This document has been scanned from hard-copy archives for research and study purposes. Please note not all information may be current. We have tried, in preparing this copy, to make the content accessible to the widest possible audience but in some cases we recognise that the automatic text recognition maybe inadequate and we apologise in advance for any inconvenience this may cause. ~~Il'ilCDn $~IJ~alfu:~ $~alDIl'il alll'il~ ~ , i: I~ I I I' 'I 1 1...-----1 LC(Q)jglI5Sjj~W rcIOJMMDSSn~~ ~[F ~aSaWa SVdnlSV 4l~HS12 ,------. 1\ .~ I 1 PENCIL STREAK, STAIN AND DECAY IN EUCALYPTUS AND SYNCARPIA IN NEW SOUTH WALES AUTHOR D. W. Edwards Forestry Commission of New South Wales Sydney 1982 1 Forestry Commission of N.S.W., Wood Technology and Forest Research Division Post Office Box 100, Beecroft, 2119 New South Wales, Australia Published 1982 National Library of Australia Card Number ISSN 0729-5340 ISBN 0 7240 6668 3 Printed in Australia by Star Printery Pty. Ltd. 21 Coulson St., Erskineville, N.S.W. 2043 2 CONTENTS Page ABSTRACT 4 INTRODUCTION 5 PENCIL STREAK IN EUCALYPTUS 7 OBLIQUA Macroscopic features Microscopic features Distribution Nature of pencil streaked wood Fungi in pencil streaked wood Production of pencil streak in vitro Role of insects DECAYS ASSOCIATED WITH PENCIL 14 STREAK AND TREE INJURIES OF E. OBLIQUA Macroscopic features Microscopic features Fungi causing decay PENCIL STREAK, STAIN AND DECAY IN 16 E. PILULARIS Initial investigation Whole cross section study Materials and methods Results Discussion Conclusions PENCIL STREAK, STAIN AND DECAY IN 18 E. NITENS Materials and methods Results Discussion Conclusions DARK HEART IN TURPENTINE 19 ACKNOWLEDGEMENTS 20 REFERENCES 21 APPENDICES 24 1 Forester's classification of Eucalyptus obliqua test logs cut at"Banshea State Forest 2 Culture media 3 Cultural and other characteristics of basidio mycetes isolated from Eucalyptus spp. 3 PENCIL STREAK, STAIN AND DECAY IN EUCALYPTUS AND SYNCARPIA IN NEW SOUTH WALES D. W. Edwards ABSTRACT The condition of pale-coloured eucalypts known as pencil streak is a reaction between the living tree and wound invasion by pioneer fungi introduced by ambrosia beetles. The nature, development and significance of pencil streak is discussed and the condition shown to be an unreliable indication of loss of wood strength. The relationship of decay to pencil streak in living eucalypts and the nature of stains in eucalypt logs are also discussed. Some other work on the stains and decays of eucalypts and turpentine is reported. 4 1. INTRODUCTION Occasional problems concerning the presence of Pencil streak i an old term fir t applied to thin black stain and decay in sawn wood of native species have lines of stain injarrah (Tamblyn 1937; Rothberg 1937). been brought to the attention of the Forestry This report deals primarily with pencil streak in Commission of New South Wales over the last 20 Eucalyptus ob/iqua L. Herit. (messmate), E. pilularis years. Many of these defects have been accepted as Srn. (black butt) and E. nitens Maiden (shining gum), normal for the species concerned and most work on but other investigations by Commission officers are them has not concerned itself with amelioration of the also reviewed. problem in living trees. Figure 1. Eucalyptus obJiqua mining timbers showing (a) pencil streak, (b) brown stain, and (c) an insect gallery. (LC 388) Figure 2. Eucalyptus obliqua mining timbers showing (a) brown stain, and (b) decay pockets. (LC 388) 5 .------ b Figure 3. Pencil streak in Eucalyptus obliqua heartwood. Insect galleries occur at (a) and pencil streak at (b). (LC8713) 6 PENCIL STREAK, STAIN AND DECAY IN EUCALYPTUS AND SYNCARPIA IN NEW SOUTH WALES D.W. Edwards Research Paper No. 1. 1982 ERRATA Caption on page 7 is deficient in data. The appended amending slip may be superimposed. 4a,b. Eucalyptus obliqua l. Herit. (messmate) 4c. E. viminalis Labill (manna gum) 4d,e. E. fastigata Deane et Maiden (brownbarrel) 4f,g. E. sieberi l. Johnson (silvertop ash) 4h,i. E. microcorys F. Muell. (tallow wood) The Commission apologises for the oversight. 2. PENCIL STREAK IN E UCA LY PTUS in timber u ed either to support cantilevered platforms OBLlQUA at the ore face, or as prop and shoring. The platforms, which must support six men and heavy drilling In 1964, trial shipments of messmate were sent from machinery up to 7 m above the tunnel floor, are Oberon to Broken Hill, New South Wales, for tests exposed to blast and are constantly wet. Their collapse below ground in the silver-lead mines. Because of mine would almost certainly cause serious injury or death. safety requirements any loss of strength due to hidden Props and shoring have less critical criteria but causes was considered to be completely unacceptable freedom from incipient decay is mandatory. a b I Figure 4. Pencil streak and stain in wood of five Eucalyptus spp. (Insect galleries may be seen in Figures 4c, 4e , 4g, 4h , 4i.) (Fe 3384) 4a,b. Eucalyptus obliqua L. Herit. (messmate) E. viminalis Labill. (manna gum) E. fastigata Deane et Maiden (brown barrel) 4f,9. E. sieberi L. Johnson (silvertop ash) 4h ,i. E. microcorys F. Muell. (tallow wood) 7 The quality of messmate available from Oberon in transparent and did not appear granular even in older 1964 was not good, pencil streak, decay and brown heartwood. It does not appear to be a cause of pencil stain being commonly found alone or together. The streak. mine management was concerned with problems of Within vessels close to insect tunnels aggregations of safety, and the prominent appearance of pencil streak thick-walled, dark-brown septate hyphae were found. and its apparent association with decay led them to These hyphae were irregular in shape, anastomosis was use it as an indicator of unsound wood (Edwards frequent, and in some specimens clamp connections, 1964). which indicate decay fungi , were present. However A field inspection of mess mate logs from Banshea there may have been several fungal species present S. F. was made by a group of forestry, sawmilling and including decays unrelated to pencil streak. mining officers in 1964 (Appendix I). It revealed a high A fe ature of most pencil-streaked zones was the incidence of decay, and defective timber, which was rarity- or absence of fungal or bacterial attack, reflected in the quality of mining timbers cut from determined either by the presence of fungal hyphae or similar material from the same forest (Figs. I and 2). bacterial cells, or by the breakdown pattern of cell Material of this type was rejected at the pit top. walls. 