SIZE 20. Intervessel pits scalariform 21. Intervessel pits opposite 22. Intervessel pits alternate 23. Shape of alternate pits polygonal Intervessel pit size (alternate and opposite) 24. Minute - ^ 4 im 25. Small 4-7 im 26. Medium - 7-10 im 27. Large - ^ 10 im 28. Range of intervessel pit size (J.Lm) Definitions: Intervessel pits = piLs between vessel elements. Scalariform intervessel pus = elongated or linear intervessel pits arrange.d in a ladderlike series, fig. 36, e.g., Dillenia reiiculara (Dilleniaceae), Michelia compressa (Magnoliaceae), Laurelia spp. (Monirniaceae), R/zizophora spp. (Rhizophoraceae). Opposite intervessel pits = intervessel pits arranged in short to long horizontal rows, i. c., rows orientated transversely across the Iength of the vessel, figa. 37, 44, C. g- Liriodendron spp. (Magnoliaceae), Nyssa ogeche (Nyssaceae). Alternate intervessel pits = intervessel pits arranged in diagonal rows, figs. 39, 40, 42, 43, e. g., Aceraceae, Mappia racemosa (Icacinaceae), Leguininosae, Mel iaceae, Salix spp. (Salicaceae), Sapindaceae, Shape of alternate pits polygonal = outline of intei-vessel pits, as seen in surface view (longitudinal sections), angular and with more than 4 sides, fig.40, e. g., Salicaceae, most Leguminosae. [ntervessel p11 sim (alternate and opposite) = horizontal diameter of a pit chamber at the broadest point
Z3(

& .--]=-.

: --;
2

40

Procedure:
Generaily, surface views of intervessel pits are easiest to find in tangential sections because radial multiples are the most frequent type of vessel multiple, and so intervessei pits are rnost frequent in tangenlial walis. When vesseis are in tangential bands and/or clusters, radial sections also provide surface views of intervessel pits. In woods with (almost) exelusively solitary vesseis, intervessel pits will be extimely rare, and often not visible in a single longitudinal Fig. 36 Intervessel pits scalariform (feature 20), Michelia compressa, x 115. - Fig. 37. Intervessel pits opposite (feature 21), Liriodendron tulipifera, x 115. Fig 38. Intervessel pits scalariform to opposite (features 20 and 21), Ilex laurina, x 350. --- Fig. 39. Intervessel pits alternate (feature 22), Mappia racernosa, x 112. Fig. 40. Shape of alternate pits polygonal (feature 23), Salix sp. Note also features 22 (pits alternate) and 26, 27 (intervessel pits medium to large); x 290. Fig. 41. Shape of intervessel pits circular t o oval (feature 23 absent). Note also plIs opposite (feature 21), Nothofagus moorei, x 300. Fig. 42. Intervessel pits minute, less rhan 4 p.m (feature 24), Polyalthia oblongifolia. Note also pits alternate (feature 22), x 300. Fig. 43. Vesturesl intervessel pits (feature 29), Terminalia sp., x 825. Fig. 44. Pits seerningly vestured due to presence of soluble deposits (feature 29, vestured pits, ahsent). Note also pits opposite (feature 21). Ilex cyniosa, x 800.

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section. In such woods, intervessel pit shape and size must be observed in overlapping end wall portions of vessel elements in a single vessel. However, in woods with vessel multiples, lhe pit arrangemcnt, shape, and size is best determined frorn lhe middle of lhe larger vessel clements. Measure ten pits, avoiding exceptionally large or small pits, and record those size classes that encompass lhe range ofpit size. Comments: Alternate intervessel pitting is lhe most common, and opposite and scalariform intervessel pitting are found in relatively few groups. When alternate pits are crowded lhe outlines of lhe pits tend to be polygonal in surface view; if alternate pits are not crowded thcn lhe outlines are usually circular to oval. \Vhen opposite intervessel pits are crowded lhe outlines of lhe borders tend to be rectangular in surface view. Some spccies have both polygonal and circular to oval intervessel pit outlines, record featurc 23 present for such woods. Combinations of different pitting patterns and/or intergrad.ing types occur (e.g., figs. 38, 41, alternate and opposite in Buxus - Buxaceac, opposite and scalai-iform in Liquidam bar - Hamamelidaceae) and may be indicated by using combinations of Lhe different pit fcatures. Pit size can help distinguish between genera within a family and between fainilies, e. g_ many Meliaceae have minute pits, while many members of the Anacardiaceae have large pits. The most widely used convention for determining pit size is to measure horizontal pit diame ter. To enable use of existing data this pararneter is included in this feature list. However, within several taxa, particularly those with some or ali opposite to scalaiiform pits, vertical diameter is a more constant and diagnostic feature, and it is recommended that this dirnension be recorded in a description. Cauzion: Do n'oL mistake vessel-vasiceniric tracheid pitting for intervessei pitting. VESTURED PITS 29. Vesturcd pits Definition: Vestured pits = pits with lhe pit cavity and/or aperture wholl y or partly lined with projections from lhe secondary ccli wall, fig. 43, e. g., Comhretaceae, Lythraceae, Myrtaceae, Rubiaceae, most Legurninosae. Procedure: Vestures are best viewed in water or glycerin mounts (or SEM). Bleaching is recornmended 50 as to remove encrusting materiais that nlay be mistaken for vestures (fig. 44 shows pseudovestures), i.e., soak sections (or, for SEM observation, wood blocks) in any household bleach until Lhe section or surface has lost its colour, rinse in water, and finish saniple preparation. Comments: Vesturing may occur in intervessel, vessel-ray or vessei-axial parenchyma, intertracheid, or interfibre pits. Vestures generaily are characteristic of entire farnilies, or groups within a famiiy. The number, size, and distribution of vestures varies considerably and these variations may be of diagnosuc value (Baiiey 1933; Ohtani eral. 1984; Van Vliet 1978; Van Vliet & Baas 1984). When intervessel pits are large and Lhe vestures are coarse (e. g., Terminalia spp. - Combretaceae), vestures are relatively easy to see with an oii-immersion objective of a good compound microscope. But when lhe vestured pits are minute (4 irn or less) as in lhe Apocynaceae or Rubiaceae, lhe vestures are difficuit to see with a compound microscope, and only clearly visible with a scanning electron microscope.

30. Vessel-ray pits with distinct borders; similar to intervessel pits in size and shape throughout the ray ceil 31. Vessel-ray pits with much reduced borders to apparently simple: pits rounded or angular 32. Vessel-ray pits with rnuch reduced borders to apparently simple: pits horizontal (scalariform, gash . like) to vertical (palisade) 33. Vessel-ray pits of two distinct sizes or types in the sarne ray celI 34. Vessel-ray pits unilaterally compound and coarse (over 10 jm) 35. Vessel-ray pits restricted to marginal rows Dejlnitions: Vessel-ray pits = pits between a ray celi and a vessel eicmcnt. linilaterally compound pita = pits in which one pil abuts two or more smalier pits in the adjacent celi. Other features as per descriptors, examples follow. Vessel-ray pits with distinct borders; similar to intervessel pts in size and shape throughout the ray cell, figs. 45, 46, e.g., Aceraceae, Leguminosae, Meliaceae, Jiex aquifoliwn (Aquifoliaceae), Betula spp. (Betulaceae), Camprostemon philippinense (Bombacaceae), Couratari cf. oblongifolia (Lecythidaceae). Vessel-ray pita with much reduced borders to apparently simple: pita rounded or angular, figs. 47, 48, e. g., Elaeocarpus calomala (Elaeocarpaceae), Clinostemon spp. (Lauraceae), Eucalyptus spp. (Myrtaceae). Popular spp., Salix spp. (Salicaceae). Vessel-ray pits with much reduced borders to apparently simple: pits horizontal (scalariform, gash-like) to vertical (palisade), figs.49, 50, e.g., Trigonobalanus vertici/lata, Quercus spp. (Fagaccae), Atherosperma moschata, Laurelia aromatica (Monimiaceae), HorsJieldia subglahosa (Myristicaceae), Syzygiwn spp. (Myrtaceae). Vessel-ray pita of two distinct sizes or types in the sarne ray cell, figs. 51, 52, e.g., some species of Erythroxylum (Erythroxyiaceae), Anacolosa spp., Chaunochiton spp. (Olacaceae), Santa/um spp. (Santalaceae), Planchoneila spp. (Sapotaceae). Vessel-ray pits unilaterally compound and coarse (over 10 Lm), fig. 53, e.g., Michelia champaca (Magnoliaceae), Ceriops spp., Kandelia spp., Rhizophora spp. (Rhizophoraceae). Vessel-ray pits restricted to marginal rows, fig. 47, e. g., Carpinus bezulus (Corylaceae), Aesculus hippocasranum (Hippocastanaceae), Populus spp., Salix spp. (Salicaceae). Comrnenrs: Vai-ious combinations of lhe above features may occur and should be recorded. Vessei-ray pits in lhe body of lhe ray may differ from those in lhe ray margins (e. g., Pala quium gaiatoxylum - Sapotaceae). Record lhe features for both types of pits. lia wood has predoniinantly solitary vesseis, comparison of vessel-ray pits with intervessel pita often is not possibie. li lhe vessei-ray parenchyma pits in such woods are uniform in size and shape and have borders, then use feature 30; ifnot, any of features 31-35 may apply. Vessel-axial parenchyma pitting usually resembles vessel-ray parenchyrna pitting, and is therefore not inciuded as a separate iist of alrnost identical descriptors.

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255

Kffi

V!1i *

'jpT
Fig. 52. Vessclray pits of two distinct sizos or types iii the sarne ray ce.l1 (feature 33); thc large pits (arrowcd) resembie perforations, Chaunochiton brevifloruin, x 290. - Fig. 53. Vesselray pits uni]aterally compound and coarse (feature 34), Ceriops raa1 (differentiai interference contrast), x .450,

Figs. 45 & 46. Vesselray pits with distinct borders, similar to intervessel pits (feature 30). 45: Courazari cf. obIongfolla, x 290. - 46: Carnpwste,non philippinense, x 75. Figs. 47 & 48. Vesselray pits wir.h rnuch reduced borders to apparently simple, pit outline rounded (feature 31), 47: Salix sp. (Salicaccae). Note also feature 35 (vesselray pits restrictcd to marginal rows); x 290. - 48: Elaeocarpus calornala, x 290. - Figs. 49 & 50. Vessel-ray pits with rnuch reduced borders co apparently simp]e, pits horizontal (gash-Iikc) to vertical (palisade), fc.ature 32. - 49: Pits horizontal, Atherosperrna ,noschata, x 450. - 50: Pics vertical, Trigonobalanus verticillara, x 290. - Fig. 51. Part of the vessel-ray pits with much reduced borders, and pits of Iwo distinct sizes or types in the sarne ray ccli (features 32 and 33), I-Iorsfieldia suhglobosa, x 115.

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HELICAL THICKENINGS 36. Helical thickenings in vessei elements present 37. Hetical thickenings throughout body of vessel element 38. Helical thickenings only in vessel element tails 39. Hetical thickenings only in narrower vessel elements Definitions: Helical thickenings in vessel etements = ridges on the inner face of the vessel elemeni wall in a roughly helical pattern. Synonym: spiral thickenings. Other features as per descriptors, exarnples follow. Helical thickenings throughout body of vessel etement, figs. 54, 55, e.g., Acer spp. (Aceraceae), Aesculus spp. (Hippocastanaceae), Cyrisus scoparius (Papilionaceae), Prunus spinosa (Rosaceae), Tilia spp. (Tiliaceae). Helical thickenings onty in vessel element tails, figs. 56, 57, e. g., Cercidiphyllum japonicwn (Cercidiphyllaceae), Liquidambar styraczflua (Hamamelidaceae). Helical thickenings only in narrower vessel elements, e.g., Robinia pseudoacacia (Papilionaceae), Ulmus americana (Ulmaeeae). Commenrs: Helicai thckenings are rather variable in terms of thickness (fine to coarse), inclination angle (nearly horizontal to steeply inclined), branching (branched or unbranched), and spacing (dose to wide). li is recomxnended that observations on these features be included in wood descriptions. Feature 36 'helical thickenings in vessel elements' is included as a general category to 1) adcommodate information frorn existing databases that indicate whether helical thickenings are present, but not their specific location; and 2) to help with the identification of small wood fragments in which vessel elements have helical thickenings, but because of the small sample size it cannot be determined whether Lhe helical thickenings are in ali vessel elements, or just in Lhe narrower ones. Feature 36 should be recorded in combination with other appropriate features (features 37, 38 or 39) for helical thickenings. Helical thickenings can also occur in vascular! vasicentric tracheids, and in ground tissue fibres (feature 64), and very rarely in axial parenchyma. Caution: Do not confuse coalescent pit apertures with helicai thickenings.

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Figs. 54 & 55. Helical thickenings throughout body of vessel elernent (features 36 and 37). 54: Prunus spinosa, x 290. - 55: cyrisus scoparius, x 220. Figs. 56 & 57. Helical thickenings only in vessel elemeni tails (features 36 and 31), Cercidiphvllurn japonicuin. - 56: x 150. 57: x 240.

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IAWA List of microscopic features for hardwood identification VESSELS PER SQUARE MILLIMETRE 46. 47. 48. 49. 50. 51. !^ 5 vessels per square millimetre 5-20 vesseis per square miltimetre 20-40 vesseis per square millimetre 40-100 vesseis per square millimetre ^ 100 vessels per square millimetre Mean, +I Standard Deviation, Range, n = x

259

TANGENTIAL DIAMETER OF VESSEL LUMINA Mean tangential diameter of vessel lumina 40. ^ 50 jim 41. 50-100 gm 42. 100-200 im 43. ^- 200 tm
44. Mean,

+I .

Standard Deviation, Range, n =

Procedure:
Vessel diameter is measured in transverse sections. Vesseis are selectcd for measurement with care not to bias the selection towards the larger or smaller vesseis. The rangential diameter of the vessei luniina, excluding the wali, is measured at the widest part of Lhe opening. At least 25 vesseis should be measured. Ining-porous woods (feature 3) and woods with 'vcssels of two distinct diameter classes, wood not ring-porous' (feature 45), only measure and record the larger size class. Information about tangential diameters of the smailer vesseis wouid be useful in a description. In semi-ring-porous woods, measure along a radial transect through a growth ring. For semi-ring-porous woods, it is recommended that more than 25 vessels be measured; a larger standard dcviation is expected for such woods. Use the category(ies) in which the mean(s) fali(s). Comments: In trees, mean tangential diameters of 100-200 im are more cornrnon than mean tangential diarneters greater than 200 im or mean tangential diameters less than 50 J.inl. In shrubs, mean tangential diameters of less than 50 jim are conimon.

Procedure:
Ali vcssels are counted as individuais, e. g., a radial multiple of four would be counted as four vesseis (Wheeler 1986). Count ali the vesseis in at least tive (and preferably ten) fields of appropriate size (depending on vessel diameter and distribution), and convert to number per square rniliimetre, i. e., for woods with small diameter vessels use fields 1 mm x 1 mm or iess; for woods with large vesseis that are widely spaced use whole fields of view at low magnifica. tion (e. g., 4 x objective lens). Of the vesseis that are partially in Lhe field of view, only 50% are included in the count. If vessei frequency is very low, examine enough fields to account for local variations, and preferably count at least 100 vesseis. Use the categories that include the total range of vessel frequency. Cornmenrs: Vessel frequency is not computed for ring-porous woods, or for woods with their vesseis ia definite tracts with vascular/vasicentric tracheids, e. g., dendritic pattern as seen in Rhamnu.s cathartica (Rhamnaceae), or tangential bands as seen in Ulmus (Ulmaceae).

MEAN VESSEL ELEMENT LENGTH 45. Vesseis of two distinet diameter classes, wood not ring-porous Definirion: Vesseis of two distinct diameter classes, wood not ring.porous woods with a bimodal distribution of tangential diameters of vessel lurnina, fig. 58, e. g., Actinidia spp. (Aclinidiaceae), Capparis maroniensis (Capparidaceae), Derris hylobia Papilionaceae), Serjania subdenrata (Sapindaceae), Congea ro,nenrosa (Verbenaceae). Comrnen ts: Vines and xerophytes often Baas & Schweingruber 1987). 52. 53. 54. 55. ^ 350 l.m 350-800 ^ 800 im Mean, +1 Standard i)eviation, Range, n = x

Procedure:
Measure the whole iength of each vessel element from one tail end to thc other, preferably in a maceration. AL least 25 vessel element lengths are measured to derive the mean and range. Use the categoly(ies) in which the mean(s) fali(s).

have

vesseis of two distinet diameter classes (Carlquist 1985;

TYLOSES ANO DEPOSITS IN VESSELS 56. Tyloses common 57. Tyloses sclerotic Definitions: Tyloses common = outgrowths from an adjacent ray or axial parenchyma ccli through a pit in a vessel wall, partially or completely blocking the vessel lumen, and ofcQmmon OccUlTence

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261

(except in outer sapwood), figs. 59, 60, e. g_, Anacardium occidentalis (Anacardiaceae), Acanrhopanax spp. (Araliaceae), Cercidiphyllum japonicum (CercidiphyUaceae), Eucalyptus acmeniaides (Myrtaceae), Sti-ombosia pustulata (Oiacaceae), Robinia pseudoacacia (Papiionaceae). Selerotie tyloses = tyloses with very thick, multilayered, lignified walis, figs. 61, 62,
e, g., Chaetocarpus schoni burgki anus, Micrandra spruceana (Euphorbiaceae), Cantleya cornicu-

lata (Icacinaceae), Eusideroxylon zwageri (Lauraceae). Brosimum guianense (Moraceae). Procedure: In ring-porous woods, it is best to examine the earlywood vesseis for tyloses because ty10es are often absent froni small diarneter latewood vesseis. Avoid sapwood when determining the presence of tyloses or gums. Commenrs: Tyloses may be few or many, ranging from ali vesseis fihled with many tyloses to a few vesseis with a few tyloses. Feature 56 applies only when tyloses are not of sporadic occurrence. Tyloses may be thin-walled or thick-walled, pitted or unpitte, with or without starch, crystals, resins, gums, etc. Such information should be recorded in a description. Woods with selerotie tyloses usually have thin-walled tyloses as well, and both features 56 and 57 tnay apply Some woods may have both tyloses and guni deposits (feature 58), and both features 56 and 58 may apply. Caution: Absence af tylases is not diagnostic Do um code lraumatic tyloses such as occur in wound heartwood and be careful rim to confuse tyloses with foamy deposits, masses of fungi, ar other deposits.
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58. Gums and other deposits in henrtwood vesseis Cornments: In cross sections, deposits appear to completely fihi some vessel lumina (fig. 63); in longitudinal sections, deposits often appear to collect at the end of vessei elements. Deposits often can be seen more clearly by exarnining the woods with a hand lens; sectioning and rnounting techfiques may remove some of the deposits. 'Gums and other deposits' includes a wide variety of chemical compounds, which are variously coloured (white, ye]iow, red, brown, black). Ia a description it is appropriate to indicate their abundance and colour. See Hillis (1987) for more informadon on the chemistry of deposits. Cauton: Use feature 58 positively oniy. Do not confuse masses of whitish fungi, which ma y be packed in a vessel cavity, ar sclerotic tyloses with deposits.