2.1 Macroscopic features In transverse log sections of messmate from Banshea S. F. pencil streak followed the direction of the rays. 2.3 Distribution Thus, if a transverse section was cut in exactly the right Examination of 738 authentic wood samples of horizontal plane, long, thin black streaks were seen to eucalypts in the Forestry Commission of New South extend from pith to outer heartwood. Pencil streak has Wales collection, supplemented by several thousand not been detected in sapwood (Fig. 3). wood samples used for teaching and field collections On the tangential face of boards from pencil which were made during this study showed pencil streaked logs the defect occurs as lenticular flecks of streak to be present in 23 eucalypt species (Table I). varying length, depending upon the number of ray cells From these results it appears that pencil streak, intercepted by the saw-cut. occurs chiefly in the subgenus Monocalytus but some In the eucalypts listed in Table I the appearance of affected species are scattered throughout pencil streak is the same except for jarrah, a dark red Symphomyrtus. timber, in which the black markings are larger and more diffuse. However, Tamblyn (1937) gives a description of pencilled jarrah very similar to that for Table I. Occurrence of pencil streak< in Eucalyptus E. obliqua. Figs. 4 and 5 show the appearance of pencil Spp.1 streak in a number of species. Similarities between pencil streak and black stains Subgenus Species around insect galleries in mess mate suggests that the two may have a common origin. 2.2 Microscopic features Monocalyptus E. marginata Donn ex Srn. Examination of pencil streaked wood under the E. jacksonii Maiden microscope showed that the dark flecks were due to E. muellerana Howitt coloured deposits in the vessels and surrounding ray E. laevopinea R. T. Bak. parenchyma cells (Fig. 6). Some of this material had E. baxteri (Benth.) Maiden and penetrated into the surrounding fibre lumina by way of Blakely the pits. It is formed originally in the ray parenchyma, E. eugenioides Sieb. ex Spreng. and in the early stage of deposition appears as a E. cameronii Blakely and McKie hemispherical granular mass with a pit opening at its E. oblonga DC apex. E. pilularis Srn. E. obliqua L'Herit. These deposits were very similar to heartwood E. delegatensis R. T. Bak. crystals described by Chattaway (1952) but were E. jastigata Deane and Maiden seldom found to be associated with fungal hyphae. No E. sieberi L. 10hnson evidence of pencil-streaked wood was found more than E. radiata Sieb. ex DC. a few fibre cells from rays, but individual fibres were E. piperita Srn. occasionally filled with similar dark material in an E. campanulata R. T. Baker and apparently random fashion. The deposition of these H. G. Smith substances follows the normal path of water movement in hardwood. Symphomyrtus E. guilfoylei Maiden E. erythronema Turcs. Amorphous, brown, non-birefringent deposits were E. camaldulensis Dehnh. found in large amounts in vessels, whether tyloses were E. viminalis Labil!. present or not. When tyloses occurred, deposits were E. badjensisde Beuz. and A. Welch frequent in the spaces between them. They were found E. microcorys F. Muel!. also in ray parenchyma and fibres and it was a Uncertain Dum-Cors. combination of the two types of deposits which E. scabra produced pencil streak. Abundant polyphenol-like substances also occurred t Eucalypts classified in order suggested by Pryor and Johnson in ray parenchyma and fibres. This material was ( 1971 ) 8 The occurrence of pencil streak in E. guilfoylei, E. kino which is a dark-coloured phenolic exudate found jacksonii and E. marginata, all Western Australian in veins or in pockets in which the parenchyma bridges species and its absence from plant groups other than have broken (Hillis and Brown 1978). Sub lances eucalypts suggests that the plant responses which give which appear similar but are not kino occur in E. rise to this feature are of great antiquity. obliqua. It is most likely that they are produced by ray parenchyma cell at the sapwood j heartwood 2.4 Nature of pencil streaked wood transition zone following stimulation by fungi present Tamblyn (1937) described substances associated in the sapwood. with pencil streak in the red timber E. marginata as a e • , .- t• Figure 5. Pencil streak and stain in wood of seven Eucalyptus spp. (Insect galleries may be seen in Figs. 5b and Se .) (FC 3387) Sa . Eucalyptus laevopinea R. T. Bak. (si lvertop stringbark) 5b. E. baxteri (Benth,) Maiden et Blakely (brown stringybark) 5c. E. marginata Donn. ex Sm . (jarrah) 5d. E. pilularis Sm . (blackbutt) Se. E. punctata DC (grey gum) Sf. E. siderophloia Benth. (grey ironbark) 5g . E. fastigata Deane et Maiden (bro wnbarrel) 9 a b c d e Figure 6. Deposits in cells of Eucalyptus obliqua L. herit. associated with pencil streak. (Polaroid - D. W. Edwards) a. Fungal hyphae in vessel affected by pencil streak, b-e. dark deposits in vessels, ray parenchyma and fibres. 10 COOH COOH Sugars - HoO-OH OO-OH OH OH Shikimic acid pool 5-dehydro shikimic acid o \ COO·R COOH ellagitannins ~ HO OH ~ HOVOH~ -~HOVOH OH OH ellagic acid gallic acid esters gallic acid Figure 7. A possible biosynthetic pathway for the formation of ellagic acid. (After Birch 1963; Brown 1966; Geissman 1963a, 1963b; Hathway 1957; Hathway and Seakins 1957, Humphries 1965; Towers 1976.) The introduction into wood of additional oxygen wider distribution among eucalypts than present following insect injury or fire may be a necessary knowledge indicates, and Bate-Smith (1963), who condition for this transformation which might also be noted that it has not been found in ferns, gymnosperms due to metabolic changes brought about by drought or or monocotyledons, thought that it had ''the disease. possibilities of being a remarkably accurate taxonomic Gardner and Hillis (1962) reported large amounts of guide". ellagic acid in eucalypts (Table 2) and said that the Clusters of crystals thought to be ellagic acid were amount of free ellagic acid varies both between and found in the heartwood of many eucalypts in within species. The outer heartwood contains more association with fungal hyphae (Chattaway 1952). For ellagic acid than inner heartwood, especially in large these eucalypts which contain large amounts of trees and some occurs in sapwood (Hemingway and ellagitannins or ellagic acid, she suggested that, as the Hillis 1971). water content of the heartwood decreases during heartwood formation, the resultant super-saturated Table 2. Occurrence of ellagic acid in Eucalyptus solution of ellagic acid could be "nucleated" by the obliqua L. Herit. (After Gardner and Hillis 1962.) hyphae to form crystals. In this study little evidence of fungal hyphae associated with ellagic acid could be found, and this may be explained by the observation of Hillis (1953) that ellagic acid is present in all eucalypt Age of Trunk Ellagic acid tissues even when fungal hyphae have not been Sample tree Diameter (% oven-dry detected, suggesting that ellagic acid is a normal (yrs) (mm) wood) metabolite. Inner heartwood 220 508 1.15 Biosynthetic pathways for ellagic acid have been Outer heartwood 2.85 discussed (Geissmann 1963a, 1963b; Hathway 1957; Inner heartwood 180 406 0.50 Hathway and Seakins 1957; Neish 1964; Brown 1966) Outer heartwood 1.85 and their ideas are summarised in Figure 7. Brown Sapwood 0.10 warned that our knowledge of these steps is not satisfactory but agrees that the formation of gallic acid from glucose via the shikimic acid path is likely (Brown Tannic acid and tannins (either condensed or 1966). Gallic acid - sugar esters could then be formed hydrolysable) are present in eucalypts and both but the esterification reactions are unknown. Finally gallotannins and ellagitannins may be present in the Hathway (1957) and Hathway and Seakins (1957); same eucalypt species (Hillis 1953, Hillis and Carle have shown that mushroom polyphenoloxidases 1959, H umphries 1965). Hillis and Carle say that many convert gallic acid to ellagic acid quantitatively. A pale-<:oloured eucalypts contain large amounts of possible pathway for the formation ' of extra ellagic ellagitannins, most of which can be removed from acid and ellagitannins in which fungi play a role fresh wood by prolonged extraction with water and therefore exists in living trees. alcohol. Polyphenols of red-<:oloured eucalypts are Other possible causes of stains around beetle largely insoluble in these solvents. galleries have been suggested (Gadd & Loos 1947). With the exception of E. moluccana Roxb., the Verrall (1943) attributed similar stains to ambrosia recorded distribution of ellagic acid in eucalypts is fungi intensified by insect secretions or other confined to the Macrantherae and Renantherae. organisms but Gadd & Loos think it more likely to be Whether this is significant in terms of phylogeny is not due to the strain of ambrosia fungus carried by the known. It is possible that ellagic acid may have a much vector beetles. 11 2.5 Fungi in pencil streaked wood diameter were made. The blocks were sterilised by (a) Materials and Method fumigation with propylene oxide vapour and placed in One hundred and eighty wood chips each I mm long 220 g glass jars on sterile washed vermiculite, wetted to of E. obliqua containing pencil streak were inoculated capacity. on to malt/ yeast extract agar and incubated at 27°C After 2 weeks incubation to stabilise moisture for 12 weeks. content each hole was inoculated with one of the (b) Results following test fungi: Fungi isolated are shown in Table 3. Geosmithia swiftii Leptographium sp. Paecilomyces varioti Table 3. Fungi isolated from pencil streaked areas of Phialophora (Prob. P. Jastigiata) (two isolates) Eucalyptus obliqua L. Herit. Poria mollusca Yeast (unidentified) There were six blocks for each isolate and one group Number of of 6 controls. The inoculated blocks were incubated for Species Isolates 12 weeks at 27° C and 70% relative humidity in glass tanks. At the end of the test the blocks were split at the Basidiomycetes (decays) holes and examined for evidence of pencil streak. - Poria mollusca (Fr.) Cooke 2 (b) Results - Unidentified species 2 With the exception of two Phialophora spp. isolates, growth was good. In Phialophora inoculated blocks Pioneer and/ or staining fungi growth was either sparse or absent. No evidence of - Cephalosporium sp. I pencil streak was found in any of the blocks. - Geosmithia swiftii (Swift) PiU' I These results suggests that pencil streak formation - Leptographium sp. 3 requires the existence of living host cells. - Paecilomyces varioti Bainer 2 - Penicillium spp. 6 2.7 Role of insects - Phialophora Sp.2 6 Association of pencil streak with insect damage - Unidentified yeasts 9 seems likely in view of the constant presence of Saprophytes and contaminants platypodid (ambrosia) beetles such as Austroplatypus - Monilia sp. I incompertus (Schedl) Browne and associated black - Mucor spp. 2 stains. This species was originally named Platypus - Pestalotia sp. I incompertus by Schedl (1968), but subsequently F. R. Browne (1971) erected a new genus, Austroplatypus, to which he assigned P. incompertus. The type I Formerly Penicillium bacillisporum Swift. specimens are in the Victorian National Museum. This 2 Probably P. jasligiala Lager. Lund. and Melin. species is one of the few platypodids associated exclusively with living, apparently healthy trees The basidiomycetes including Poria mollusca will be (Browne 1971) and it attacks a number of N.S.W. discussed later. Whilst they were found adjacent to eucalypts (Table 4). pencil streak zones they occurred also in brown areas There is an abundant literature on ambrosia beetles from the pith outwards. This appears to be a classical and their associated fungi: Baker 1963; Bakshi 1950; example of the succession of pioneer and decay fungi Batra 1963a, 1963b, 1967; Batra and Francke (Shigo 1979) as components of the decay processes Grosmann 1961 ; Batra and Michie 1963; Cooke 1977a, which follow injuries to living trees. 