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Fig. 58. Vesseis of two disdnct diameter classes, wood not ring-porous (feature 45), Serjania subdentata, x 35, - Figs, 59 & 60. Tyloses common (feature 56). 59: In transverse section, Anacardiurn occidentalts, x 220. - 60: Ia tangential secdon, Robinia pseudoacacia, x 140. Figs. 61 & 62. Tyloses sclerotic (features 56 and 57), Cantleya corniculata. 61: Transverse scction, x 75. 62: Tangential section, x 75. Fig. 63. Gums/dcposits in heartwood vesseis (feature 58), Physocalymma scaberrimum, x 110.

62

-:

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IAWA Bulietin n.s., Vol. 10 (3), 1989

IAWA List of microscopie features for hardwood identification

263

WOOD VESSELLESS 59. Wood vesselless Definition: Wood vesselless = wood without vessel elements, cornposed only of imperforate tracheary elements and parenchyma, figs. 64, 65, e. g., Amborellaceae, Tetracentraceae, Trochodendraceae, Winteraceae. Comnients: Vesselless dicotyledonous woods are relatively uncornmon and are distinguished from coniferous wood by tall multiseriate rays. For vesselless woods, it is not necessary to cocle for type of imperforate tracheary elements (fibres or vascular/vasicentric tracheids) and impossible to code for vessel features.

TRACHEIDS AND FIBEES 60. Vascular/vasicentric tracheids present Deflnitions, Vascular tracheids = imperforate edis resembling in size, shape, pitting, and wall ornamentation narrow vessel elements and intergrading with the lattcr, figs. 66, 67, e.g., Sambuctis nigra (Caprifoliaceae), Sop hora arizonica (Papiionaceae), Phellodendron amuren.se (Rutaceae), Lycium europaewn (So!anaceae), Celtis occidenralis (Ulinaceae). Vasicentric tracheids = imperforate celis with numerous distinctly bordered pits in their radial and tangenhial walis, preserir around the vesseis, and different from ground tissue fibres, often, but not always of irregular shape, fig. 68, e. g., Castanea spp., Quercus spp. (Fagaceae), rnany Shorea (Dipterocarpaceae) and Eucalyprus (Myrtaceae) species. Comments: Vascular tracheids often occur in association with extensive vessel rnultiples or clusters, especially in the latewood. A very thorough search of macerations will reveal their presence iri tnany species. 1-lowever, for wood identification purposes use rhein only when they are commonly present. The intergradation of vascular tracheids with narrow vessel elements implies thar there are some celis with a single, often very small, perforation. Some anatomists would prefer to cail such edis vascular tracheids, others would prefer to cail them narrow vessel elements, probably terminating a ves.sel. Because they have a perforation such cells are best referred to as vessei elements; tracheids are imperforare edis. The IAWA Glossary (1964) includes shortness and irregular form in the definition of vasicentric tracheids, but these criteria do nor always apply (e.g., iii Eucalyptus spp., cf. lhe 1987). Since vascular tracheids are often intermixed with vessels (i.e., in a vasicentric position) in many taxa, they can also be considered as vasicentric iracheids. s1

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67 Figs. 64 & 65. Wood vesselless (feature 59), Trochodendron aralioides. - 64: Transverse section, x 38. - 65: Tangential section, x 38. - Figs. 66-68. Vascular/vasicentnc tracheids (feature 60). 66: Radial section, vessei elements surrounded by vascular tracheids (note also helicai thickenings in vessel elements, features 36 and 37), Phellodendron a.'nurense, x 290, 67: Maceration, vessel element flanked by two vascular tracheids, Celtis occidentalis, x 300. 68: Radial section, vasicentric tracheids, Castanea sariva, x 170.

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GROUND TISSUE FIBRES 61. Fibres with simple to minutely bordered pits 62. Fibres with distinctly bordered pits 63. Fibre pits common in both radial and tangential walis Defini rions: Fibres with simple to minutely bordered pits = fibres (libriform fibres) with simple pits or bordered pits with the chambers less than 3 tm in diameter, flgs. 71, 72, e. g., Swietenia spp. (Meliaceac), Inga spp. (Mimosaceae), Fraxinus spp. (Oleaceae), Populus spp. (Salicaceae). Fibres with distinctly bordered pits = fibres (or fibre-tracheids or ground tissue isacheids) with bordered pits with chambers over 3 p.m in diameter, figs. 69, 70, e.g., 1/ex spp. (Aquifoliaceae), Dilienia spp. (Dileniaceae), Illiciu.m spp. (llliciaceae), Xanrhophyllum spp. (Polygalaceae), Carneilia spp. (Theaceae). Fibre pits common in both radial and tangential walis = fibre pits, either bordered or simple, common in radial and tangential walis, e. g., [lex spp. (Aquifoliaceae), Dilienia spp. (Dilleniaceae), 11/id um spp. (Illiciaceae), Xanthophyllum spp. (Polygalaceae), Clematis vitalba (Ranunculaceac), Carne/lia spp. (Theaceae). Procedure: Determine the nature and distribution of fibre pits only in longitudinal (radial and tangential) sections, because in cross section many fibre walis are not strictly radial or tangendal. Both longitudinal and cross sections are suitable to determine if the pits are bordered or (almost) simple. Commen:s: The feature 'fibres with distinctly bordered pits' partly overlaps with the descriptors 'tracheids' sensu Bailey (1936) and Carlquist (1986a, 1986b, 1988) and 'fibre-tracheids' sensu Baas (1986). It usually coincides with 'fibre pits common in both radial and tangential walis' (feature 63). The following combinations are of very sporadic occurrence: 1) fibre pits not distinctly bordered, ie., pit chambers less than 3 j.un or pits simple, feature 61 present, and pits comrnon in radial and tangential walls, feature 63 present, e. g, Capparis spinosa (Capparidaceae), Nyctanthes arbor-tristis (Oleaceae), Vitis vin(fera (Vitaceae); 2) fibres with distinctly bordered pits, feature 62 present, and pits mainly restricted to the radial walis, feature 63 absent, e. g.. Elaeagnu.s angu.safolia (Elaeagnaceae). Two types of fibres with respect to wall pitting (both features 61 and 62 present) may occur (e. g, some species of Vaccinium - Ericaceae). Fibres with simple to minutely bordered pits (feature 61 present), mainly confined to the radial walis (feature 63 absent) are libriform fibres in the deflnition of Baas (1986) or libriform fibras and/or fibre-tracheids in the deflniuon ofCarlquist (1986a, 1986b, 1988). The terms libriform fibras, fibre-tracheids, and 'true tracheids' have been deliberately avoidcd as descriptors in this list because there is no consensus on their deflnitions. ii

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Figs. 69 & 70. Fibres with distinctly bordered pits (feature 62), common in both radial and t:mgential walls (feature 63), tangential sections. - 69: lilicium carnbodianu,n, x 230. - 70: Xant/i phyllurn lanceatu,n, x 230. - Fig. 71. Fibrcs with simple to minutely bordered pOs (feature 61), tangential section, phase-contrast, Populus sp. (Salicaceae). Note also .scarcity ofpits in tangential walis (feature 63 ahsent); x 410. Fig. 72. Fibrcs with simple to minutely bordered pOs (feature 61), common in both radial and tangcntial walis (feature 63). Tangential seclion, phase-contrast, Clernatis viralba, x 410. - Figs. 73 & 74. Helical thickcnings iii ground tisue fibres (fet:re 64). - 73: 1/ex cjnerea. x 850. 74: JIe. chinensis, x 850.

Er266 IAWA Balietin n.s., Vol, 10 (3), 1989 1 AWA List of rnicroscopic features for hardwood identification 267

64. Helical thickenings in ground tissue fibres Definition.Helical thickenings in ground tissue fibres = as per feature descriptor, see definition of helical thickenings, feature 36, tigs. 73, 74, e.g., Euonynus europaeus (Celasti-accae), Haname1isjaponica (1 larnamelidaceae), Cercocarpus ledifolius (Rosaceae), and tcmperate species of 1/ex (Aquifoliaceae) and Syringa (Oleaceae). Com ments. Fibres with helical thickenings usualty occur in woods that also have he]ical thickenings in the vessel elements. However, the opposite is not true, Le., many species with helical thickcnings in their vessel elernents do not have helical thickeriings in the ground tissue fibres. Helical thickenings are rnuch more cornmon iri fibres with distinctly bordered pits than in tibres with sinnple to rninutcly bordered pits. They also occur more frequently in tempera te woods thari in tropical woods,

SEPTATE FIBRES AND PARENCHYMA-LIKE FIBRE BANDS 65. Septate fibres present 66. Noriseptate libres present 67. Parenchyma-Iike fibre bands alternating with ordinary tibres Definitions: Septate fibres = fibres with thin, unpitted, transverse wall(s), flgs. 75-78, e.g., Spondias mombin (Anacardiaceae), A ucournea kaineana (Burseraceae), Buchenaia capitara (Combretaccae), E!aeocarpus spp. (Elacocarpaceae), Aglaia spp. (Meiaeeae), Virex orinocensis (Verbenaceae). Nonseptate fibres = tibres without septa. Parenchyma-like libre bands alternating with ordinary fibres = tangential bands of relatively thin-walled fibres alternating with bands o thicker-wallcd fibres, fig. 79, e. g., Cassine spp., Mayxenus obtusifoila (Celastrace ae), Dubauria spp. (Compositac), Lagersiroemia tomernosa, Physocalymma spp. (Lythraceae), Cupania americana (Sapindaceae). Com menu: Septa are formed after lhe secondary fibre waJls have been deposited; they therefore do not extend to Lhe compound middle larnellae between adjacent fibres, Septa are usua]Iy unlignified and veiy thin (cf. Pararneswaran & Liese 1969). In some woods, ali fibres are septate (feature 65 present, feature 66 absent), e.g., fig. 76, Lainea welwirschii, Spondias niombin (Anacardiaceae), Canariwn schwemnfiirthii (Burseraceae). In other woods, both septate and nonseptate flbres occur together (features 65 and 66 boih present), eg., fig. 78, Buchenavia capirata (Conibretaceae), E/aro carpus spp. (Elaeocarpaceae), Swie:enia macrophylla (Meliaceae). The septate fibres rnay then either be scaitered irreguIarly, situated near the vesseis or the rays (feature 67 absent), or arranged in tangential bands (feature 67 presem).

Figs 75-77. Septate fibres presem (feature 65). 75 & 76: Ali iibres septane, severa] septa per fibre, Aucouniea klaineana, - 75: x 290 76: x 75 - 77: Many septa per fibre, Agia/a lirtora/is. x 150 Fig, 78. Septate (arrows) and nonseptate fibres presem iii lhe sarne sanipie (features 65 and 66), Swietenja inacrophyIla, > 115. - Fig. 79L Parenchvtna-iike (ihi'e bands (feature 67). Phys cah'nnma .scabcrrimu,n, x 45.

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IAWA List of microscopic features for hardwood identification

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The fibres of parenchyma-like fibre bands (featore 67) are usuaily septate; lhe ordinary fibres they alternate with may be nonseptate (feature 66) as in Cassine maurocenia, Maytenus obrusifolia (Celastraceae), or septate as in Lagersrroernia tornentosa, Physocalymma scaberrimurn (Lythraceae). In lhe laner case, features 65 and 67 are presem, feature 66 is ahsent. Fibre septation is coded independently of fibre wall pining (features 61-63). Thc number of septa per fibre can vary from 1 to many. This number can be taxon specific (e.g., Van Vliet 1976b), and so lhe average number of septa per taxon shouid be given in a description.

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Caurions: Do not confuse tom ccli wali fragments, ceil wail deformations, gum deposits, fungal hyphac, or tyloses in fibres (seen in some Magnoiiaceae, Lauraceae) with septa. Sometimes parenchyma strands can also easily be confused with septate fibres. Avoid tension wood, because gelatinous fibres are nonseptate.

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FIBRE WALL THICKNESS 68. Fibres very thin-walled 69. Fibres thin- to thick-walled 70. Fibres very thick-walled DeJinitions: Fibres very thin-wailed = fibre lumina 3 or more times wider than lhe double wall thickness, fig. 80, e. g., in Bursera simaruba (Burseraceae), Te:ra,neles nudzflora (Datiscaceae), Neubergia corynocarpa (Loganiaceae), Ti/ia japonica (Tiliaceae). Fibres Ihin- to thick-walled = fibre lumina less than 3 limes the double wall thickness, and distinctly open, fig. 81, e, g., in 1/ex spp. (Aquifoliaceae), Michelia compressa (Magnoliaceae), Salix alba (Salicaceae). Fibres very thick-walled = fibre lumina a]most compietcly closed, fig. 82, e.g., in Goupia glabra (Ceiastraceae), Lophira spp. (Ochnaceae), Strombosia pusxulwa (Olacaccac). Kr/4giodendronferreurn (Rhainnaceae), Rhizophora tnangle (Rhi zophoraceae). Commen:s: Measurement of lhe actual thickness of fibre walls usually invoives an amount of work ou[ of ali proportion to lhe iimited diagnostic value of the figure obtained. Therefore, lhe classes for fibre wall thickness are based on lhe ratio of lumen to walt thickness (Chattaway 1932). The ratio proposed is that of lhe width of lhe lumen to lhe combined thickness of lhe walis between it and lhe lumen of lhe next ccli as viewed in cross section. When celis are flattened radiallv the lumen becomes oval and will give a different ratio with lhe wail according to whether it is measured radiaily or tangentially; lhe radial measurement is suggested. Chattaway (1932) proposed four categories; three are used here. Feature 68 roughiy corresponds to her category very thin'-walied; feature 69 includes her two categories 'thin' and 'thick'; feature 70 is identical to her category 'very thick'. Fibre wall thickness in many species is variable and there may be more than one category of fibre wall thickness in a species.

,
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1 8' '

Fig. 80. Fibre.s very thin-walled (feature 68), Neuburgia corynocarpa, x 290. - Fig. 81. Fibres thin- to thick-walled (feature 69), Michelia compressa, x 290. - Fig. 82. Fibres very thickwalled (feature 70), Rhizophora rnangle, x 290.

Cauzions: In woods with distinct growth rings, fibre wail thickness changes throughout lhe growth ring, and rnay be particularly thick aI lhe end of lhe growth ring. When describing fibre wali thickness, do ,tot consider these iast latewood fibres. Also, do nol dcscribe gelatinous fibres (tension wood fibres), which usual]y liave thick walls with an unlignified gciatinous iayer,

MEAN FIE3RE LENGTI IS 71. 900 j.un 72. 900-1600 j.irn 73. ^ 1600 j.tm 74. Mean, +I Standard Deviation, Range, n

= x

Procedure: Use macerations of mature trunk wood, and measure lhe length of at lcast 25 fibrcs to de termine lhe mean, range, and standard deviation. Use lhe category(ies) in which lhe mear-i(s) fali(s). For woods with distinct growth rings, sample from lhe middie of lhe growth ring. Because of lhe importance of ccli length in wood quality studies, a variety of rnethods have bccn developed to insure random selection of edis for measurement. It is recommended that one of these rncthods be used (Dodd 1986; Hart & Swindel 1967). There are very few woods in which fibre len h :In Tnca1Ired acc uratclv from sccion s. so such a method n O rLc( ,rnrncndcd.

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AXIAL PABENCHYMA General commenls: When identifying an unknown, use the most obvious type of parenchyma pattern first and then the less evident type or types. Be sure to use a broad ficld of view when deterrnining the predorninant parenchyma pattern(s) from the n-ansverse section. Various combinations of the three general types (Apotracheal, Paratracheal, and Banded) described below may be prescrit in a given wood.

75. Axial parenchyma absent or extremely rare Definizion: Axial parenchyrna absent or extremely rare = as per feature descriptc,r, fig. 83, e.g., Berberidaceae, Punicaceae, Violaceae, Homaliu,nfoetidwn, Sconellia coriacea (Hacourtiaceac), Sonneratia spp. (Sonneratiaceae). Comment: li is necessary to study longitudinal sections in combinajion with transverse sections to be sure axial parenchyma is absent or extremely rare (i.e., very difficult to find; only a few stands per section). This feature may be used in combination with 'axial parenchyma scanty paratracheal' (feature 78) and/or 'axial parenchyma diffuse' (feature 76), if, despite the scarcity of parenchyma strands, the distribution is clear.