1977b; Davidson 1955; Farris 1963 ; Farris and Funk The role of Geosmithia swiftii is unknown but it is 1965; Faulds 1973; Fernando 1960; Finnigan 1963; commonly found as a pioneer fungus of wounded Francke-Grosmann 1956a, 1956b, 1957, 1963, 1965, trees. The genera Phialophora and Leptographium 1967; Gadd and Loos 1947; Jones 1959; Lagerburg, contain a number of fungi which cause dark stains in Lundberg and Melin 1927; Leach 1940; Litchwark timber, and species of the latter are frequently 1978; Norris and Baker 1967; Schneider-Orelli 1911, imperfect stages of Ceratocystis which cause blue 1913; Verralll943; Webb 1945; Wilson 1959). Between stains of both eucalypts and pines. them a composite picture may be built up of fungi associated with such stains. There is considerable Yeasts are common components of wound-altered confusion as to the identity of many of the organisms, wood and as contaminants which grow rapidly on but Batra (1967) has done much to clarify this freshly cut, wet timber. situation. 2.6 Production of pencil streak in vitro Ambrosia beetles are widely distributed in Australia To determine whether pencil streak could be attacking sta.nding fire-killed trees (Hogan 1948) produced under laboratory conditions the following freshly felled logs, and green, sawn timber. They do not test was made: attack seasoned wood. Black halos surrounding insect (a) Materials and method tunnels are a feature of platypodid beetle attack, and Blocks of E. obliqua, 30 mm3, with a moisture result from introduction of fungi by adult females. content of approximately 40% were cut from non These fungi, which may be specific to a particular stained areas of a pencil streaked log. On the tangential insect species, line the tunnel walls and provide food face of each block four holes 25 mm deep by 2 mm for developing larvae. (Cooke 1977a, 1977b). 12 Figure 8. Mycetangia of Austroplatypus incompertus (Schedl) Browne. a. Mycetangia on pronotum of female. b. Mycetangial orifices. (Photos R. Johnson). 13 The mechanism by which fungal spores are Colony viability requires maintenance of an transferred from tree to tree was once thought to be unobstructed gallery to the outside air'. To do this the accidental, e.g. deposition of spores on insect hairs. insect must move to the surface at frequent intervals to Batra (1963a), Farris (1963) Francke-Grosmann eject frass, and a short tube of chewed up wood is often (1963), and others showed however that many built out from the bark. ambrosia beetles have special organs - mycetangia in which spores and hyphal fragments are kept viable but inactive until needed. Mycetangia may be located In E. ob/iqua from Banshea SF a long, rambling on the pronotum, or behind the coxa below the sides of type of gallery has been observed (Fig. 3). The galleries the pronotum. The mycetangia of A. incompertus are are circular to oval in cross-section and lined by a jet heart-shaped, and located on the upper surface of the black layer of fungal hyphae. Dark deposits similar to pronotum (Figure 8). those in pencil streaked areas extend about 2 mm into the wood. The brood chambers have a similar lining. A. incompertus (usually male adults) tunnel horizontally through bark into inner heartwood.' Secondary tunnels are then bored, often at right angles The length of various stages of this insect's life cycle to the entrance tunnel, i.e. in a tangential direction. is not known. The beetles are strong flyers and their Above and below the main galleries brood chambers distribution has interesting parallels with that known are then bored. The entrance hole is probably the exit for pencil streak as can be seen by comparing (Tables I hole. and 4). Table 4. Eucalyptus spp. attacked by Austroplatypus incompertus (Schedl) Browne. Subgenus Species Authority Monocalyptus E. muellerana Howitt I E. laevopinea R. T. Bak. I E. macrorhyncha F. Muel!. ex Benth. 2 E. baxteri (Benth.) Maiden and Blakely 2 E. pilularis Srn. 1,4 E. obliqua L'Herit. 1,2,4 E. delegatensis R. T. Bak. 2,4 E. Jastigata Deane and Maiden 1,2,4 E. consideniana Maiden 3 E. sieberi L. 10hnson 1,2 E. dives Schau. 3 Symphomyrtus E. botryoides Srn. 3 Uncertain E. scabra Dum-Cours. 2 I. Campbell, 2. Harris, 3. Harris (probable) (Campbell, K. Forestry Commission of New South Wales - Personal communication), 4. Browne (1971). 3. DECAYS ASSOCIA TED WITH PENCIL This is a typical example of the decay process (Shigo STREAK AND TREE INJURIES OF E. OBLlQUA 1979) where initial invasion by pioneer fungi is 3.1 Macroscopic Features followed by decolourization and finally decay. Decay in pencil streaked wood is best seen in freshly sawn logs as a water-soaked irregular heartwood zone which may extend almost to the sapwood. It appears to 3.2 Microscopic Features enter the tree around knots, and this is clearly shown Hyphae associated with these decays are thin, during conversion. Affected wood differs in colour hyaline and with or without clamp connections. from normal heartwood, being pale to very dark Irregularly shaped chlamydospores occasionally brown or sometimes bright yellow. In extreme cases occurred in the vessels. There are no other softening occurs, and there may be severe decay near distinguishing microscopic features except that where the pith. In freshly awn timber such damage is severe yellow staining occurs brown thick-walled prominent but is more difficult to see as the timber hyphae with infrequent clamps are present in both dries out. vessels and rays. I Campbell , K. Forestry Co mmission of ew South Wales - Personal communicatio n. 14 3.3 Fungi causing Decay other eucalypts are uniquely Australian, others are Identification of decays in logs and sawn timber is cosmopolitan. For this reason, and despite the lack of seldom possible unles sporophores are present. published data on the cultural characteristics of Despite efforts by Waiters and Da Costa there is as yet Australian species it was decided to record the insufficient information on Australian wood-rotting Australian isolates in terms of Noble's key, with fungi to allow satisfactory