APOTRACHEAL AXIAL PARENCFIYMA 76. Axial parenchyma diffuse 77. Axial parenchyma diffusein.aggregates Dejinirions: Apotracheal axial parenchyma = axial parenchyma not associated wth the vessels. Axial parenchyma diffuse = single parenchyma sirands or pairs of strands distributed inegularly among the fibrous elements of the wood, fig. 84, e. g., Aspdosperrna polyneuron (Apocynaceae), A/nus glutinosa (Betulaceae), Goupia glabra (Celastraceae), Cornus mas (Cornaceae), Apodytes di,nidiata (Icacinaceae), Crataegus spp. (Rosaceae), Santalum album (Santalaceae). Axial parenchyma diffuse-in-aggregats = parenchyma strands grouped into short discontinuous tangential or oblique lines, fig. 85, e.g., Durio spp. (Bombacaceae), Hura crepitans (Euphorhiaceae), Ongokea gore, Strombosia pusrulata (Olacaceae), Agonandra brasiliensis (Opiliaceae), Dalbergia sreven.sonii (Pap ilionaceae), Prerospermum spp. (Sterculiaccae), Tilia spp. (Tiiiaceae). Fig. 83. Axial parenchvnia absent or extremely rare (featurc 75), Homaliu,nfoezidum, x 75. Fig. 84. Axial parenchyma diffuse (feature 76), Alnus glutinosa,x 115. - Fig. 85. Axial parenchyma diffuse-in-aggregates (feature 77), Agonandra brasiliensis, x 45. - Fig. 86. Axial parenchyma scanty paratracheal (feature 78), and aiso scanty diffuse (feature 76), Dilienia pulcherrima,x 115.

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Co,nmenrs: Because there is a continuous range from parenchyma extrernely rare, diffuse, diffuse-inaggregatcs, to parenchyma in narrow bands (feature 86) or scalariform (feature 88), for some taxa it wil] be necessary to record more than one feature for apotracheal parenchyma. Diffuse and diffusc-in-aggregates frequently occurincombination. Record (1944) referred to diffusc-inaggregates parenchyrna as reticulate. This list does not follow that usage, but uses rcticulate to describe a type of barided parenchyma, sce feature 87. Caurions: Although by defiuiition apotracheal parcnchyma is not associated with vesseis, woods with abundant diffuse or diffuse-in-aggregate parenchyma rnay exhibit several strands touching ihe vessels. Such random contacts should not be recorded as parairacheal parenchyma. Apotracheal diffuse parenchyma sornetimes occurs primarily near the rays ('ray adjacent parenchyrria' of Carlquist 1988), and shou]d not be confused with sheath celis in rays (feature 110),

PARATRACHEAL AXIAL PARENCHYMA 78. Axial parenchynia scanty paratracheal 79. Axial parenchyma vasicentrie 80. Axial parenchyma aliform 81. Axial parenchyma lozeuge-aliform 82. Axial parenchyma winged-aliform 83. Axial parenchyma confluent 84. Axial pa renchyma unilateral pa rat racheal Definitions: Axial parenchyma paratracheal = axial parenchynia associated wih the vesseis or vascular tracheids; Lypes of paratache.al parenchyma are scanty paratracheal, vasiccntric, aliforrn (subtypes: lozenge-aliform, winged-a]ifonn), confluent, and unilateral paratracheal. Axial parenchyma scanty paratraclieal = occasiojial parenchyma celis associated with the vesseis or an incornplete sheath ofparenchyma around Lhe vesseis, lig. 86, eg. Pistacia vera (Anacardiaceae), Scierolohium spp. (Caesalpiniaceae), Dillenia puicherrima (Dillerriaccae), Eiythroxylu,n ,nannii (Erythroxylaceae), Laurus nobilis (Lauraceae). Axial parenchyma vasicentric = parenchyma celis forining a complete circular to oval sheath around a solitary vessel or vessel multiple, lgs. 87, 88, e. g. Tachigali myr,necophylla (Caesalpiniaceae), Octomeles sunwrana (Datiscaceae), Phoebe porosa (Lauraceae), Khaya grand(foliola (Mcliaceae), Anadenanrhera spp., Ent-erolobiwn cyclocarpwn, Pipradeniastru,n africanum (Mimosaceae), Olea europaea (Oleaceae). Axial parenchyma aliform = parenchyma surrounding or to one side of the vessel and with lateral extensions. For examples see Lhe two subtypes below. Axial parenchytna lozenge-aliform parenchyma surrounding or to orie side of the vessels with lateral extensions forming a diamond-shaped outline, fig. 89, e.g., Albizia lebbek. Parkia gigantocarpa (Mimosaceae), Arlocarpus chaplasha (Moraceae), Microberlinici brazzavillensis, Ormosia fiava, Vataireci spp. (Papilionaceae), Quaiea rosca (Vochysiaceae) Figs. 87 & 88. Axial parernhyrna vasjcentric (featurc 79). - 87: Parenchvuia sheath broad,
Pip fade niastrum africanum, x 26. - 88: Parenchyma sheaih narrow, Khavn grandifoliola, x 45.

flii .tiIJ(

- Fig. 89. Axial parenchyma lozenge-aliforrn (features 80 and 81). Microberlinia brazzavillensts, x 29. - Fig. 90. Axial parenchyma winged-aliform (features 80 and 82), Brosimu.m rubescens, x 45,

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Axial parenchyma winged-aliform = parenchyma surrounding or to one side of the vesseis with the ]ateral extensions being elongated and narrow, fig. 90, e, g. Jacaranda copaia (Bignoniaceae), Termina/ia superba (Combretaceae), Brosimum spp. (Moraceae), Quassia amara (Simaroubaceae), Gonystylus spp. (Thymelaeaccae). Axial parenchyma confluent = coalescing vasicentrie or aliform parenchyrna surrounding or to one side of two or more vesseis, and often forming irregular bands, figs. 91, 92, e. g., Kigelia africana (Bignoniaceae), Caesalpiniaferrea, Peito gyne confertifiora (Caesalpiniaceae). Marmaroxyion racemosum, Parkia pendula (Mimosaceae), Ch/orophora tinctoria (Moraceae), Bowdichia nitida,Vatairea guianensis (Papilionaceae). Axial parenchyrna unilateral paratracheal = paratracheal parenchyma forming semicircular hoods or caps only on one side of the vesseis and which can extend tangentially or obliqucly in an aliform or confluent or banded pattern, fig. 93, e.g., Aspidosperrna desman. thum (Apocynaceae), Caraipa grandiflora (Borinetiaceae), Peito gyne conferiflora (Caesalpi fiaceae), Mammea bongo (Guttiferae), Dilobeia thouarsii (Proteaceae). Comments: Scanty paratracheal includes what has been described iti the literature as iricomplete vasiceninc. Some woods have vasiccntric, aliform, and confiuent paratracheal parenchyma. ConflueLu often intergrades with banded and may be recorded or used in combination with features for hand width (85-86). Feature 80, 'Axial parenchyma alifornf, is included as a general catcgory to 1) describe those woods that clearly have aliforni parenchyma, but in which it is difficult to decide whether it is lozenge- or winged-alifonn, and 2) allow use ofinformation from the literature that does not differcntiate between these two types. Features 81 and 82 are used in combination with frature 80. Feature 84 'unilateral paratracheal parenchyma' is used in cornbination with aliform and/or confluent when the unilateral parenchynia extends lateraily or obliquely. Unilateral includes both abaxial and adaxial because generally ii is not possible to discinguish between the two in a wood fragment. Woods with several types of paratracheal parenchyma co-occurring and/or intergrading have been assigned the general descriptor 'parenchyma predominantly paratracheal' by several authors (fig. 94). Caution: Vasicenti-ic/vascular tracheids are often thinner-wal]ed than ground tssue tlbres, and in cross sections may be confused with axial parenchyma. Examine longitudinal sections to detennine whether vasicentric/vasctdar tracheids or axial parenchyma surrounds the vesseis,

1-'igs. 91 & 92. Axial parcnchyrna conflucnt (frature 83). 91: Parkiapen(Iuia arrowhead, note also features 80 and 81, parenchyma ]ozenge-aliforrn), x 36, 92: Peltogyne confertiflora (note also feature 84, unilateral parenchyma), x 45. Fig. 93. Axial parenchyma unilateral paratracheal (feature 84), Caraipa grandiflora, x 45. - Fig, 94. Axial parcnchyma prcdominantly paratracheal, Garcinia iwissirna (note also features 80, alifonii, and 83, confluent), x 45.

276 BANDED PARENCHYMA 85. 86. 87. 88. 89.

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Axial parenchyma bands more than three cells wide Axial parenchyma in narrow bands or tines up to three cells wide Axial parenchyma reticulate Axial parenchyma scalariform Axial parenchyma in marginal or in seemingly marginal bands

Definitions: Parenchyma bands more than three edis wide = as per feature descriptor, fig. 95, e.g., Dicorynia paraen.sis (Caesalpiniaceae), Entandrophragma candoliei (Meliaceae), Ficus retusa (Moraceae), Lophira alata (Ochnaceae), !Jasyloxylon brasiliensis (Sterculiaceae), Erisma uncinarwn (Vochysiaceae). Parenchyma in narrow bands or lines up to three cells wide = as per feature de-

scriptor, figs. 96, 98, e.g., Dialium guianensi.s (Caesalpiniaceae), Endospermum malaccensis
(Euphorbiaceae), Bertholletia excelsa (Lecythidaceae), Dysoxylum frasera num (Meliaccae), Aurranelia congolensis (Sapotaceae), Hanwa klaineana (Simaroubaceae). Parenchyma reticulate = parenchyma in continuous tangential lines of approxirnately the sarne width as the rays, regularly spaced and forming a network with them. The distance between the rays is approxirnately equal to the distance between the parenchyma bands, lig. 97, e. g., Cleistopholjs spp. (Annonaceae), Diospyros discolor (Ebenaceae), Bertholletia excelsa, cariniana spp., Couratari guianensis, Eschwei lera spp. (Lecythidaceae). Parenchyma scalariform = parenchyma in fairly regularly spaced fine lines or bands, arranged horizontally or in arcs, appreciably narrower than thc rays and with them producing a ladder-Iike appearance in cross section. The distance between the rays is greater than the distance between parenchyma bands, figs.99, 100, e.g.,Anisophyllea spp. (Anisophylleaceae), Onycho. petalum sp. and most other Annonaceae, Cardwellfa sublimis, Embothrium mucronatum (Proteaceae), Rhopalocarpus spp. (Rhopalocarpaceae). Parenchyma in marginal or in seemingl y marginal bands = parenchyma bands which fonn a more or lcss continuous Iayer of variable width at the margins of a growth ring or are irregularly zonate, figs. 101, 102, c. g- Jntsia hijuga (Caesalpiniaccae), Juglans regia (Juglandaceae), Cryptocarya moscJzaza (Lauraceae), Liriodendron zulipifera, Michelia compressa (Magnoliaceae), Cedrela spp., Swierenia spp. (Meliaceae), liorsfieldia subglobosa (Myristicaceae). Comments: Parenchyma bands may be mainly independent of the vesseis (apotracheai), definitely associated with the vesseis (paratracheal), or both. Bands may be wavy, diagonal, straight, contifluous or discontinuous (the latter often intergrading with confluent). The nurnber of bands per mm varies and may be useful as a diagnostic feature in some groups. Bands over three celis wide are visible to the unajded eye. For woods with reficulate, scalariform, or marginal parenchyma, the band width (either f-atures 85 or 86) also should be recorded. In the past, some anatomists (e. g., Record 1944) have used the term reticulate for abundant diffuse-in-aggregates parenchyma with numerous short interrupted lines. Sometimes marginal parenchyma bands are associated with axial interceliular canais. In some temperate woods there are discontinuous bands/lines of parenchyma at the growth ring margin; this condition also should be described as 'marginal'. Marginal parenchyma includes terminal and initial parenchyma, and seemingly marginal includes what has been called irregular zonate bands. Fig. 95. Axial parenchyrna iii hands more than three edis wide (feature 85), Ficus retusa, x 45. - Fig. 96. Axial parenchyma in narrow bands or lines up to three edis wide (feature 86). .4utranella congolensis, x 45.

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O,

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Figs. 101 & 102. Axial parenchyma in marginal or seetningly marginal hands (feature 89). 101: Inisia bijuga, Note also features 80, 81 (lozenge-aliform parenchyma), 83 (confluent), and 86 (narrow bands); x 29. - 102: Michelia compressa, x 45.

t?f

td2

A.

Fig. 97. Axial parenchyrna reticulate (feature 87), 3ertholleria excelsa (note also feature 86, parenchyma in narrow bands), x 45. - Fig. 98. Axial parenchyrna intermediate between reticulate and scalanform (featurcs 87 and 88 variable), Couratari guianensis, x 45. - Figs. 99 & 100. Axial parenchyrna scalariforrn (feature 88). - 99: Onychopetalum sp., x 45. - 100: Parenchyma bands also 'festooned', Cardwellia sublirnis, x 15.

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AXIAL PARENCHYMA CELL TYPE/STRAND LENGTIT 90. 91. 92. 93. 94. Fusiform parenchyma cells Two edis per parenchyma strand Four (3-4) celis per parenchyma strand Eight (5-8) edis per parenchyma strarid Over eight ceiis per parenchyma strand

Definition.: Fusifoi-m parenchyma = parenchyma celis derived from fusiform cambial initiais without subdivisions or tip growtli. In shape they resembie a short fibre, fig. 103, e.g., Capparis spp. (Capparidaceae), Aeschynornene elaphroxvlon, Erythrina spp., Lonchocarpus spp. (Papilionaceae), Trplochiion scieroxylon (S tercu] iaceae), Buinesia spp., Guiacuin spp., Zygophyllurn spp. (Zygophyllaceae). Parenchyma strand = a series of axial parenchyrna cells forrned through transverse division(s) of a single fusiform cambial initial cd. Two cells per parenchyma strand, figs. 103, 104, e.g., Dalhergia spp., Lonchocarpus spp., Pterocarpu.s spp.(Papilionaceae), Four (3-4) celis per parenchyma strand, fig. 104, e.g., Termina/ia spp. (Combretaceae), Ligusrtim spp., Syri;iga spp. (Oleaceae), Nesogordonia spp. (Sterculiaceae). Eight (5-8) edis per parench yma strand, fig. 105, e.g., Nerium oleander (Apocynaceae), Macaranga spp. (Euphorbiaceae), Fraxinus spp. (Oleaceae). Over eight celis per parenchyma strand e.g., Bhesa spp. (Celastraceae), Lophira spp. (Ochnaceae), Minquartia spp., Tetrasylidum spp. (Olacaceae). Commenrs. Type ai' parenchyma, fusiform vs, sirand, is dctermined frorn tangential sections. Fusiforni parenchyma celis are relatively uncomrnon and generaily occur in woods with storied structure and short axial elements. In some species, cornbinations of the above fea[ures occur, e. g. fiisifopn cdlls' and two cel]s per parenchyma sirand', or 'twa cel]s per parcnchyma srxand' and 'four (3-4) celis per parenchyma strand'. Strand length can diferbetwcen earlywood and latewood of the sarne ring, ar between vessel-associated parcnchyma and parenchyrna which is not in concact with the vesseis. Record ali cornrnonly occurring strand lengths. Cawion.' Be careful not to confuse uniseriate rays or septate fibres with strand parenchyrna. Do nor determine number of ce]ls per strand from chambere.d crystalliferous sirands. 95. Uniignifled parenchyma Definition: Uniignitied parenchyma = as per feature descriptor, fig. 106, e. g, Apeiba spp., Ente/ca arborescens, Heliocarpus spp. (Tiliaceae), Laporteo srimtdans (Urticaceae). Conuneni: Unlignified parenchyma usually occurs in broad bands, and is resu-icted to a small number a taxa. Fig. 103. Fusiform parenchyma ce]Is (feature 90, left and right) and twa celis per parenchyrna su'and (feature 91, left centre). Note also feature 120 (axial parenchyrna storied) and fcatiire 142 (crystals in chambered axial parenchyina). Aeschynomcne e/aphroxylon, x 115. - Fig. 104. Two (feanire 91) ia fom (feature 92) edis per parenchyma strand, Cordia abyssinica, x 115. Fig, 105. Figiu (feature 93) ar over eight ceils (feature 94) per parenchyma strand, Bertholleia excelsa, < 115.Fig. 106. Un]ignified parcnchyma (feature 95), Ente/ea arborescens, x 45.