identifications on the basis variations described by Fritz (1923); Davidson et a/ of cultural characteristics as is done in Canada (Nobles (1938); Etheridge (1957), and Jorgenson and Vejlby 1965). This situation is likely to persist for some years. (1953). Isolates FC 859, 860 and 911 from E, pi/u/aris failed to survive in culture and isolates FC 2841,3047 (a) Iso/ation of Decay Fungi and 3417 were substituted for them. Discs 150 mm thick were cut across full trunk cross sections of selected logs of either E. obliqua or E. Cultures were grown on malt/ yeast extract agar as pilu/aris. Other pieces 150 mm )( 50 mm2 were taken described by Nobles except that "Oxoid" agar and J from brown-stained areas of several flitches of the malt extract were substituted for "Difco-Bacto" same two species. All samples were wrapped products. immediately in clean polythene sheet and isolated the following day. All microscopic examinations were made on material mounted in 5% aqueous potassium hydroxide From each piece a number of small samples (50 mm and stained with 1% aqueous phloxine. Noble's gum )( 250 mm2) were cut, depending on the size of the disc, guaic solution was used to test for formation of but never less than three. Each sample was flame extracellular oxidases. sterilized on all faces, split with a sterile chisel and from the centre of each, three chips were transferred on to Results are shown in Appendix 3. each of three petri dishes containing malt/ yeast extract/ glucose agar (Appendix 2). (c) Sporophore production Etter (1928), Koch (1962), Niederpreum (1963), The fungi isolated from E. obliqua which had Plunkett (1953), and Raper and Krongelb (1959) basidiomycete features are shown in Table 3. Isolates studied the role of environment and nutrition in the FC 859, 860 and 911 from E. pi/u/aris ex Manning formation of wood decay fruiting bodies. Despite these River National Forest also showed typical decay investigations successful laboratory production of features. sporophores of wood decays remains an art rather (b) Cultural characteristics of decays isolated from than a science. The technique which has proved most E. obliqua and E. pilu/aris successful to date is that of Badcock (1941, 1943) as N obles (1965) in her definitive work on cultural modified by Tamblyn and da Costa (1958). This characteristics of wood-inhabiting Hymenomycetes method was further modified for this study. Other gives a mUltiple choice key for identification of 149 changes suggested by Yurchenko and Warren (1961) Canadian species of wood decays based on published were considered and rejected as being too specialised, data on their cultural characteristics. Although it is or likely to be useful with a very limited range of probable that some wood decays of E. obliqua and species. Figure 9. Modified 8adcock sporophore test. (LH 19225) 15 A basal medium of sawdust plus nutrients was (b) Unidentified Basidiomycetes. Source: Isolates prepared (Appendix 2). The medium was wetted until FC 518 , FC 519, and FC 705 were from E. it just failed to absorb further water, then a small obliqua in Banshea State Forest, Oberon, and amount of dry medium was added to absorb any free Isolates FC 859, FC 860 and FC 911 from E. water. It was then loosely packed into 300 mL bottles pilularis, Manning River National Forest, until level with the top of the neck and the bottles Taree, New South Wales. capped with aluminium foil and autoclaved at 103 kPa (c) Coniophora sp. (probably c. olivaceae (Fr. ex for 20 minutes. Pers.) Karst.). Source: Division of Building When cool the sterile media was inoculated with 5 Research, CSIRO. Cartwright and Findlay mm cubes of the test fungus taken from 10-20 days old (1958) describe the species as a brown cubical malt/ yeast extract Petri dish cultures, and incubated rot of heartwoods, which grows well in warm at 27°C until the mycelia had penetrated throughout climates and is common in central and northern the containers. Europe and America. Cunningham (1965) says From clear sapwood of Pinus radiata D. Don, 45 it occurs in Australia, New Guinea and New mm x 50 mm 2 blocks were cut so as to expose two Zealand. transverse faces at right angles to the short axis of the Screwcap jars (110 g) with a wide mouth were filled tree. The blocks were routered to give a 45 mm with 5 g of vermiculite, and 30 mL of distilled water. diameter hole, 19 mm deep, which allowed the block to On this wetted bed of vermiculite were placed 20 cm2 be fitted tightly over the neck of the culture bottle. A feeder strips of flame kurrajong veneer 2 mm thick and similar hole 5 mm deep was made in the top of the the assembly autoclaved at 103 kPa for 20 mins with block to act as a water reservoir. the caps loosely screwed in place. The feeder strips were then inoculated with the test organism, the metal When the fungus had grown sufficiently the caps screwed down but not so as to exclude all air. The aluminium foil caps were replaced under aseptic assembly was then incubated for 14 days at 27°C. conditions by the wooden blocks which had been previously autoclaved at 103 kPa for 20 minutes, and Test blocks were cut from inner heartwood of E. the block and bottle neck covered with sterilized obliqua from Oberon, N.s.W. and air dried to give a polythene sheet. The assembly was incubated at 27° C final weight of about 4 g. All were selected to give a until hyphae appeared at the edges of the blocks. The similar density group and cut to present a machined assembly was then transferred to glass tanks, the water tangential face in contact with the test organism. The well in the top of each block filled with sterile water, test blocks were dried at 105° C for 24 hours to and the tank covered with sterile polythene sheet. The determine oven dry weight, then soaked in tap water assembly was incubated at room temperature in for 10 days to give a moisture content of about 80%. subdued light until sporophores appeared. (Figure 9.) The blocks were then autoclaved at 103 kPa for 30 Occasional replenishment of the water reservoir was mins. necessary. After the test fungus had grown sufficiently on the Of the six wood decays tested only two produced