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RAYS

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RAY WIDTH
96. Rays exclusively uniseriate 97. Ray width 1 to 3 celis 98. Larger rays commonly 4- to 10-seriate 99. Larger rays cornmonly > IO-seriate 100. Ra ys with multiseriate portion(s) as wide as uniseriate portions Definitions: Ray width in celi numbers as per feature descriptors. Rays exclusively uniseriate, fig. 107, e. g, Lophoperalwn beccarianum (Celastraceae), Termina/ia superba (Combretaceae), Hura crepitans (Euphorbiaceae), Castanea sariva (Fagaceae), Populus spp. (Salicaceae). Ray width 1 to 3 celis, fig. 108, e.g., Aucoumea klaineana (Burseraceae), Dialium guianense (Cacsalpiniaceae), Alseodaphne costa/is (Lauraceae), Albizia (Samanea) saman (Mimosaceae), Malus co,nmunis (Rosaceae). Larger rays commonly 4- to 10-seriate, fig. 109, e.g., Acer saccharum (Aceraceae), Spondias mombin (Anacardiaceae), Anisoprera laevis (Dipterocarpaceae), Khaya anthotheca (Meliaceae), Celtis sinensis (Ulrnaceae). Larger rays comrnonly > 10-seriate, fig. 110, e.g., Quercus spp. (Fagaceae), Foraqueiba guianensis (Icacinaceae), Rapanea spp. Myrsinaceae), Platanus spp. (Platanaceac), Cardwe/lia sublimis, Grevi/lea robusta (Proteaceae), Tamarix aphyl/a (Tamaricaceae), Jacquinia revoluta (Theophrastaceae). Rays with multiseriate portion(s) as wide as uniseriate portions, fig. 111, e.g., Anthodiscus amaznicus, Caryocar cosraricense (Caryocaraceae), Srrombosja pustu/ata (Olacaceae), Ad/na cordifolia (Rubiaceae), and many Apocynaceae, Sapotaceae, and Euphorbiaceae. Procedure: Determine ray width on the tangential section by counting the number of celis in the widest part of the rays, perpendicular to the ray axis. When rays are of two distinct sizes (feature 103), record the width of the larger sire class in the database. Commen Is: 'Exclusivel y uniseriate rays' and 'rays> lO-seriate' are the Ieast comnion of the ray width features. The categories for ray width match those of the Clarke (1938) and the Princes Risborough multiple entry keys (Brazier & Franklin 1961). There are some taxa that may be separated by uinerdistinctions within these categories, and in adescription more detailed information than provided by these categories would be desirable. Caution: These features for ray width do not apply tu rays containing radial canais (feature 130) or to the rays composing an agegate ray (feature 101). Thus, for a wood with exclusively unisenate rays, except for the rays with radial canais, describe the wood by recording feature 96 for exclusively unisenate rays and feature 130 for radial canais.
104

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Fig. 107. Rays exclusivcly uniscriate (featiire 96), Lophopetalum beccarianuin, x 75. - Fig. 108. Ray width one to three celis (feature 97), Alhizia (Samanea) sarnan, x 29. - Fig. 109. Largcr rays commonly 4-10-seriate (feature 98). Note also shcath ecOs (arrow, feature 110); Celtis sinen-sis, x 29. - Fig. 110. Larger rays commonly> lO-seriate (feature 99), Cardwellia subi/mis, x 19. Fig. 111. Rays with miiltiseriate portion(s) as wide as uniseriate portions (feature 100), Caryocar costaricense, x 115.

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AGGREGATE RAYS 101. Aggregate rays DeJlnirion: Aggregate ray = a number of individual rays so closely associated with one another that they appear macroscopica]ly as a single large ray. The individual rays are separated by axial elements, e.g., many species ofAlnus (Bctulaceae), Carpinus, Cory!us (Corylaceae), Casuarjna (Casuarinaceae), Necepsia afzelii (Euphorbiaceae), Castanopsis, Lithocarpus, Quercus - evergreen species (Fagaceae), Enimotum orbiculatum (Jcacinaceae), Cryptocarya densflora (Lauiaceae). Commenrs: There is variation in the size of the individual rays of aggregate rays. In some specics the aggregate rays are composed of narrow rays (figs 112, 113, e. g., Carpinus spp. - Corylaceae), while in others they are composed of broad rays (figs. 114, 115, e. g, Empnotum orbiculatu,n lcacinaceae). Aggregate rays occur in few taxonomic groups. Caution: Aggregate rays may be relatively infrequent inche taxa in which they occur, so they may be easily overlookcd or absent in a small sample; therefore, this feature should preferably be used posiive1y only.

RAY HEIGHT 102. Ray heiglit > 1 mm Definition: Ray height> 1 mm = the large rays commonly exceeding 1 mm in hcight, e.g., Guatteria schomburgkiana (A nnonaceae), Anisoptera laevis (Dip terocarpaceae), Uapaca guineensis (Euphorbiaceae), Scotzellja coriacea (Flacourtiaceae), Barrington(a asiatica (Lecythidaceae), Piaianus occidentalis (Platanaceae), Paypayroia guianensis (Violaceae). Procedure: Determine total ray height in tangential section, along the ray axis. Comnzent: In this Iist of features, only one category for total ray height is used as was done in some of the earlier multiple entry keys (Clarke 1938; Brazier & Franklin 1961). More detailed ray height data generally are given in descriptions and may be helpful in distinguishing between taxa in some groups. Ray height is quite variable in some woods (particularly woods with markedly heteroceilular rays), bui quite uniform in others (particularly woods with storied strucwre). Figs. 112-115. Aggregate rays (feature 101). - 112 & 113: Carpinus herulus, aggregatc r,r: (ar) composed of narrow rays. - 112: Transverse section, x 20. - 113: Tangencial sectr. x 52. - 114 & 115: E,nmozum orbicularum. aggregate ravs (ar) composcd of rnu1tisriate 45. - 114: Transverse section. - 115: Tangential seclion.

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RAYS OF T\VO DISTINCT SIZES 103. Rays of two distinct szes Definition: Rays of two distinct sizes = when viewed in tangential section, rays form two distinct populations by their width and usually also by thcir height, figs. 116, 117, e.g., Acer saccharum (Aceraceae), Poga oleosa (Anisophylleaccae), Ilex aquifolium (Aquifoliaceae), Dillenia pentagyna (Dillcniaceae), Quercus spp. (Fagaceae), Dendrobwgia boliviana (Icacinaceae), Scaphiwn macropodum (Sterculiaceae), Ternsrroernja spp. (Theaceae). Commenis: There are no limits for the size classes - the smaller rays may be 1- or 2- or 3-seriate, the ]arger rays may be less than 5-seriate. Generaily, to fit the feature definition, intcrmecliate rays should not exist between the two populations or be quite rate. Thus, when very large rays occur with a fcw rnedium-sized and more numerous small rays (e. g, Fagu.$), feature 103 'rays of iwo distinct sizcs' may still be applied. Cautions: Vhether a wood has rays of two distinct widths cannot be deterrnined from the cross section because in this view the long uniseriate wings of heteroceilular multiseriate rays might bc interpreted incorrecdy as narrow rays. Aggregate rays per se houId not be considercd as a separate ray size class. Only iri those species where the aggrcgate rays are composed of much broader rays than thc nonaggregate rays does feature 103 apply, e. g,, severa] species of Ca.suarina (Casuarinaceae) and Quercus (Fagaceae).
1 ..

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E

-k

Figs. 116 & 117. Rays of two distinct sizes (feature 103). - 116: Ternstroemia sp., x 45. - 117;
Quercus gilva, x 29.

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28')

RAYS: CELLULAR COMPOSmON 104. Ali ray cells procumbent 105. Ali ray celis upright and/or square 106. Body ray ceiis procumbent with one row of upright and/or square niargina! ceils 107. Body ray ceiis procunibentwjth moslly 2-4 rows of upright and/er square marginal celis 108. Body ray celis procumbent with over 4 rows of upright and/or squarc marginal ceils 109. Rays with procumbent, square and upright celis mixed throughout tlie ray Definjrfons: Procumbent ray cefi = a ray parenchyrna ceil with its longest dirnension radial as seen in radial section. square as seen in radial section. Upright ray celi = a ray parenchyrna cel] with its lorigest dirnension axial as seen in radial section. AR ray cetls procumbent, fig. 118, e. g., Acer spp. (Aceraceae), Tabebuia spp. (Bignoniaceae), Alhizia spp. (Mimosaceae), Hannoa klaineana (Sirnaroubaccae) Ali ray celis upright and/or square, fig. 119, e.g., Hedyosmum scabrum (Chloranthaceae), Aucubajaponica (Cornaceae). Body ray ceils procumbent with one row ol upright and/or square marginai celis, fig. 120, e.g., Kalopanax pictus (Araliaceae), Aucm.mea klaineana (Burseraccae), Pseudocedrela kanschyi (Meiiaceae). Body ray celis procumbent with mostiy 2-4 rows of upright and/or square marginal celis, fig. 121, e.g., Liq uidainbarstyrac,Jlua (Hamameljdaceae), Carapa guianensis (Meiiaceac), Trecu lia africana (Moraceae), A/seis peruviana (Rubiaceae), Euscaphf.s spp, (Staphyleaceae). Body ray eeils procumbent with over 4 rows of upright and/or square marginal cells, fig. 122, e.g., Wein,nannia descendens (Cunoniaeae), Quintinia spp. (Escailortiaceae). HomaIiumfoetid (Flacourtiaceae), Hu.'niria spp, (Humiriaceae), O?oschulzj spp. (Icacinaceae), Coffea spp. (Rubiaceae), Turpinia spp. (Staphvleaceae). Rays with procumben, square and upright celis mixed throughout the ray, fig. 123, e. g, Guaerja spp. (Annonaccac),Xan,/jop1/u, lanceatum (Polygalaceae), Pometia pinna:a (Sapindaceae), Heliocarpus spp. (Tiliaceae). Square ray celi = a ray parenchynia ccli approximatel y

'T.
I1tlii t
Fig. 118. Ali ray edis procurnbent (feature 104), Acer cwnpcslre. x 115.-- Fig. 119. Ali ray ecOs upright anrifor squarc (feature 105), Aucuba japonica, x 29. Fig. 120. Body ra y cells procumbeni with oric row of upright and/ar square marginal edis (feature 106). Fseudocedrelu kotschyi, x 115. Fig. 121. Body ray celis procumhent with most]y two to oui' rows of upright and/or square marginal cells (eature 107), Campa guianen.sis.. x 114. Fig. 122. Body ray celis procurnbent with mostly over 4 rows of upright aiidlor squarc marginal celis (feature 108), Hornalium j.'etidwn. x 115. - Fig. 123. Rays with procumbeni, square and upright cclls rnixed throughout thc mv (feature 1091, Xwtthophyl!wn lanceatwn, x 90.

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Use radial sections LO determine the cellular composition of rays because types of ray celi (procurnbent, upright, and square) are defined on the basis of their appearance in radial section. Generaily, upright and square cens, if presem in eomhinatjon with procumberiL celis, are located ri Lhe marginal rows, i.e,, those rows at lhe top and bottom of lhe ray, and procumbcnt cells are ]ocatcd in lhe body (centre) of lhe ray. In woods wih uniseriate and rnultiscriate rays - desenhe lhe celhila.r cocnposition of l he muitiseni ate rays, not lhe uniseriate rays. Some woods have more than onc category of ray type with rcspect to celiular composition, only record lhe rclatively common categonies.
Cornmen13-

Cautions:
Ray composition often varies betwecn juvenile and mature wood. Ia many species, rays mar the pith may be composed entirely of upnight celis, while rays distant from lhe pith are composed largely of procumbent edis with only a few rows of upnight and/or square edis. When creadng a database, only examine mature wood samples or, for shrubs, lhe penipheral wood of lhe thickest available stems. When an unknown wood fragrnent is from a thin branch, do nor use ray composition. Although in tangential section, marginal rows of upright and/or square edis often will appear as uniseriate margins, lhe presence of uniseriate margins alone is not a reliable indicator of heteroceliular rays. Ia some woods (e, g., Carya spp. Juglandaceac), there are uniseniate mar-' -. ginal rows visibie in tangential section, and these celis appear larger xhan the body edis, but wheri viewed in radial section these edis are procumbent, as are lhe edis of the rnultiseriatc portion. Sheath celis (feature 110) or tile cells (feature 111) are not considered when deternilning ray ceiluiar composition. Feature 109 only appiies if there is a mixture ar alteration of different ray cdl shapes xhroughout the ray, irrespective of whether it is in uniseriate or nnildseniate rays or ray portions. Feature 109 does not appiy to woods with vertically fused rays (ar 'rays with alternating uniseriate and multiseriate portions') where the uniseriate portions may be composed of square and upright cells and lhe multiseniate portions ofprocumbent edis. Just lhe presence of sheath ce]is (feature 110) or tile edIs (feature lii) also does not qua]ify a wood for frature 109.

The ceiluiar colnposition of lhe niultiseriate and uniseniate rays in [f i e sarne wood is not necessariiy the sarne. In some woods, their uniseniate rays are cornposed oniy of upright celis, while thcir multiseniate rays are composed of both upright and procurabeni celis. Hornoceilular rays are rays coniposed of a single ccli type; heteroce]lular rays are composed of two ar more ccli types. Thc tens homogencous and hetemgenenus are used to describe ray tssue as a whole. Designating Krihs types (Kribs 1968) rnay be useful when describing a wood, but are not used in this Iist. Roughly spealdng, Kribs homogeneous types correspond with feature 104 (ali ra y celis procurnbent), Kribs heterogeneous 111with feature 106 (one row of upright and/ar .square marginal edis), Kribs heterogeneous II with fcature 107 (2-4 rows af upright and/or square marginal celis), Krihs heterogeneous 1 with feature 108 (more than 4 rows of upright and/or square marginal celis). H&wever, feature 108 also may partly ovenlap witb Kribs heterogcneotjs 11 II' ffie body of the rays is very high.

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SHEATH CELLS 110. Sheath celis DeJlnition: Sheath celis = ray cells that are located along thc sides of broad rays (> 3-seriate) as viewed in tangential section and are larger (generaily tailer than broad) than the central ray celis, figs. 124, 125, e. g., Ceiba pentandra (Bombacaceae), Cordia altiodora (Boraginaceae), Sainbucus nigra (Caprifoliaceae), Dipterocarpus lowii (Dipterocarpaceae), Stemonurus luzoniensis (Icacinaceae), Ailanthus altissima (Simaroubaceae), Sterculia oblonga (Sterculiaceae). Commenrs: Presence of sheath celis should be deterrnined from tangential sections. There is variability in the frequency and distinctiveness of sheath celis. In some species most, if not ali, multiseriate rays have sheath cells which are much larger than the other ray celis, while in others sheath celis are not frequent and/or slightly larger than the adjacent cells. When identifying an unknown wood sample, do not use this feature as a flrst lime of approach unless it is well mai-ked. Cautwn: Do not confuse sheath celis with tile cells (feature 111), which are always found in the body of the ray as weli as the edges and are visible in both tangential and radial sections.

IiuI11

TILE CELLS 111. Tile celis Definition: Tile celis = a special type of apparently empty upright (rarely square) ray celis occurring in intermediate horizontal series usually interspersed among rhe procumbent celis, figs. 126-129, e. g., Durio spp. Neesia aitissinia (Bombacaceae), Guazutna spp., Kleinhovia hospira, Prerospermum spp. (Sterculiaceae), Despiarsia spp., Moi/ia spp., and some species of Grewia (Tiliaceae). Commenrs: Tile edis sometimes have been classified into two groups: type Durio when they have the sarne height as the procumbent ray celis, and type Pterospermum when they are higher. However, this distinction is dubious because there are intergradations between the two, as in Guaruma (Sterculiaceae), and Grewia (Tiliaceae). Tile edis do not occur in uniseriate rays, and as far as is known are restricted to the order Malvales.

,L, ''11fr1

Figs. 124 & 125. Sheath celis (feature 110). - 124: Ceiba pentandra, x 75. - 125: Stemonuru luzoniensis, x 75. Figs. 126-129. Tile celis (feature 111). - 126-128: Durio type, Neesia alrissima, radial, transverse, and tangential section, x 115. - 129: Pterospermum type, Pterospermwn grewiaefoliurn, radial section, x 120.

Iii

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PERFORATED RAY CELLS 112. Perforated ray celis Definition: Perforated ray celis = ray celis of Lhe sarne dimensions or larger than the adjacent edis, but with perforations, which generaily are on the side walis connecting Iwo vesseis on either side of the ray, figs. 130-132, e. g., Combretwn Ieprosrachiuni (Combretaceae), Richeria racemosa (Euphorbiaceae), Chaunochiton breviflorum (Oiacaceae). Commenrs: The type of perforation in a perforated ray ccli may be simple, scalariform, reticulate, or forarninate, and does not necessarily coincide with the type of perforation plate occurring in the vessel elements of the sarne wood. For instance, Sloanea monosperma (Elaeocarpaceae) and Richeria racemosa (Euphorbiaceae) have simpie perforations in the vessei elernents and scalariform perforations in Lhe perforated ray cells. In Siparuna (Monimiaceae) there is a range of multiple perforations in thc ray celis, but the vessel elemcnt perforations are simple and/or scaiariform with variations depending on species. Pcrforated ray celis have bordcrcd pits similar to the intervessel pits. They can occur mdividually or in radial ortangential rows. Radial rows ofperforated ray edis with perforations in the tanendai walls have been described as radial vesseis (Van Vliet 1976a). Caution: Use this feature positively only and with some caution because spccies that have perforated ray celis may have thein in such low frequency that they could easily have bcen overiooked when creating a database, or exarnining an unknown.
1

DISJUNCTIVE RAY PARENCHYMA CELL WALLS 113. Disjunctive ray parenchyrna ccli walis Definirion: Disjunctive ray parenchyma ceil walis = ray parenchyma edis partialiy disjoined but with contacts maintained through tubular or compiex wall processes, fig. 133, e. g., Funtuinia africana (Apocynaceae), Buxus sempervirens (Buxaceae), Crozon oligandrus, Glycydendron a,nazonicum, Suregada laurina (Euphorbiaceae), Malpighia incana (Malpighiaceae), Gardenia imperialis, Randia armara (Rubiaceae). 135 Comrnent: Axial parenchyma nay also be disjunctive Figs. 130-132. Perforated ray celis (feature 112). - 130: Simple perforation in radial wai.I, Chaunochiron breviflorum, x 115. - 131: Reticulate perforation in radial wali, Richeria racemora, x 115. - 132, Sirnple perforation in tangential wali, Com brerum leptosrachium, x 290. -Fig. 133. Disjunctive ray parcnchyma ccli walis (feature 113). Malpighia incana, x 290. Figs. 134 & 135. Wood rayless (feature 117), t'eronica tr2erSii, x 115. 134: Transverse section. 135: Tangential section.
ft . u

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RAYS PER MILLIMETRE 114. 54/mm 115. 4-12/mm 116. ^ 121 mm Definirions: As per feature descriptors, examples follow. ^ 4/ mm, e. g., Guarreria schomburgkiana (Annonaceae), Cussonia arborea (Araliaceae). 4-12/mm, e. g., Acer rubrum (Aceraceae), Acacia spp. (Mimosaceae). ^: 12/ mm, e. g., Diospyros ,nesp il(for?nis (Ebenaceae), Randia armara (Rubiaceae). Procedure: The number of rays per linear unit is best determincd frorn a tangential section along a une perpendicular to the ray's axis; it can also be deterrnined from a cross section. Make ten measurements and record the categories the range falis within. Com ments: The feature 'rays 4-12 per mm' is more common than the features 'rays ^. 4 per mm' or 12 per mm' (Metcalfe & Chalk 1950). The number of rays per mm cannot sensibly be determined in woods with aggregate rays, or woods with very broad rays and mo distinct size classes, e. g., Quercus spp. (Fagaceae).