feeder strips (14 days) a test block was placed on each sporophores. One of these was identified in 1967 by strip. The lids were again partially screwed down and Walters l as Poria mullusca (Fr.) Cooke, who was of the jars incubated in glass tanks over a free water the opinion that it agreed in all details with surface at 27°C for either 9 or 10 weeks. Cunningham's description except for the rot type. Results - See Table 5. Waiters however, does not consider that Cunningham's rot types are always correct. The All isolates tested on wood block substrates identity of the other was not established. produced some decay after 9-10 weeks but no significant differences were obvious between them. (d) Decay potential of isolates from E. obliqua and The low level of decay (4-6%) suggests the need for E. pilularis caution when interpreting the results. The ability of a fungus to decay timber in service is not always the same as in living trees, Duncan (1958), Feist et al (1971), Hartley (1958). Many heart rots do 4. PENCIL STREAK, STAIN AND DECAY IN E. not persist after the tree is felled and the wood must be PlLULARIS transformed by pioneer fungi before serious damage occurs. 4.1 Initial investigation At the request of the Commission's Engineering Seven basidiomycetes, four from E. obliqua and Aspects Research Group, fifty-seven samples of E. three from E. pilularis were compared with two pilularis were examined for bacteria, fungi and other common wood-decaying fungi supplied by CSIRO symptoms of wood biodeterioration (Edwards 1979a). Division of Building Research. These samples were from complete log cross-sections The fungi used were: taken from butt and top logs of six trees supplied by (a) Poria mollusca (Pers. ex Fr.) Cooke. FC 511. that Group. Source: E. obliqua-Banshea State Forest, Hand cut sections in transverse, radial longitudinal Oberon, N.S.W. This identification is tentative and tangential longitudinal directions were made from but Cunningham (1965) records it from both each sample and mounted in lactophenol aniline blue. Europe and New Zealand and Lloyd (1910) All samples were examined for deposits within the from Australia. It produces a brown rot. cells, tyloses, fungi and bacteria. 'N. C. WaIters Division of Forest Products CSIRO. Personal Communication. 16 Table 5. Relative decay potential of six basidiomycetes grown on Eucalyptus obliqua heartwood at 27° C for 9 and 10 weeks. Growth Number of Average Isolate Origin time replicates weight loss s.d. (weeks) (%) Poria Sp.1 E. ob/iqua 10 10 4 1.65 FC 518 9 6 4 1.26 FC 705 " 9 6 5 2.59 FC 859 E. pilularis" 10 8 5 1.64 FC 860 10 10 6 3.00 FC911 " 10 10 4 1.32 Control " 10 8 )2 1.39 " I Probably P. mollusca. 1 Probably due to contaminants. Cell deposits were found in most sections examined samples, which is presumptive evidence that they were and were readily identifiable in all three planes. Similar not wood decays. In a few samples dark coloured deposits had been previously described by Dadswell hyphae of a type commonly associated with ambrosia (1972), who distinguished them from kino-like beetles were noted. No fruiting bodies were produced substances. There was no apparent relationship so this could not be confirmed. between these deposits and variations in the Well developed tyloses were present in most samples mechanical strength of the samples. and in the genus Eucalyptus this is a reliable indication The occurrence of ellagic acid deposits similar to of heartwood. In many cases the interfaces of the those described by Chattaway (1952) was very variable tyloses were covered with extraneous materials. and did not appear to be associated with fungal hyphae. However, as Chattaway notes, ellagic acid is deposited only in cells which are already dead and 4.2 Whole cross section study from which hyphae may long since have disappeared Samples from a further 5 trees of regrowth black butt by autolysis. Ellagic acid is very common in pale supplied by the Engineering Aspects Group were next coloured eucalypts and in some species is responsible examined to determine whether wood destroying fungi for problems during conversion to paper pulp. and/ or bacteria were associated with changes in mechanical properties at two levels in each tree. The Fungal hyphae were common in sapwood and more history of these samples and their mechanical and particularly in heartwood. In nearly every case the physical properties have been reported elsewhere hyphae were thin and hyaline and without clamp (Ghali et al 1979). connections. Since no fruiting bodies were produced it was not possible to determine whether these hyphae were of pioneer fungi or wood decays. No 4.3 Materials and Methods deterioration of cell walls either from the lumen Complete cross-sections were cut from butt and top inwards, or within the cell wall itself was found in any logs of five trees as shown in Figure 10. interior dark coloured zone exterior exterior 2 3 4 5 6 Figure 10. Diagram of a typical discoloured sample selected from a flitch of Eucalyptus pilularis (After Ghali et a/1979). From each available position, thin hand-sections 4.5 Discussion were made and examined from the transverse, radial From Table 6 it can be seen that the samples were and tangential wood faces as before. Some flitches chiefly heartwood, i.e., tyloses were present in most sampled had fewer and some more sample blocks than samples. Fungal hyphae were widespread in the wood shown in Fig. 10. of all five trees but there was no obvious relationship with any previously determined wood property. 