Wood rayless = wood with only axial clements, figs. 134, 135, e.g., Arrhrocne,nu,n ,nacrostachyum (Chenopodiaceae), Heimerliodendron brunonianum (Nyctaginaceae), Hebe sahczjolia, Veronica rraversii (Scrophulariaceae). Commenr: Rayless woods are restricted to a small number of families (Carlquist 1988). Caurion: In rayless woods with included phloem (e.g., several Chenopodiaccac. the conjunctive parenchyma (i.e., the parenchyma linking mo or more phloem strands) may forni radiai extensions which resembie rays. In such woods there rnav be a continuum from short radi1 wedges tu long radial strips to 'normal' rnuitiseriai ravs lhn r al. ) tl teLLue fd\ 1es' should he iised i[li caution in such 'oods

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118. 119. 120. 121. 122. 123.

Ali rays storied Low i-ays storied, high rays nonstoriect Axial parenchyma and/or vessei elemcrits storied Fibres storied Rays andlor axial elements irregularly storied Number of ray tier: par axial mm

Definitions: Storied structure = cclls arranged in tiers (horizontal series) as viewed on lhe tangenrial surface. Ali rays storied, 1lgs. 136, 138, e. g, Dalbergia bariensis, Prerocarpus sansalinoides (Papiiionaceae), Quassia amara (Simaroubaceae). Low rays storied, high rays nonstoried, fig. 137, e.g., Scaphium spp., Triplochiton scleroxylon (S tercul iaceae), Cercis canadensis (Caesalpiniaceae). Axial parenchyma and/or vessel elements storied, fig. 136, e.g, Balanites aegypdaca (Balanitaceae), Dalbergia bariensis, Sparziwnjunceum (Papilionaceae), Tarnarix spp. (Tamaricaceae). Fibras storied, 6g. 138, e. g, Quassia amara (Simaroubaceae), Ocwmeles sumacrana (Datiscaceae), Zygophyllu.in dumosum (Zygophyllaeeae). Rays and/or axial elements irregularly storied = stories of rays and/or axial elements not horizontal or straight, but wavy or oblique (synonym: in echelon), or only locally presem, fig. 139,e.g., certain Leguininosae (Monopetalantlius, Teiraberlinia), Entandrophragrna cylindricwn (Meliaceae), Fraxinus alba (Oleaceae), Tieghemella spp. (Sapotaceae). Number oF ray tiers par axial mm = as per feature descriptor. Co,nments: lhe The presence of storied structure should be deterrnined &om lhe tangential section, foi radial section. These features can be recorded singly or ia combination as in some woods ali elements are storied (e. g., Hibiscus tiliaceus -- Malvaccae, Cenrrolobiu,n paraense, Afrormosia data - Papilionaceae), whiie in cahers various combinations of elernents are storied. Generally, if parenchyma is storied, the vessel elements also are storicd. Featare 122, 'rays and/or axial elements irregularly storied', is used in combination with lhe other features when appropriate. There is variability within species and saniples. For instance, ia some samples ofSwietenia (Meliaccae) rays are defiriitely storied, in others irregulariy storied, and in still others rays are aol storied. Tiers of rays are visibie at low magnification, or with lhe unaided eye or a hand lens, and appear as fina horizontal striations or ripple marks on the tangencial surface. Especiafly ia lhe Leguminosae, which has many taxa with storied rays, the number of ray tiers per mm (feature 123) can be useful in distinguishing genera and specics. Absence of storied snuclure (features 118-123 abseni) is also oU diagnostic vaJue. Caution: Storying of wide vessel elements may be obscured because oU ceil enlargement during vessel development, and se it is best to examine lhe narrower vessel elements te determine whether vcssel elements are storied.

Fig. 136. Ali rays storied (fearure 118) and axial parenchyrna and/ar vessel elements storied (feature 120), Dalhergia bariensi.s, x 75. Fig. 137. Low rays storied, high rays nonstoried (feature 119), Triplochiton scleroxylon. Note also features 120 (parenchyma storied) and 121 (fibres storied); x 45. - Fig. 138. Fibras storied (feature 121), Quassia amara, x 45. - Fig. 139. lrregularly storied structure (feature 122, hera expressed as rays ia echeion), Entandrophragma cylindricwn, :x 45

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OIL AND MUCILAGE CELLS 124. Oil and/or mucilage celis aSsociated with ray parenchyma 125. 011 and/or mucilage edis associated with axial parenchyma 126. Oil and/or mucilage celis present anlong fibres Definitions:
011 celi = a parenchymatous idioblast fihled with ou; mostly, but not always, enlarged and rounded in outhne, occasiona]ly of considerable axial extension, figs. 140, 141, e.g., Nectandra grandis, Ocotea glaucinia, O. tendia, Phoebe porosa and many other species of Lauraceae, Talaur,ia spp. (Magnoliaceae),

ii !

.1ttIiI Ij mtt

Mucilage ccli = a parenchymatous idiob]ast fihlcd with rnucilage; typically enlarged and rounded in outline, occasionally of considerab]e axial extension (resernbling fibres), e.g., figs. 142, 143, axial iii some specics of Endlicheria (Lauraceae), and in ray parenchyma of some species of Persea (Lauraceae). Co,nments Both oil cells and mucilage celis are commonly associated with axial and/or ray parenchynia, but may also occur among fibres. They are limired to very few woody dicotyledons and are similar to one another, except for iheir contents, which are easily removed during rnicrotechnical procedures (Richter 1977). Because ii is not practical to distinguish beween oil and mucilage edis by thcir appearance, they are listed together. Various combinations of these features eccur togciher (see Baas & Gregory 1985; Gregory & Baas 1989),

11J II

1.
1

4 'j

Fig. 140. Oil and/or mucilage cdlis associated with ray pare nchyma (feature 124), Ocoiea giaucinia, x 115. Fig. 141. Oil and/or mucilage edis associated with axial parenchyma (feature 125, arrows), Ocotea glaucnfa. Note also feature 124, ujl and/or mucilage edis in rays; x 45, - - Figs. 142 & 143. Oil and/or mucilage cells presem among the fibres (feature 126), Ocorea ccneilci (elongated inucilage celis), x 115. 142: Transverse secdon. - 143: Radial section.

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1NT1TRCELLI5LAR CANALS 127. 128. 129. 130. 131. Axial canais in long tangential lines Axial canais in short tangential lines Axial canais diffuse Radial canais Intercellular canais of traumatic origin

D/initions, Interceilular canal = a tubular interceilular duct surroundcd by an epitheiium, gencrally containing secondary piam products such as resins, gums, etc., secretcd by the epitheliai celis. Interceiluiar canais may be oriented axialiy (axial/vertical interceliular canal), or radially (radial/ horizontal interceilular cana], within a ray), figs. 144-151. Synonyms: gum duct, resin duct. Axial canais in long tangential iines = more than five canais in a une, fig. 145, e.g., Copaifera spp., Sindora spp. (Caesaipiniaceae), Dryobalanops spp, flopea spp., Neohalanocarpus spp., Parashorea spp., J-'enracme spp., Shorea spp. (Dipterocarpaceae). Synonym: concenic axial canais. Axial canais in short tangential lines = two to five axial canais in a une, fig. 146, e. g., Diprerocarpus spp. (Dipterocarpaceae). Axial canais diffuse = randorniy distributed soiitary canais, fig. 147, e. g., Prioria copaifera (Caesaipiniaceae), Anisoptera spp., Cotylelohium spp., Upuna spp., Vaiaria macrocarpa, Varica spp. (Dipterocarpaceac). Radial canais = canais presem in rays, figs. 148-150, e.g., Pistacia spp., Tapirira guianen.sis (Anacardiaceae), Bursera gummzfera (Burseraceae), Shorea section Richetia (Dipterocarpaceae), and rnany other genera of Anacardiaceae and Burseraceae. Synon ym: horizontal canais. Trauniatic canais = canais formed in response to injury, arranged in tangential bands, generaiiy irregular in outline and ciosely spaced, fig. 151, e. g., Terminaliaprocera (Combrctaceae), Liquidambar sryracflua (Harnamciidaceae), Carapa procera (Meliaceae), Prunus serorinu (Rosaceae), Balfourodendrorz riedelianwn, Murraya exolica (Rutaceae), Quassia amara (Simaroubaceae). Cornments: It is possibie to have a mixture of thesc features in one wood. In some species of Dipterocarpaceae, the size of the axial canais is useful in differentiating species, i.e., whether the canais are small (diameter < 100 xm) or largc (diameter> 100 im). The colour of resins in canais of Dipterocarpaceae can also be usefui in identif-ication. The effect of radial canais on ray shape (i.e., whether the canal niakes the ray fusiform in shape or not), size, and number of canais per ray are also usefui features. Caurion: Traumatic canais may not occur consistently in a species, therefore, when identifying an unknown, never use the absence of traurnatic canais.

Li
,_, '

$:

FI 1 z nu; :

Fig. 144. Axial interceliular canal, Parashrea srn)rhiesii, x 290, Fig. 145. Axial intcrcellular canais in iong tangential hnes (fcature 127), Shorea parsfo1ia, x 29. - Fig. 146. Axial interceiluiar canais in short tangential lines (feature 128), Dipterocarpus grandiflorus, x 20. -- Fig. 147. Axial canais diffuse (feature 129), Vareriamacrocarpa, x 29.

1
304 IAWA J3ul]tin n. s., Vol. 10 (3), 1989 IAWA List of rnicroscopic feawres for hardwoud identification
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II

Itf

1
l

v
149 ,. ol
I ' ^1,^ 1

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i

151 ,

Figs. 148, 149. Radial canais (feature 130). 148: Bursera gurnmifera, pefo1ia (arrow), x 140.

70: - 149: Shorea h-

Fig. 150. Radial canais (feature 130), Parahorea smythiesu. x 75. - Fig. 151. Jnterce11t11r canais of u-aurnatic origin (feanire 131), Murraya e.xotica, x 115-

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TUBES / TUBULES 132. Laticifers or lanninierous tubcs

Dejlnitio:
Tubes/tubules = ce].]s or series of cclls of indeterniinate lcngth, extending radially rir veriically (among fibres); according to specific contents mo Lypes can be distinguished. Laticifers = tubes containing latex, the latex may be colourless or light yellow to hrown; laticiers rnay extend either radially (in geriera of Apocynaceae, Asclepiadaceae, Campanulaceae, Caricaceae, Euphorbiaccae, Moraceae) or axial]y (interspersed aioong fibres and so Lar known ori]y from Moraceae), figs. 152-155. Synonyms: latex tubes, latex canaIs. Tanniniferous tubes = tubes containing tannins, whih are reddish-hrown, in rays (so Lar known only froni Myristicaceae), Figa. 156, 157.

Corn,nenrs:
Although latex is often light-coloured, and tannins are dark, colour is not a reliable diffcrence, and chenical tesis for tannin are needed to verify tube contents. Structural differences between the laticifcrs and tarininiferous tubes appear minor (Fujii 1988). Thereforc, thesc mo features are combiried into one descriptor. Latex traces are included in this clescriptor. Tanniniferous tubes oftcn are difficult to recognise in tangcntial sections because in that vim their dirnensions may appear similar to the ray edis; examining radial sections shows tanniniferous tubes to be longer than ray celis. 15

It;

Figs. 152-157. Laticifers (latex tubes) or tarininiferous tubes (feature 132). - 152: Alsionia scholaris, laticifer, x 140. - 153: Dyera costulata, wide laticiers, x 180. - 154: Crotorr panarnensis, laticifer with dark-staining contents, 290. - 155: Artocarpus conjnwjis, axial laticifcr. x 75. - 156 & 157: Horsfieldia subg/ohosa, tanniniferous tube in transverse and radial seclion (arrows), x 290.

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CAMBIAL VARIANTS 133 lncluded phloem, concentric 134. Included phloem, diffuse 135. Other cambial variants

Definrions:
tncluded phloem, coricentric = phloem sirands in tangential bands alternating with zones a xylem aiicl/or conjunctive tissue, fig. 158, e.g., Avicennia spp (Avic.enniaceac, Suaeda monoica (Chenopodiaceae). Incluided phloern, dilTuse = scattered, isolated phloeni strands. The phloem strands mav be surrounded by parenchyma ar imperforate tiacheary elements, fig. 159, e, vomica (Loganiaceae). Synonym: included phloem, foraminate ar island type. Other cambial variants = caegory for a variety of cambial variants inclj:rr cal, lattened, and furrowed in cross section; axes with lobed ar furrowed xy1cn; isircc x:. 1cm; compound, divided, corded and cleft xylem masses.

Comrnents:
The features for includcd phloem type are based on the appearance a [he wood, and do not have devclopmental inferences - they are not defined on the basis of whether there is a sing]e permanent ca'mbium, ar successive cambia, ar whethcr the tissue surrounding the phloeni swands is xylern or conjunetive tissue. As pointcd oul by Mikesell and Popham (1976) and Carlquist (1988), it is desirable to restrici ihe term interxylary ph]oem to chose cases in which the phloem has been produced internal]y by a single cambiurn. lncluded phioem of the concentric type very often intergrades with diffuse included ph]oem (e.g,, in many Chenopodiaceae); in all cases ofdoubt use both fcatures in species with concentric included phloem the phloein bands may branch and anastomose, and the conjunctive parcochyma somedrnes fornis radial extensions resembling rays. Because included phloem and other cambial variants are of regular occurrence iii the taxa in which they are founcl, lhe cerrn 'anornalous' niust be considered a misnoiner. Feature 135 'other cambial variants' most frcqucntly occurs in iiana: for more inforination sie Carlquist (1988),

Fig. 158. lncluded phloern, concentric (feattire 133, arrows), Suaeda ,nonoica. Note also radial extensions of conjunclive parenchyrna (feature 117, wood rayless, variable-): x 38. Fig. 159. lncludcd phloern, diffuse (feature 134, arrows), Strychnos nux-votnica, x 45.

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MINERAL INCLUSIONS PRISMATIC CRYSTALS 136. Prismatic crystals present 137. Prismatic crystals in upright and/or square ray cells 138. Prismatic crystals in produmbent ray celis 139. Prismatie erystals in radial alignment in procumbent ray cells 140. Prisrnatic crystals in chambered upright and/or square ray celis 141. Prismalie crystals in nonchambered axial parenchyrna celis 142. Prismatic crystals in charnbered axial parenchyma cells 143. Prismatic crystals in fibres Definirions: Prisrnatic crystals = solitary rhombohedral or octahedral crystals coniposed of calcium oxalate, which are birefringent under polarised light (fig. 160). Synonym: rhornboidal cxystal. Chambered ceil = an axial parenchyma strand ceil or ray parenchyma ccli subdivided by septa or b' thin to tliick celi walls. Features 137-143 as per descriptors, examples foilow. Prismatic crystals in upright and/or square ray cells, fig. 161, e.g.. Astronium spp. (Anacardiaceae), Bursera spp. (Burseraceae), Khaya anthorheca spp. (Meliaceae), 1-lelicostylis spp. (Moraceae). Prismatic crystals in procumbent ray cells, fig. 162, e.g., Anogeissus laufolia (Combretaceae), Carpinus spp. (Corylaceae). Prisrnatic crystats in radial aligument in procumhent ray celis, fig. 162, e. g., Aspidosperma quebracho-bianco (Apocynaceae), A nogeissus bufo lia, Bucida buceras (Combretaceae), Secw-i nega perrieri (Euphorbiaceae), Ouratea su.rinarnensis (Ochnaceae), Gonystylus spp. (Thymclaeaceac). Prismatie crystals in chambered upright andfor square ray edIs, fig. 163, e.g.. Elaeocarpus cabomala (Elaeocarpaceae), Glycydena'ron a,nazonicum (Euphorbiaceae), Banara nitida (Flacourtiaceae), Byrsonima laevigara (Malpighiaceae), Fagara fiava (Rutaeae).

lf,,

1'

:.1FJ\

Fig. 160. Prismatic crystals (feature 136), Drypetes keyensis, x 230. Fig. 161. Prismatic crystals in upright and/or square ray edIs (feature 137), Astronium graveolens, x 115. Fig. 162. Prismatic crystals in procumbent ray celis (feature 138) and in radial alignmcnt (fe.ature 139), Anogeissus laufolia, x 75. - Fig. 163. Prismatic crystals in chambered (or divided) upright and/or square ray cells (fearure 140), Elaeocarpus cabornala, x 290. - Fig. 164. Prsrnatic crystals in nonchambered axial parenchyma edis (feature 141), Drypetes gerrardii, x 290. Figs. 165-167. Prismatic crystals in chambered axial parenchyma celis (feature 142). - 165: Crysrals in short chains, Lithocarpus edulis, x 290. - 166 & 167. Crystals in long chains. 166: Tangential section, Parkiapenduln,x 115, 167: Radial section, tvfalpighia incana, x 115. F. 1 6. ['nsmatic crvstals in flhrcs (feature 14) Ranara reja. .' 115.

I1

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IAWA List of microscopic features for hardwood identification DRUSES 144. Druses present 145. Druses in ray parenchyma cells 146. Druses in axial parenchyma celis 147. Druses in fibres 148. Druses in chambered cells Dejlnirions:

313

Prismatic crystals in nonchambered axial parenchyma cells, fig. 164, e.g., Ceiba spp., Ocliroma spp. (Bombacaceae), Drypetes gerrardii (Euphorbiaceae), Ficu..s spp. (Moraceae). Prismatic crystals in charnbered axial parenchyma celis, figs. 165-167, e.g., Gilhertiodendron preusii (Caesalpniaceae), Pentacrne contorta (Dipterocarpaceae), Lilhocarpus edulis (Fagaceae), Juglans nigra (Juglandaceae), Couratari spp. (Lecythidaceae), Malpighia incana (Malpighiaceae), Parkia pendula (Mimosaceae), Zanthoxylum hygrophila (Rutaceae), Manilkizra spp. (Sapotaceae). Prismatic crystals in fibres, fig. 168, e.g., Heniandradenia chevalieri (Connaraceae), Banara regia (Flacourtiaceae), Triplaris americana (Polygonaceae), Punica granatum (Punicaceae), Majidea zanguebarica (Sapindaceae). Co,nments: Prismatic (rhomboidal) crystals are the most common type of ciystal in wood (Chattaway 1955, 1956). The relative abundance of prismatie crystals is variabie. In some species, crystals are consistently abundam; in others, they are consistently prcsent, but not abundant; and in still other species, they are present in some samples, but absent in other samples. In some taxa, crystais occur in only one ceil type; in other taxa, they occur in more than orle ccli type. In the iatter case, record ali features that apply. Information on the specific location of crystals often is not available from the literature. Consequentiy, fature 136, 'prismatic crystals present' is useful in constructing a database from existing information. Feature 136 is recorded in combination with other applicable crystal features (137-143). In some species, crystals occur throughout Lhe ray (for heteroceliular rays: features 137 and 138 both present). In other species with heteroceliular rays they are restricted to the marginal rows of upright and/or square celis, upright and/or square cells in the body of the ray, or sheath celis (feature 137 present, feature 138 absent). In yet others, crystals are restricted to thc procumbent body ray celis (feature 137 absent, feature 138 present). This last condition is not common and usually occurs in combination with crystals in a radial alignment (feature 139). The latter feature may appiy tu ciystals m nonchambered as weli as chambered celis. l'he descriptors 'prismatic crystais in chambered upright and/or square ray edis' (feature 140) and 'prismatic crystais in chambered axial parenchyma celis' (feature 142) include a considerabie diversity in types of chambcred or subdivided celis (cf. Parameswaran & Richter 1984) and, particularly for axial parenchyma, in length of the chains of crystalliferous chambers or subdivisions. In some taxa there are oniy a few charnbers in a series, in others there are long chains. Such inforrnation should be recorded in a description. Cauzions: There are many genera in which prismatic crystals are regularly absent (e. g,, Dipterocai-pus spp. - Dipterocarpaceae, Betula spp. - Betuiaceae, Liriodendron Spp. - Magnoliaceae, and Ti/ia spp. - Tiiaceae). But, when identifying an unknown, using absence of crystals is no[ recommended because crystals are of sporadic occurrence in many other taxa (e. g., Acer spp. Aceraceae, Quercus spp. - Fagaceae, and Ulmus - Ulrnaceae). Care is needed to distinguish between crystals in septate fibres and crystals in charribered axial parenchyma celis.

Druse = a compound crystal, more or less spherical in shape, in which the many component crystals protrude from Lhe surface giving the whoie structure a star-shaped appearance, figs. 169-17 1, e. g., Hibiscus tiliaceus (Malvaceae). Syrionym: cluster crystal. Features 145-148 as per feature descnptor, exampies foliow. Druses in ray parenchyma edIs, fig. 169, e.g., Gledirsia triacanthos (Caesalpiniaceae), Macaranga barreri, M. heudelotrii Euphorbiaceae), Colubrinaferruginea Rhamnaceae),

Arnygdalus cammunis (Rosaceae), Celtis paniculata (Ulmaceae). Druses in axial parenchyma cells, fig. 170, e.g., Dacryodes edulis (Biirseraceac), Terminalia catappa (Combretaceae). Druses in flbres, e. g., Combretum fruticosum (Combretaceae). Druses in chnmhered cells, fig. 171, e.g., Macaranga barteri, M. occidenralis (Euphorbiaceae), Banara regia (Flacourtiaceae). Comrnenrs: Most of the existing rnultiple entry ke ys and the literature do not provide information on the location of druses. Consequently, the feature 'druses preseni' is included so that this information can be used. In a given species, druses may occur in one or more of Lhe ccli types, and the celis may be cnlarged as well (feature 156).

OTHER CRYSTAL TYPES 149. 150. 151. 152. 153. Raphides Acicular crystals Styloids and/or elongate crystals Crystals of other shapes (mostiv small) Crystal sand

DeJinition.',.Raphides = a bundie of long needle-likc erystais, fig. 172, e. g .. Dilienia rericulara. Terra cera boliviana (Diileniaceae), Pisonia Spp. (Nyctaginaceae), Psyc/lot ria recordiana (Ruhiaccac), Tetrarnerista crassfolia (Tcuamcristaccac). Viris vinife ra (Vitaccae).

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Acicular crystals = small needle-like crystals, not occurnng in bundles, fig. 173, e.g., Tecorna stans (Bignoniaceae), Cryprocarya glaucescens (Lauraceae), and Gmelina arborea (Verbenaceae). Styloids = large erystais at least four times as long as broad with pointed or square ends,
fig. 174, e. g., Mayrenus obrusifolia (Celastraceae), Terminalia amazonica (Combretaceae), Gel-

semium senpervirens (Loganiaceae), Me,necylon membranfilium (Melastomataceae), Ga/lesia inregrfo1ia (Phytolaccaceae), Gonystylus bancanus (Thymelaeaceae). Elongate crystals = crystals two to four times as long as broad with pointed ends, fie. 175, e g., Siphonodon pendulwn (Celastraceae), Ligustrum vu/gare (Oleaceae), Vires glabrat (Verbenaceae). Crystals of other shapes (mostly smaH) = includes ali other shapes of crystals, figs. 176, 177, e. g., cubic (e. g,, Aporusa viliosa - Euphorbiaceae), navicular (boat.. shaped) (e. g., Litsea rericulara - Lauraceae), spindie-shaped (e. g., Dehaasia spp. - Lauraceae), pyramidal (e. g., Caryodaphnopsis ronkinensis - Lauraceae), tabular (e. g., Aniba spp. - Lauraceae), indented (e. g., Forestiera segregaza Oleaceae), twinncd (e. g., Nesregis spp. Oleaceae), etc. i .- ''... Crystai sarid = a granular mass cornposed of very small crystals, fig. 178, e. g., Cordia suhcordata (B oraginaceae), Acrinodaphne hookeri (Lauraceae), Bumelia obtusifolia (Sapotaceae), and Nico riana cordzfolia (Solanaceae). Synonym: microcrystais. Com,nents: Crystals, particularly the small ones, are best detected with polarised light These crystals are not comrnon, and their occurrence may be sporadic. Therefore, these features should only be used in the positive sense. Raphides and styloids often occur in enlarged celis, feature 156. For more information on crystal types. see Chattaway (1955, 1956) and Richter (1980).

1 73

f
lik
174

L: -

IL'

Caurions: Care must be taken not to interpret a cross section of an elongatc or acicular crystal as a cuhic crystal. lhe crystals in raphide hundies often scparate during sectioning.

OTHER DIAGNOSTIC CRYSTAL FEATURES 154. 155. 156. 157. 158. More than one crystal of about Lhe sarne size per celi or chamber Two distinct sizes of crystals per ccli or chamber Crystals in enlarged celis Crystais in tyloses Cystoliths

Fig. 169. Druses (feature 144) in ray parenchyrna celis (feature 145), Hibiscus riliaceus, x 145. - Fig 170. Druses in axial parenchyma cells (features 144 and 146), Terminalia carappa, x 170. - Fig 171. Druses in chambered ray celis (features 144, 145 and 148), Banara regia, x 290. - Fig. 172. Raphides (feature 149) in procumbent ray ceil, Vitis vinifera, x 290. Fig. 173. Acicular or needle-shaped crystals (feature 150), Gmelina arborea, x 675. - Fig. 174. Styloids (feature 151), Mernecylon memhrarnifolium (in included phloem), x 95. - Fig. 175. Elongate crystals (feature 151), Ligustrum vulgare, x 220. - Fig. 176. SmaiI cubic crystais (feature 152), Aporu.sa villosa (in ray celis), x 290. - Fig. 177. SniaII navicular crystals (f':iture 152), T,itsca reticulata. x 675.

Definir jons: Features 154-157 as per feature descriptor, examples follow. More than one crystal
of

about Lhe sarne size per celi or chamber, figs. 179,

182, e. g., Bouea opposirifolia (Anacardiaceae), Garcinia latissima (Guttiferae), Aniba dzsckeii

(Lauraceae). Ligustrum vulgare (Oleaceae), Gmelina arhorea. Vires divaricara (Verbenaceae).

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IAWA List of_microscopie features for hardwood identification

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Two distinct sizes of crystals per ceil or chamber, fig. 181, e. g_ Mangifera altissima (Anacardiaceae), Cordia bantamensis (Boraginaceae), Pentacme conrorta (Dipterocarpaceae), Zanthoxylum juniperinum (Rutaceae), Bumelia glomerara (Sapotaceae), Gonyszylu.s bancanus (Thymeiaeaceae). Crystals in enlarged celis (idiobiasts), fig. 183, e.g., Carpinus carolinianum (Corylaceae), Juglans nigra (Juglandaceae), Pyrts communis (Rosaceae), Citru.s auranrium (Rutaceae), Carne/lia japonica (Theaceae), Zelkova serrata (Ulmaceae). Crystals in tyloses, fig. 184, e. g., Astronium graveolens (Anacardiceae), Cordia gharaf (Boraginaceae), Pera bumeliaefolia (Euphorbiaceae), Chlorophora tincroria, Pseudolmedia spu'ria (Moraceae), Chrysophyllum aurarum (Sapotaceae). Cystoliths = internal stalked outgrowths of the ccli wali that project into the ccii lumen and are composed of celiulose impregnated with calcium carbonate. They are irregular in shape and sometirnes cornpletely 611 a ccli, fig. 185, e. g., in some Trichanrhera spp. (Acanthaceae), Spararra,uhelium spp. (Hernandiacae), and Opiiiaceae. Comments: Generaily, there is ordy one crystal per ccli or charnber. However, two or more similar-sized crystals, especially acicuiar and/or navicular, and cubic and/or rectangular crystals, may Occur in the sarne ccli or chamber. li is rare that there are two distinct sizes of crystals in the sarne ccli or chamber. For more information, see Chartaway (1956). For feature 156, 'crystals in enlarged edis (idiohlasts)', the enlargcd edis can be either ray or axial parenchyrna edis, or more rarely both. The crystals in enlarged edis may he prismatic crystals, druses, raphides, or any other crystal type. Cystoliths, as far as is known, occur only in the exampies given (Ter Weile 1980). Caution: Raphides are bundies of crystals, but the whole bundie is considered as a singie unit, and so feature 154, 'more than one crysta.l per ccli or chamber', does not apply to raphides. Crystal sand should also not be coded under feature 154.

I
1

i. 1
1

Fig. 178. Crystal sand (feature 153, arrowheads), Cordia suhcordata (in ray celis), x 290. Figs. 179 & 180. More than one erystal of about the sarne size per ccli or charnber (feature 154). - 179: In axial parenchyrna celis, Garcinia lati ssima, x 75. - 180: In ray edis, Bouea opposiiifolia, x 290. Fig. 181. Two distinct sizes of crystals per cdl or chamber (feature 155), Cordia banramensis (in ray celis), x 290. - Fig. 182. Different size classes of crystals intergrading in some ray edis (feature 155 variable), but more than one crystai of about the sarne size per ccii in others (feature 154 present), Cordia abyssinica, x 115. - Fig. 183. Crystais in enlarged celis (featm-e 156), Citru.s aurantium, x 135. - Fig. 184. Crystals in tyloses (feature 157), Cordia gharaf, x 140. - Fig. 185. Cystoliths (feature 158), Trichanthera gigantea, x 220.

318 SILICA

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159. Silica bodies present 160. Silica bodies iii ray celis 161. Sulca bodies in axial parenchynia celts 162. Silica bodies in fibres 163. Vitreous silica Defitjtjons: Silica bodies = spheroidal or irregu]arly shaped particles composed of silicon dioxide. Synonyms: sulca grains, silica inciusions. Features 160 -162 as per feature descriptor, exampies follow. Silica bodies in ray celis, fig. 186, e.g., in Tratrfnickia burserfolla, T. demararae (Burseraceae), Licania leptostachya (Chrysobaianaceae), Shorea lameilara (Dipterocarpaceae), Mezilaurus irauba (Lauraceae), Vitex compressa (Verbcnaceae). Silica bodies in axial parenchyma cells,fig. 187, e.g., inBomhax nervosum, Distemonanjhus spp. (Bomhacaceae), Apuleia leiocarpa, D ialium guianense (Caesalpiniaceae). Silica bodies in fibres, fig. 188, e.g., Canariwn hirsutum, Prorium neglectum, Trartinickia burserfaIia (Burseraceae), Ocotea pube ruia (Lauraceae). Vitreous silica = silica that coats ccli walis or cornpletely fihis the ccli lurnina, fig. 189, e.g., Srereospermum chelonoides (Bignoniaceae), Hydnocarpus gracis (Flacourtiaceae), Artocarpus vriesianus (Moraceae), and Gynot-roches a.rlllw-is (Rhizophoraceae). Procedures: Slica hodies: Silica bodies are observed with Lhe light microscope in radial sectioris of eithcr permanent or temporary mounts or in celis that have been macerated, LI largo amounts of extractives are present and the sulca bodies are difficult to see in section, hleach with a domcstic blcaching agent, rinse in water, heat in carbolic acid, and mount iri clove ou, or macerate a few chips in any macerating fluid that removes most of the exlracives and lignin but not the silica, At low magnification (4-10 x objective lens), si]ica bodies genera]ly appear as smaIl dark nonbirefringcnt particles. At higher magnification (25-40 x objective lens), they have a 'giassy' appearance. Vitreou.s slica.' Thoroughly macerate chips or splinrers, leave the wood iii the maceratirig soludon until ii is white. Decant the macerating tluid, add water, rinse, decant, and repeat mui] Lhe solution is clear. Place some macerated wood on a slide: warrn ilie slide on a hotplate until the macerated wood is dry. Aliow the slide to cool, and then add 2 to 3 drops of concentrated su]furic acid to dissolve ihe ceilulose. Add a cover s]ip and observe the cel]s under a light microscope ai low magnification. Vitreous silica appears like pieces of translucent vessel clements and fibres. To distinguish undissolved celis from vitreous silica use polarised light. Undisso]ved edIs are bircfringent, whereas vitreous silica is not. Vitreous sulca can also be recognised in weil-bleached sections bccause of irs 'glassy' appearance.

Fig. 186. Silica bodies present (feature 159) in ray edis (feature 160), Shorea lamellata, x 150. - Fig. 187. Sulca bodies in marginal ray celis and axial parenchyma edIs (features 159, 160, and 161), Apuleia lelocarpa, x 75. - Fig. 188. Silica bodies iii fibres (features 159 and 162), Ocotea ci. puberula. x 115. - Fig. 189. Viu-enus silica (feature 163), Hvdnocarpus gracilis, x 5(}.

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IAWA List of microscopic features for hardwood identification APPENDIX Non-anntomical Information GEOGRAPH1CAL DISTRIBUTION (fig. 190)

321

Cornmenss:

Silica bodies rnost often are rcstrictcd to ray celis, particularly the marginal or upright edis. Sometimes they are rcsicted to axial parerichyma; somctimes thcy oceur in both ray and axial parenchyma. Slica bodies rarely occur in fibres, but if lhey do lhe fibres usually are septate. Wheiher silica occurs in aggregations, as irregularly shaped or globular bodies, or whether the silica bodies have a smooth or verrucose surface may be diagnostic in certain groups and needs to be re.corded in a description. For more information on silica bodies, see Amos (1952), Ter Weile (1976), and Koeppen (1980).
Cautions: When looking for silica bodies, do not use glycerin as a mounting medium because

its refractive index niakes it difficult to detect silica bodies. Hydrofluoric acid, which is sometirries used to soften wood, wili dissolve the silica bodies.