4.4 Results There was no evidence of clamp connections (which The results are summarised in Table 6. indicate potential decay fungi) but these structures are 17 frequently absent and more reliance is placed on composition. These appeared structureless and were evidence of cell wall breakdown. No degrade of cell not birefringent. They caused wood discolouration but walls by decay fungi, soft rots or bacteria was seen and their presence could not be related to any mechanical again no link could be established between presence of properties of the samples. fungal hyphae, or heartwood crystals associated with such hyphae, and breakdown of cell wall structure. The fungi present were therefore probably pioneer 4.6 Conclusions organisms associated with the initiation of the decay No link could be established between physical and processes described by Shigo (1967, 1979). mechanical properties of the samples and presence or In many samples vessel and fibre lumina were partly absence of fungi, bacteria, heartwood crystals or or completely filled by dark deposits of unknown amorphous cell deposits. Table 6. Presence of fungi, tyloses, heartwood crystals and dark amorphous deposits in wood of re growth blackbutt. Tree Sample Heartwood Poly- Amorphous No. Height Fungi Tyloses Crystals phenols Deposits Top +++++' -++++ +++++ ++-++ -++++ Butt +-+-- ++++++ -++++- --+ -++++- 2 Top - NS-+ ++NS++ ++NS-+ ++NS++ ++NS++ Butt ++++-+ ++-+++ -+- ++++++ --+++- 3 Top -++-+ ++++- ++++- ++++- ++++- Butt 1111 111111 --+++t+ ++- -++++++ 4 Top +-+- +++++ +++++ +++++ +++++ Butt +-++++ ++-+++ ++-+++ +++++- 6 Top +++++ +++++ -++++ ++-++ +++++ Butt -+--+++- 11111 -+-++ -++++++- +1111111 I Each symbol represents a sample position across a tree from bark to bark. NS means no sample was available. S. PENCIL STREAK, STAIN AND DECAY IN Group D - Heavy Stain: Limited slope of grain as E. NITENS above, borer holes and stained areas in excess of Pencil streak, stains and decays of the same general Group C. type as for E. obliqua and E. pilularis also occur in From each sample hand sections were made from shining gum (E. nitens Maiden). the transverse and radial longitudinal faces. These As part of an examination of the effects of such were mounted in lactophenol aniline blue stain and stains, the mechanical properties of four visual grades examined at 600x for evidence of biological degrade. of shining gum were determined (Ghali 1981), and material from each grade was examined S.2 Results microscopically for evidence of biological degrade The results obtained are shown in Table 7. (Edwards 1979b). S.3 Discussion S.l Material and Methods Ghali et al (1981) showed significant differences Four groups of samples were selected from impact between means of the impact toughness tests at a T toughness test blocks after these had been broken on a value of 1.96 as follows: Denison tension testing machine. These were classified Visual No Light Medium Heavy into the following visual grades:- grade stain stain stain stain Group A - Clear: Normal, clear, defect-free material Mean (loul) 20.91 18.96 18.24 16.42 I without imperfections, but allowing sloping grain in Before interpreting the biological results some 15. qualifications must be borne in mind. The type of data Group B - Light Stain: Defect-free material, allowing obtained comes from a search for gross differences, sloping grain as in Group A, borer holes up to 2 mm such as a clearly graded series of decay intensities diameter, but not exceeding 10 in any lOO cm2• Black between each of the visual grades. This did not occur, stain distribution density less than I cm2 j 100 cm2. and the results could not be quantified because: Group C - Medium Stain: Limited slope of grain as (a) Both dark deposits in the lumina of wood cells above, borer holes up to 3 mm wide, but not exceeding and the presence of dark hyphae will cause 20 in any 100 cm2• Black stain distribution density stain. In these tests the relative contribution of between 1-2 cm2j 100 cm2• each factor could not be determined. Further 18 Table 7. Occurrence of fungi and associated materials in unseasoned shining gum (Eucalyptus nitens) impact toughness (Denison) samples. stain indicators decay indicators visual No. of dark dark cell wall hyaline grade samples freq. deposits hyphae degrade clamps hyphae (A) N 9 14 1 0 8 no 54 %A 17 26 2 0 15 stain % Total 8 12 1 0 7 Ill) (B) N 10 13 0 2 10 light 53 %B 19 25 0 4 19 stain % Total 9 12 0 2 9 '~ (C) N 28 12 3 1 16 medium 51 %C 55 24 6 2 31 stain % Total 26 11 3 1 15 (D) N 32 27 5 2 36 heavy 53 %D 60 51 9 4 68 stain % Total 29 24 5 2 32 the area sampled was minute in relation to the and heavy stain. Differences between the first two total stained area, which makes an accurate and last two groups were marked. assessment of the total effect of the stain fungi difficult. • The presence of dark hyphae appeared to be quantitatively indistinguishable in the groups, no (b) Cell wall degrade was the only reliable stain, light stain, and medium-stain, but there was quantitative measure of decay available in this a large difference between these groups and heavy study. The presence of clamps indicates that stain. This may be due to severe insect attack with potential decays were present but whether they its related dark fungal hyphae. Again there was no were active, or could grow further in the wood obvious link with mechanical properties. after pioneer organisms had prepared the way was not determined. Again some of the hyaline • Small differences (less than 10%) between samples and dark hyphae which were seen to be without showing decay are probably not significant in clamps could be potential decay fungi since tests of this type since the decay was confined to a clamps are not always produced. few cells in each sample and was not made up of large significant patches of damaged wood. A link (c) A number of samples in all groups contained between mechanical characteristics of these fungal fruiting bodies in the lumen of the samples and the number showing signs of decay vessels. These were Phialophora m elin ii, a would be difficult to establish. Trichocladium sp. and a Ceratocystis sp. Phialophora spp. are common pioneer fungi • A link is possible between the number of samples and also saprophytes of plant material and they showing hyaline hyphae and those with dark occur in soil. They are often found in deposits. However it is not known whether the association with dark stains of wood and wood dark hyphae contributed to the dark deposits. pulp and may occur in black zones around borer holes. There is no evidence that they cause significant wood decay. Whether these 5.4 Conclusions were in the wood before felling or entered after On the microscopic evidence available there did not sawing and storage is not known. Much of the appear to be any correlation between either visual hyphae present may have been of these species grades or impact toughness tests and the presence of but this could not be determined decay, nor was there any evidence of a relationship microscopically. Ceratocystis spp. commonly between visual grading and the presence of staining cause blue stain in timber and are widespread in fungi and dark wood-cell deposits. slash on the forest floor. Trichocladium spp. include pioneer species (Shigo, 1967). 