---
1B2<

164

164. Europe and temperale Asia (Brazier and Franklin region 74) 165. Europe, excluding Mediterranean 166. Mediterranean including Northern Africa and Middle East 167. Temperate Asia (China), Japan, USSR 168.. Ceniral South Asia (Brazier and Franidin rcgion 75) 169. India, Pakistan, Sri Lanka 170. Burma 171. Southeast Asia and the Pacific (Brazier and Franklin region 76) 172. Thailand, Laos, Vietnam, Cambodia (Indochina) 173, Indomalesia: Indonesia, Philippines, Malaysia, Brunei, Singapore, Papua New Guinca, and Solomon Islands 174. Pacific Islands (including New Caledonia, Samoa, Hawaii, and Fiji) 175. Australia and New Zealand (Brazier and Franktin region 77) 176. Australia 177. New Zealarid 178. Tropical mainland Africa and adjacent islands (Brazier and Franklin regiori 78) 179. Tropical Africa 180. Madagascar & Mauritius, Runion & Comores 181. Southern Africa (south of the Tropic of Capricorn) (Ilrazier and Franklin rcgion 79) 182, North America, nonh of Mexico (Brazier and Frarddin rcgion 80) 183. Neotropies and temperate Brazil (Brazier and Franklin region 81) 184. Mexico and Central America 185 Carrihbean 186. Tropical South America 187. Southern Ilrazi] 188. Temperate South America including Argentina, Chile, Uruguay, and S. Paraguay (Brazier and Franklin region 82) C,nments.' There is no single ideal way of dividing the world. Thc abovc is a mixture of political and hiogeographica] criteria. It retains the major geographical regions of Brazie.r and Franklin (1961), but some regions are subdivided so that more prccision is possible.

IIs:fZ
Fig. 190, Geographical distribution (feaures 164-188).

HABIT 189. Tree 190. Shruh 191. Vine/liana Definjtion.: Tree = woody perennial piam with one main stem usuatly over 3 Tnetres tal],

R
322 IAWA Builetin n.s., Vol. 10 (3), 1989 IAWA List of microscopic features for hardwood identification 323 Shrub = a woody pererinial plant usual]y with several stems and usually Iess than 3 meti-es uilI at maturity. Vine/liana = any plant witli a long relativel y thin siem that c!imhs along a suppnrt or trails along lhe ground. Comments: There wifl be overlap between these categorics in some species, and two or more, rarely ali three,may apply.

which specific gravity is detemiined. Basic specific gravity (Bsg), which is based on the green volume (wood fully swollen, moisture coritent of fibre saturation point or higher) is one of lhe most commonly cited values (Panshin & DeZeeuw 1980). Qffier values often given for wood include basic density (Bd) which is equal to lhe oven-diy weight of woodfgreen volume and which has units (g/cm 3 , kg1m 3 ' or lbs/ft3 ). To convert from basie specific gravity (Bsg) to basie density (Bd) rnultiply lhe basic s.g, by lhe density of water as is shown below: Bd in gfcni 3 = Bsg x 1 glcm3 Bd in kg /M3 = Bsg x 1000 kg/m3 Bd in Ibs/ft 3 = Bsg ' 62.4 1b51ft3 Therefore Bsg of 0.4 = Bd of 0.4 g/crn 3. 400 kg/m 3 , or 25 lbs/ft3 Bsg of 0.75 = Bd of 0.75 g/cm 3 , 75 kg1m3 , or 46.8 1b5/ft3

WOOD OF COMMERCJAL HPORTANCE 192. Wood of commercial importancc This category is intended for woods of both historical and current commercial importance. The term ofcommereial importance' is somewhat vague, and should be used with caution when identifying an unknown. But when identifying certain wooden artefacts, C. g., fiirniture. ii can be helpful to segregate comrnercial species from noncommercial species.

HEARTWOOD COLOUR
196. Fleartwood colour darker than sapwood colour 197. Heartwood basically brown or shades af brown 198. I-teartwood basically red or shades of red 199. Ileartwood basically yellow or shades of yellow 200. Heartwood basically white to grey 201. Fleartwood with streaks 202. Heartwood not as above

SPECIFIC ORA vrry 193. Basic specific gravity 10w, ^ 0.40 194. Basic specific gravity medium, 0,40-0.75 195. Basic speeilie gravity high, ^ 0.75 Defini:ion.s: Basic specifie gravity = ratio of lhe oven-dry weight of a piece of wood to the weight of lhe water displaced by lhe wood when it is completely swollcn (i.e., green volume). Examples for lhe feature categories follow. Basic speciic gravity Iow, :!^ 0.40, e. g., Dyera co.rtulata (Apocynaceae), Ceiba spp., Qchrwn spp. (Bombacaceae), Populus balsamfera (Salicaceae). Triplochton spp. (Stercuiiaceae), Ti/ia spp. (Tiliaceae). Basie speciflc gravity medium, 0.40-0.75, e.g., Acer saccharum (Accraceae). Betuia lenta (Benhlaceac), Carapa guianensis, Khava rardjf!iiia (MeiLeae). iruns aniercana (Oleaceae), Tectona spp. (Verbenaceae). Basic speciic gravity high. ^ 0.75, e. g ..As rrwuum 2reveo1ens (Anacard iac) Ocotea rodiei (Lauraeeae), Dalbergia melancwlon (Papilionaceae), Manikarn bidenrata (Sap) taceae), Guaiacwn spp. (Zygophyl]aceae). Com rnenrs: Density is the weight of a substance (mass) per uni[ volume: speciflc gravity (s.g.) is lhe rario of the density of a material to lhe density of water and, conscquently, specific gravity does not have units. For purposes of compucing speci fie gruvily of wood, wood density uses lhe oven-drv weight of wood as lhe numerator, Because lhe volume of wood changes with changes in moisture content bclow fibre saturation point, it is neccssarv lo specify the muislure conreni ai

Definitions:
Heartwood colour darker than sapwood colour, e.g,, Astronium spp. (Anaeardiaceae), Tabebuia guayacan (Bignoniaceae), Acocia koa (M.imosaceae), Morus alba (Moraceae), Robinia spp. (Papilionaceae).

Heartwood basically brown or shades of brown, e,g., Quercus alba (Fagaccae), Albizia (Samanea) saman (Mimosaceae), Morus rubro (Moraceae), Eucalyptus giobulus (Myrtaceae), Robinia spp. (Papilionaceae). Heartwood basically red or shades of red, e. g., Brosimum rubescens (Moraceae), Prerocarpus macrocarpus (Papilionaceae), Sickingia spp. (Rubiaceae). Ileartwood basically yellow or shades of yellow, e.g., Enantia chloranrha (Annorsaceae), Buxus spp. (Buxaceae), Schaefferiafrutescens (Celastraccae), Gossypiospernium spp. (Flacourtiaceae), Ciadrastis iutea (Papilionaceae). Heartwood basically white to grey, e.g., Jiex opaco (Aquifoliaceae), Didymopana.x spp. (Araliaceae), Ceiba spp. (Bombacaeeae), Hura spp. (Euphorbiaceae), Cecropia spp. (Moraceae). Simarouba spp. (Simaroubaceae). Heartwood with streaks, e. g., Dracontomelon dao (Anacardiaceae). Heartwood not as above = colours such as black, purple, orange, green, as in Diospyros ebenum (Ebenaceae), Peltogyne Spp. (Papilionaceae).

324 Procedure:

IAWA Bulietin n.s, Vol- 10 (3), 1989

IAWA List of mioscopic feanires for hardwood identi6cation -325 ODOUR* 203. Distinet odour Definition: Distinct odour = as par frature descriptor, e.g., Vihurnwn (Caprifoliaceae), Ceraropetalum apetalum (Cunoniaceae), many specics ofLauraceae, Cedrela (Meliaceae), and Santalwn (Santalaceae). Procedure: In dricd wood samples the chemicais responsible for Lhe odour rnay have voiatised froin lhe surface, so it will be nccessary to expose a fresh surface, or take other measures to enhance Lhe odour, e. g., add moisture by breathing on the wood, or wet Lhe wood with water and warm it.

Time and exposure to light may alter Lhe appearance or vividness of Lhe colour. Thercforc, it is best to determine colour from a freshly cul tangentia] surface of a dry woocl sample. The heartwood colour of recently felled trees often differs from that of dry wood samples. These descriptors are for wood samples that are ai Ieast air-dry. Comments.' Feature 196, 'heartwood colour darkcr than sapwood colour', only can be used wben both heartwood and sapwood are present, and is recorded in combination with Lhe other heartwood colour features (197-202). The heartwood of some species, e.g., Quercus alba (Fagaceae) and Betula lenta (Betulaceae), is not markcdly darker than Lhe sapwood, but can be distinguished from it, so feature 196 applies to thcse taxa. The variety of colours, shades, and combinations of heartwood colour make it impossible to categorise ali of theni. In general, Lhe colour of heartwood is either brown, red, yellow, white, or some shade or combination of these colours. Basically brown heartwood is very corumon; basically red and basicafly yellow are rather rale; basicaily white or grey is rather frequent. The heartwood colour of many taxa is not rcstricted to one colour, bui tu a combination of colours and, when appropnate, various combinations shou]d be recorded and may be used when idendfying an unknown. Examples of these combinations include: reddish-brown in Astronium spp. (Anacardiaceae), Hyrnenaea spp. (Caesalpiniaceae), Quercus rubra, Fagus spp. (Fagaceae), Khaya spp., Swieienia spp. (Meliaceae); yellow and brown in Disternonanthus spp. (Caesalpiniaceae), Chlorophora :inctoria (Moraceae), Adina cord(folta (Rubiaceae), Faga.ra spp. (I.utaceae), tvfastichodendron spp. (Sapotaceae). Very light coloured woods would be recorded as combinations of white to grey and brown and/or yellow, c.g., Acer spp. (Aceraceae), Alstnia spp. (Apocynaceae), Anisoptera spp. (Dipterocarpaceae), Gmeilna spp. (Verbenaceae). Heartwood with streaks is always uscd in combination wth Lhe general heartwood colour, as in Microberlinia spp. which lias brown heartwood with streaks. 'Heartwood not as above' is a 'catch-all' category for taxa with heartwood colours such as black, green, orange, and purpie. This feature may be used alone (e. g., as for Diospyros ebenwn - Ebenaceae, which has distinctly black heartwood), but more commonly it will be used in combination with olher heartwood colours. For example, Lhe combinadon of basically brown and green occurs in Bucida buceras (Combretaceac), Ocoteu rodiei (Lauraceae), Liriodendron rulip (fera. Michelia spp., Talawna spp. (Magnoliaceae); Lhe combination of basically red, brown, yellow, and orange with streaks occurs in Centrolobiu,n spp, (Papilionaceae) and Aspidosperma spp. (Apocynaceae); lhe combination of brown, red, purple, black and orange with streaks occurs in Dalbergia spp. (Papilionaceae). Cautions: Do not use heartwood colour features for ancient saniples, archaeological material, or other samples whose colour has been altered by burial, time, lreatrnent, or decay. Be particularly careful when using lhe feature 'Heartwood basically white io grey', because a whitish coloured sample may represent sapwood and not heartwood.

Caucion: Odour is quite variable, and individual perceptions of odour often differ 'I'hercforc,
use this feature with caution and only ia Lhe positive sente. * Taste is deibcrately excluded because of safety considerations, particularly a concern that someone may try tasting a wood whose contents could cause a severe allergic reaclion.

HEARTWOOD FLUORESCENCE 204. Heartwood Iluorescent Definition: Heartwood fluorescent = heartwood fluorescing when ilhiminated with longwave ultraviolei light, e.g., with a strong yellowish or greenish fluorescence in Anacardium excelsum (Anacardiaceae), Asimina spp. (Annonaceae), Aspidosperina eburneum (Apocynaceae), Robinia spp. (Papilionaceae); with a slight tinge of orange fluorescence ia Mangifera indica (Anacardiaccae), Varairen lundellii and Symphonia spp. (Guttiferae),Nauclea diderrichii (Rubiaceae); weak, yet positiva, fluorescence in many Annonaccae, Lauraceae, and Magnoliaceae. Procedure: Samples for testing fluorescence must be frcshly surfaced; simply removing some shavings with a knife is sufficient for exposing a fresh surface. Place samples i.mder a longwave (365 nm) ultraviolet (UV) iight ata distance of less than 10 cm. A high-intensity longwave UV lamp and observation in a darkened room is recommended. Co,nrnents: Fluorescent samples generally appear yellowish or greenish under Lhe UV iamp, although some species show slight tinges of orange or pink. Samples that are not fluorescent rnay rellect some of Lhe UV light making the sample appear slightly blue or purpie. Some samples with a yellowish heartwood, such as Chloroxylon spp, (Rutaceae) and Gonystylus spp. (Thymeiaeaceae), are not fluorescent, but may seem tu have a weak yellow fluorescence because of reflection. Absence of fluorescence can be irnportant in some famulies, e. g., Anacardiaceae and Leguminosae. See Aveila et ai, (1989) for a survey of fluorescence ia Lhe dicotyledons.

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Caulions: This feature applies only to naturaily occurrmg fluorescence and not to fluorescence associated with decay or pathological infections. Wood infected with decay organisms may fluoresce with sireaks, spots, or a mouled appearance, e.g., wetwood of Populus rre,nuloides ( Salicaceae) produces yellow fluoresceni streaks. Naturaily occurring fluorescence appears more uni form. Oven-drying of samples or exposure to high temperatures or other extreme environmental condirions may affect fluorescence propeilies.

Examples of wood with weaker fluorescence of ethanol extracts, but still positive, include Kiggelaria africana (Flacourtiaceae), Acacia melanoxylon ( Mimosaceae), Olea capensis (Oleaceae). Sometimes the water extract of a species fluoresces, bus its ethanol extract does not (e. g., Leucaena glauca - Mimosaceae). More often, the ethanol extract fluoresces, whi!e the water extract does not (e. g., Afzelia quanzensis Caesalpiniaceae, Lysiloma baha,nensisMimosaceae). Commenrs & Examples for water & alcohol exrract colour: Water extract basically colourless tu brown or shades of brown is the most common of the water extract feature colours. Examples of woods with water extract basically red or shades of red (feature 206) include Brasilettia spp. (Caesalpiniaceae), Catha edulis (Celastraceae), Cunonia capensis (Cunoniaceae), and Mimusops caffra (Sapotaceae). Examples of woods with water extract basically yellow or shades of ye!low (feature 2( y7) include Gonioma ka,na.ssi (Apocynaceae), Albina adianthifolia, Acacia caffra (Mimosaceae). Feature 209, 'water extract not as above' includes colours such as orange, hlack, and purpie. Various combinations of water extract colours occur as well. Ethanol cxtract basically colourless to brown or shades ofbrown (feature 211) is the most common. Examples of woods with 'ethanol extract basicaily red or shades of red' (feature 212) include Rhus integrifolia ( Anacardiaceae), l3aikiaea plurijuga, Peltophorum dubium, Swartzia madagascariensis (Caesalpiniaceae), and Berchemia discolor (Rhamnaceae). Exarnples of woods with 'ethanol extract basically yellow or shades ofyellow' (feature 213) include Gonioma kamassi (Apocynaceac), Praeroxylon obliquwn, Zanrhoxylwnflavum ( Rutaceae), Balanites ,naugharnii (Zygophyllaceae). Feature 214, 'Ethanol extract not as above', includes co!ours such as orange, black, and purp!e. For more information, see Dyer(1988) and Quirk (1983).

WATER & ETHANOL EXTRACTS: FLUORESCENCE & COLOUR 205. 206. 207. 208. 209. 210. 211. 212. 213. 214. Water extract fluorescent Water extract basically colourless to brown or shades of brown Water extract basically red or shades of red Water extract basicalty yellow or shades of yellow Water extract not as above Ethanol extract tluorescent Ethanol extract basically colourless to brown ar shades of brown Ethanol extract basically red or shades of red Ethanol extract basically yellow or shades of yellow Ethanol extract not as above

Procedures: Preparation of extracts: Add enough thin heartwood shavings to cover the bottom of a clean vial which is approxirnately 20 mm x 70 mm. Do not use splinters or chips, because the extraction time is much longer than for shavings. For water extracts, cover the shavings to a depth of approximately 20 mm (approximately 5 ml) with distilled water that is buffered at a pH of 6.86. Packets of buffering agent are available from most scientific supply companies so that only the contents of a packet need to be added to 500 or 1000 ml ofdistilled water to obtain the desired pH. For ethanol extracts use 95% ethanol. Dererminingfluorescence a water & alcohol extracrs: Cover the vials and shake vigorously for 10 to 15 seconds. Allow the shavings and solution to stand for 1 to 2 minutes, and then hold the vial under a longwave (approximately 365 nm) UV lamp and check for extract fluorescence. Generaily, extracts that fluoresce are bluish, but sornetimes they are greenish. Determining colour of water and alcohol e.rrracts: Afrer determiriing the fluorescence of the water and ethanol extracts, place the viais on a hotplate and bring the solution to a boi!. As soon as the solution boils, remove the vial and immediately determine the colour. Com,nenrs & Examples for fluorescence: Examples of woods yielding water exrracts that fluoresce a hrilliant blue include Strychnos decussata (Loganiaceae), Brosimum rubescens ( Moraceae), Olea europaea subsp. africana (Oleaceae), Prerocarpus indicar ( Papiionaceae), Zanrhaxylwnfiavwn (Rutaceae). Examples of wood with weaker fluorescence of watercxtracts, but still positive, includeAcaciafarnesiana (Mimosaceae) and Lonc/iocarpus capassa (Papilionaceae). Examples of woods yielding ethanol extracts with bright fluorescence include Protorhus longifolia (A naca rdi aceae), Cordia gerascanrhus ( Boragi naceae), A cacia eriolol'a (Mimosaceae).

FROTH TEST 215. Froth test positive Procedure: FolIow the procedure for preparing for lhe water extract tests. Add enough thin heartwood shavings to cover the bottom of a clean via! approximately 20 mm x 70 mm. Do not use splinters or chips, because the extraction time is much longer than for shavings; if sawdust is used, extraction time will be less. For water extracts, cover the shavings to a depth of approximate!y 20 mm (approximately 5 ml) with distilled waler that is buffered at a p11 of 6.86. Packets of huffering agent are available from most scientific supply companies so that only the contents of the packet need to be added to 500 or 1000 ml of distilled water tu obtain the desired pH. Cover the vial and shake vigorously for 10 to 15 seconds. If natural saponins are present in large amounts, tiny bubb!es or 'froth' (like foam on a g!ass of beer) will be formed. A!low the vial to stand for approximate]y 1 minute from the end of the shaking. If 'froth' still completely covers the surface of the solution after 1 minute, the test is positive. If 'froth' or bubbles form and then disappear within 1 minute, the test is negative. Ifonly some 'froth' remains around the edge of lhe vial (i, e., forming a ring of 'froth'), but does not cover the entre surface, the test is weakly positive.

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IAWA List of microscopie features for hardwood identification Fuil or complete ash = ash more or less retaining the shape of the original splinter.

329

Com,nen:s & Exa,'nples: Positive 'froth' test reactions are produced by, e. g., Mora spp. (Caesalpiniaceae), Enrerolobium cyclocarpum, Lysiloma baharnensis, Pseudosamanea spp. (Mimosaceae), Dipholis spp., Mastichodendron spp. (Sapotaceae). Wcaldy positive reactions (ring of froth) are produced by, e. g., Pelrophorum spp. (Caesalpiniaceae), Kiggelaria spp. (Flacourtiaceae), Ekebergia spp., Enrandrophragma spp. (Meiaceae), Acacia nigrescens (Mimosaceae), Millerria spp. (Papilionaceae), and llerchemia spp. (Rhamnaceae). For more information, see Dyer (1988), Quirk (1983), and Cassens and Milier (1981).

Partial ash = ash that shrinks in size in comparison to the original splinter, lias a tendency to drift away, and usually feels gritty when rubbed between the fingers. Procedure: Prepare match-size (approximately 2 x 2 x 50 rum) splinters from sound outer heartwood, insure the wood is aI least air-dry. The splinter must be ignited with a match, and devices (e. g., Iighters) producing higher temperatures must be avoided. Ignite the splinter while it is held in a vertical position wiffi a pair of tweezers/forceps. While the splinter is burning, hold it in a hoij -zontalpsidurowly. Some timber species will hurn with relative case (e. g., Populus spp. - Saiicaceae), while others rnay show considerable reluctance (e. g., Eucalyptus paniculara - Myrtaceae). If it appears that the flame will extinguish before the splinter lias burned fully, combustion may be aided by gently returning the splinter to a vertical position and then back to horizontal. After Lhe flame extinguishes, it is important to aliow the giowing part of the splinter to extinguish before placing Lhe remnant on a cold surface. Comments: Certain splinters may crackle or produce bright sparks (e. g., Termina/ia carappa - Combretaceae), whilc others may produce a characteristic smoke coloration (heavy black smoke in Fimdersia iaevicarpa - Rutaceae) or exude coloured compounds while Lhey burn. Ali these features rnay be recorded ia a description. The descriptive classifications for appearance of the burnt splinter are those first recommendcd by Dadswell and Burneli (1932). In some iiterature, buff is used to describe spiinters that have Lhe colour of pale tanned leather, a yellowy brown, e. g., Eucaiyptus panicuiara (Myrtaceae). Apart from its use in CS1RO keys, Anonymus (1960) lias impiemented the feature and suggests that it is of little vaiue except in distinguishing between some timbers which are closely related anatornically. For further information on the burning splintcr test, which so far has only been used on a very limited scale, see Mann (1921), Welch (1922), Swain (1927), Dadswell and Burncli (1932), and Mennega (1948). Caution: Code greyish-white ash as 'other than above', as the white is reserved for obviously (bright) white ash.

CITROME AZUROL-S TEST 216. Chrome Azurol-S test positive Procedure: Prepare a 0.5% solution of chrome azurol-S reagent by dissolving 0.5 g of the dry chrome azurol-S granules and 5.0 g of sodium acetate (buffer) in 80 ml of distiiled water. After the chemicais are compietely clissolved, add enough distilled water to make 100 ml of reagent. This solution is stable and can be used over a number of years. To test dry wood samples, use ali eyedropper to appiy one or two drops of the solution to a freshly exposed end-grain. In highly positive woods, a bright blue colour wii develop in a matter of minutes, e.g., Poga spp. (Anisophylleaceae), Cardwellia spp. (Proteaceae), Symplocos spp. (Symplocaceae), ali Vochysiaceae. In those woods which absorb the solution very slowiy, e.g, Anisophyllea spp. (Anisophylleaceae), Goupia spp. (Goupiaceae) or contain low concentrations of a]uminum, e.g., Laplacea spp. (Theaceae), Henrieriea spp. (Melastomataceae), several hours niay be required for the blue colour to develop. Comment: Chrome Azurol-S tests for the presence of aluminum in both heartwood and sapwood. For more information on this test, see Kukachka and Miller (1980). Caution: Avoid decayed wood because chrome azurol-S is an indicator for some types of wood decaying fungi.

BIJRNTNG SPLINTER TEST 217. 218. 219. 220. 221. Splinter burns to charcoal Splinter burns to a fulI ash: Colour of ash bright white Splinter burns to a fuIl ash: Colour of ash yellow.brown Splinter burns to a fuil ash: Colour of ash other than above Splinter burns to a partial ash REFERENCES Amos, G.L. 1952. Silica in timbers. C.S.T.R.O. Australia, Buii. No. 267. Anonymus. 1960. identification of hardwoods. A lens key. For. Prod. Res. Buil. No. 25, London. Avelia, T., R. Dechamps & M. Bastin. 1989. Fluorescence study of 10,610 woody species from Lhe Tervuren (Tw) coilection, Belgium. IAWA Buil. n. s. 9: 346-352. Baas, P. 1973. The wood anatomical range in ilex and its phylogerietic and systematic significance. Blumea 21: 193-258. - 1986. Terminology of imperforate tracheary elements - in defence of libriform fibres with minutely bordered pits. IAWA Buil. n. s. 7:82-86.

Definitions: Charcoal = the blackened and charred remains of a splin ter, which usually burned slowly andfor with difficulty, or the black and charred remnant of Lhe splinter with a fine thread of black or grey ash which may remam attached.

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- , P.M. Esser, M.E.T. van der Westen & M. Zandee, 1988. Wood anatomy of Lhe Oleaceae. IAWA BulI. as. 9: 103-182. - & M. Gregory. 1985. A survey of oil celis in Lhe dicotyledons with cornments on their replacement by and joint occurrence with mucilage celis. Israel J. of Bot. 34: 167-186. - & F.H. Schweingruber. 1987, Ecological Irends ia lhe wood anatomy of crees, shrubs and clirnbers from Europe. 1AWA Buli. n.s. 8:245-274. Bailey, 1W, 1933. The canibium and its derivative tissues. VIII. Structure, distribution and diagnostic significance of vestured pits in dicotyledons. J. Arnold Arbor, 14: 259-273. - 1936. The problem of differentiation and c]assification of tracheids, 6ber-tracheids, and libriform fibers. Trop. Woods 45: 18-23. Braiier, J. & 01. Franklin, 1961. Identiflcatiort of hardwoods: a niicroscope key. For. Prod. Res. Buil. 46. HMSO, London. llridgwater, S. & P. Baas. 1982. Wood anatomy ofXanthophyllum Roxb. IAWA Buli. n.s. 3: 115-125. Carlquist, S. 1980. Further concepts in ecological wood anatotny, with corrmients on recent work in wood anatomy. Aliso 9: 499-553. - 1985. Vasicentric tracheids as a drought survival mechanism in Lhe woody flora of southern California and similar regions; review of vasicenlric tracheids. Aliso 11: 37-68. - 1986a. Terniinology of imperforate tracheary elements. 1 AWA Buli, n.s. 7: 75-81. - 1986b. Ibid.: A reply. IAWA Buil. as. 7:168-170, - 1988. Comparative wood anatorny. Systematic, ecological, and evolutionary aspecis of dicotyledon wood. Springer Verlag, Heidelberg, Berlin. Cassens, D. L. & R. B. Milier. 1981. Wood anatomy ofthe New World Pithccellobium (sensu lato). J. Arnold Arbor. 62: 1-44. Chattaway, M.M. 1932. Proposed standards for numerical values used ia describing woods. Trop. Woods, 29: 20-28. - 1955. Crystals ia woody tissues I. Trop. Woods 102; 55-74. - 1956. Crystals ia woody tissues II. Trop. Woods 104: 100-124. Clarke, S.H. 1938, A multiple-entry perforated . card key with special reference to Lhe identification of hardwoods. New Phytologist 37: 369-374. ronquist, A. 1981. An inteated system of classification of fiowering plants. Columbia Univ. Press, New York. - 1988. The evolution and classification of fiowering plants. 2nd Ed, New York Botariical Garden, Bronx, New York, Dadsweu, 1 I. & M. Burneli. 1932. Method for lhe identification of coloured woods of the genus Eucalyptus. CSIR Div. For. Prod., Technical Paper No. 5, Dodd, R. S. 1986. Fiber Iength nieasurcrncnt systems: A review and modification of an existing method. Wood & Fiber Sei. 18: 276-287, Dyer, S.T. 1988. Wood fluorescence of indigenous South African trees. IAWA Buli, as. 9: 75-87. Fahn, A., E. Werker & P. Baas. 1986. Wood anatomy and identfication a trees and slirubs from Israel and adjacent regions, Israel Acad. Sei. Hunianides, Jerusalem. Fujii, T. 1988. Structure of latex and tanniniferous tubes in tropical hardwoods. BUI]. For. Prod. Res. Inst. No. 352: 113-118 (+ 14 places; Japanese with English summary). Gouwald, H. 1983. Wood anatomical studies of I3oraginaceae (s.1.). 1. Cordioidae. IAWA Buli. n.s. 4: 161-178. Gregory, M. & P. Baas. 1989- A survey of mucilage celis ia vegetative organs of the Dicotyledons. Israel J. ofBot. 38: 125-174.

Hart, C.A. & B. Swindcl. 1967. Notes on laboratory sarnpling of rnacerated wood fibers. TAPPI 50 (7): 379 -3 8 1. Hillis, H.E. 1987. Heartwood and tree exudates. Springer Verlag, Heidelberg, Berlin, London, Paris. IAWA Committee, 1964. Multi]ingual glossary of tenns used ia wood anatoniy. Konkordia, Winterthur. IAWA Comrnittee. 1981. Standard Iist ofcharacters suitable for coinputerized hardwood identilication. IAWA Bull. n. s. 2:99-110. Ilic, Y. 1987. The CSIRO family card sorting key for hardwood identification. CSIRO Division of Chemical and Wood Technology Technical Paper No. 8. Kocppen, R. C. 1980. Silica bodies in wood of arboresccnt Leguminosae. IAWA Buil. n. s, 1: 180-184. Kribs, D.A. 1968. Commercial forcign woods on Lhe American Market. Dover Publications, New York. Kukachka. B.F, & R.B. MilIer. 1980. A chemical spot-test for aluminum and its va.lue in wood identification. IAWA Buli. n. s. 1: 104-109. Mabberley, D.J. 1987. The plant-book, a portable dictionary of the higher plants. Cambridge Univ. Press. Mana, J. 1921. Ausiralian timbers, their strength, durability and identification. 2nd Ed., Wa]ker, May & Co. Melbourne. Mennega, A.M.W. 1948. Suriname timbers 1. General Introduction, Guttiferae, Vochysiaceae, Anacardiaceae and Icacinaceae. Martinus Nijhoff, The Hague. Metcalfe, C. R. & L. Chalk. 1950. Anatomy oU the dicotyledons. Vol. I. Clarendon Press, Oxford. Mikesell, J. E. & R. H. Popham, 1976. Ontogeny and correlative relationship of the prirnary thickening meristem in four-o'clock plants (Nyetaginaceae) maintained under long and short photoperiods. Amer. J. Bot, 66: 997-1005. Miller, R, B. 1981. Explanation of Lhe codingprocedure, IAWA Buli. as. 2: 111-145. Qhtani, J., B. A. Meylan & B.G. Butterfleld. 1984. Vestures or warts - Proposed terniinology. IAWA Buil. as. 5: 3-8. Panshin, A.J. & C. DeZeeuw. 1980. Textbook of wood technology. 4th Ed.. McGraw-Hill, New Yc,rk, Parameswaran, N. & W. Liese. 1969. On Lhe formation and fine structure of septate wood fibres of Ribes sanguineum. Wood Sei. Technol. 3: 272-286. Parameswaran, N. & H.G. Richter. 1984. The ultrastructure of crystalliferous edis ia some Lecythidaccae with a discussion of their terminology. IAWA Buil. n.s. 5: 229-236. Quirk, J.T. 1983. Data for a coniputer-assisted wood identitication system. 1. Corarnercia] legumes of tropical Asia and Australia. IAWA Buli. n. s. 4: 118-130. Record, S.J. 1944. Keys to Anierican woods (continued). Trop. Woods 77: 18-38. Richter, H.G. 1977. Differential staining of oil and mucilage ia idioblasts of Lauraccae. 1AWA 197714: 76. - 1980. Occurrence, morphologv and taxonomic implications of crystalline and siliceous incliisions in the secondary xylem of the Lauraceae and related farriilies, Wood Sci. Technol. 14: 35-44. Sudo, S. 1959. Identification of Japanese woods. Buli. Govt. For. Exp. Stadon No. 118. Tokyo, Japan. Swain, E, H. 1927. Universal index to wood. Dep. of Public Lands. Queeasland For. Buli. No. 7.

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Takhtajan, A. 1980. Outline of the classificatiori of the fiowering plants (Magnoliophyta). Bot. Rev. 46: 226-359. - 1987. [Systema Magnoliophytorum.1 In Russian. Officina editoria 'Nauka', Leningrad. Thorne, R.F. 1976, A phylogenetic classification of the Angiospermae. Evolutionary Biol. 9: 35-106. Topper, S.M.C. & J. Koek-Noorrnan. 1980. The occurrence of axial latex tubes in the secondary xylem of some species of Artocarpus J.R. & G. Forster (Moraceae). IAWA Bu11. n.s. 1: 113-119. Vidal Gomes, A., L. Lopes Teixeira, E. Gomes Schaitza & R.M. Hofmeister. 1989. Perforation plates in vesseis of Citharexylum myrianthum Cham. (Verbenaceae). IAWA Buil. ri. s. 10: 27-34. Vliet, G.J.C.M. van. 1976a. Radial vesseis in rays. IAWA Buli. 1976/3-35-37. - 1976b. Wood anatomy of Rhizophoraceae. In: P. Baas, J. Bolton, & D. M. Catling (cds.), Wood sr.ructure in biological and technological rcsearch: 20-75. Leiden Bot. Series 3. Leiden Univ. Press. 1978. The vestured pits of Cornbretaceae and allicd families. Acta Bot. Neerl. 27: 273285. - 1981. Wood anatomy of the palaeotropical Melastoinaraceac. Blumea 27: 395-462. - & P. Baas. 1984. Wood anatomy and classification of the Myrtales. Ann. Mo. Boi Gard. 71: 783-800. Weber, W.A. 1982. Mnemonic three-letter acronyms for the farnilies of vascular planrs: a device forore effectivc herbarium curation. Taxon 31: 74-88. Welch, M.B. 1922. A method of identification of some hardwoods. J. Roy. Soc. New South Wales 56: 241-248. WeUe, B.J.H. ter. 1976. Silica grains in woody plants of the neotropics, especially Surinam. In: P. Baas,J. Bolton, & D.M. Catling (eds.), Wood structure in biological and technological research: 107-142. Leideri Bot, Series 3. Leiden Univ. Press. - 1980. Cystoliths in the secondary xylem ofSparattanthelium (Hemandiaceae). IAWA Buil. n.s. 1: 43-48. Wheeler, E.A. 1986. Vesseis per square millimetre or vessel groups per square millimetre? IAWA Buli. n.s. 7: 73-74. - , R. G. Pearsori, C.A. LaPasha, T. Zack, & W. Hatley, 1986. Computer-aided wood identification. Reference manual. N.C. Ag. Res. Serv. Buli. 474. Willis, J.C. 1973. A dictionary of the fiowering plants and ferns. 8th Ed., revised by H. K. Airy Shaw. Cambridge Univ. Press.

IAWA LIST OF MICROSCOPIC FEATUIRES FOR HARDWOOD IDENTIFICATION by a Cornmittee of he International Association 01 Wood Anatorniqs Veronica Angyalossy Alfonso, So Paulo, Brazil Pieter Baas, Leiden, The Netherlands Sherwin Carlquis, Claremont, California, USA Joao Peres Chimelo, So Paulo, Brazil Vera T. Rauber Coradin, Brasilia, Brazil Pierre Dtienne, Nognt-surMarne, France Peter E. Gasson, Km, UK Dietger Grosser, Mnchen, FRG Jugo flue, Highett, Victoria, Australia Kc co Kuroda, Kyoto, Japan Regis B. MiLer, Madison, Wiscons3n, USA n Ogata, Tsukuba, Japan Hans 'ieorg Richte:, Hamburg, FRO I3en J. H. zer Wlle, Utrecht, The Netherlands E1isabefl A. Wheeler, Raleigh, North Carolina, USA

Contenis Preface cknow1edgernents Explanatory Notes List of Features Name natomical Features Apper;dix Non-anatornl Inforrnation eferences

................................... ............................ ............................ .............................. .................................... ........................... ............. ................................

221 223 225 226 233 234 321 329

c ei; Sweeiiafruiicoa (Ferreir(.a spectabilis, Papilionaceae), n-ansver.se sec on, x 115. Photograph by couriesy of eter E. Gasson (Kew, IJK) and taken from a slide by Vera T. Rauber Coradin (Brasilia, Brazil).