6. DARK HEART IN TURPENTINE The incidence of the condition known as dark heart With these points in mind it is possible however to of turpentine is often associated with defective wood draw some very broad inferences. by timber users. Following an examination of the • Dark deposits in the cells appear to be correlated mechanical properties of three trees of turpentine with the degree of stain as assessed by visual (Syncarpia glomulifera (Sm.) Niedenzu) showing this grading but not as well as with the mechanical defect Ghali (1978) 25 Izod samples were tested for the properties. There was little difference between no presence of microbial deterioration. These samples are stain and light-stain grades or between medium described in Table 8. 19 Table 8. Details of izod samples taken from three turpentine trees felled in the Karuah River Management Area. (Modified from Ghali, 1978.) Billet Details Tree Date Tree Tree No. Felled Height Dia. Billet Sample Billet Billet Billet Samples Code Nos. Dia. Height Colour with Above Sapwood Ground I (m) (cm) (cm) (m) .) I 1. 10.8.78 20 44 A Al-5 44 Red to AI, A5 Pink B Bl-4 36 4 B1, B4 C Cl-4 32 8 Cl, C4 " 2. -7.78 27 95 D Dl-2 N.A. 5 Dark D1 Brown E El-6 67 5 El, E6 " 3. -7.78 24 52 H Hl-5 44 5 Dark HI, H5 Brown Ghali (1978) concluded from his study that 'the Ghalil has questioned whether brown heart in influence of the discoloration upon the impact turpentine is a form of false heartwood. We have no properties of turpentine is obvious and evidence to confirm or disprove this. distinguishable' and that further study of the problem was importinit. False heartwood is a well known phenomenon (lane 1970; Panshin and de Zeeuw 1970; Perelvgin 1965). It occurs in a number of northern hemisphere species, A loss of impact strength associated with dark stains e.g. beech, birch, maple, poplar and pines and may be of wood is sometimes associated with wood decay. To test this hand sections were cut fro.m each sample, dark or red brown, lilac, violet or dark green in colour. stained in lactophenol aniline blue mountant and The vessels within the false heartwood area are heavily tylosed. The reasons for its formation are not clear, but examined at 600x under normal and polarized light. may range from minor changes in the tree's metabolism which result in the deposition of coloured With one exception (sample D2), no pioneer, extractives in the wood and induce tyloses, up to more staining, soft rotting or decny-causing fungi could be deep seated changes due to the invasion of fungal detected. No evidence was seen of cell degrade, due hyphae. either to these organisms, or to bacteria. There was thus no detectable difference between the red and dark There are however similar staining problems not brown samples due to microorganisms. Sample D2 necessarily associated with false heartwood. Shigo showed slight traces of an unidentified fungus, but (1979) describes the processes of discolouration and only in a few cells. No identifiable damage could be decay as a complex continuum of events that merge associated with it. and overlap in time and place. He describes a sequence of events leading from biochemical changes due to There appeared to be a greater deposition of injury, to invasion by bacteria and non-decay fungi, materials in the vessels of the dark brown samples than and finally attack by decay fungi. Discolourations in the red specimens, but the composition of this which arise from such causes may occur after material was not determined. Incipient deterioration heartwood is formed. A typical example ofthe latter is of the cell wall could not be detected by polarization brown stain of E. obliqua due to incipient decay. methods because of the nature of the samples. 7. ACKNOWLEDGEMENTS The lack of fungal hyphae or bacteria does not mean The assistance of Mrs. J. L. 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No. 2. pilzzuchtenden. Obstbaumborkenenkafen Jorgensen, E., and Vejlby, K. (1953). A new Xyleborus (Anisandrus) dispar und seinen N ahrpilz. polyphenol oxidase test. Physiol. Plant. 6, 533-537. Zentr. Baketerio!' Parasitensk. 38, 25-110. 22 Shigo, A. L. (1967). The early stages of discoloration [In The Filarnentous Fungi.] Vol. 2. Biosynthesis and decay in living hardwoods in Northeastern and metabolism [Ed. Smith, J. E., and Benay, D. R.] United States: A consideration of wound-initiated Edward Arnold, London. discoloration and heartwood. XIV IUFRa. CONGR. 9 (41), 117-133. Verrall, A. P. (1943). Fungi associated with certain Shigo, A. L. (1979). Tree decay. An expanded concept. ambrosia beetles. J. Agr. Res. 66, 135-144. U.S. Dept. Agric. Forest Service Agric. In! Bull. No. 419, 73 pp. Webb, S. (1945). Australian ambrosia fungi. Proc. Roy. Soc. Vict. 51, (N.S.) 57-80. Tamblyn, N. (1937). Decay in timber with special Wilson, C. L. (1959). The Columbian timber beetle reference to jarrah (Eucalyptus rnarginata Srn.). and associated fungi in white oak. Forest Sci. 5, 114- Aust. For. 2, 6-13. 127. Tamblyn, N., and da Costa, E. W. B. (1958). A simple technique for producing fruit bodies of wood Yurchenko, J. A., and Warren, G. H. (1961). A destroying Basidiomycetes. Nature 181, 578-579. laboratory procedure for the cultivation and Towers, G. H. N. (1976). Secondary metabolites fructification of species of Hericium. Mycologia 53, derived through the shikimate-chorismate pathway. 566-574. 23 APPENDIX 1 Forester's Classification of Eucalyptus obJiqua Test Logs Cut at Banshea State Forest LOG No. LENGTH GIRTH FAULTS FAULTS FAULTS (cm) (cm) (BUTT) (TOP) (LENGTH) 1 488 158 Moderate Blk.* Heavy Blk. 2 549 137 Yz Dia. Blk. 12/16 Heavy Blk. 3 641 132 Heavy Black '-, 8 Points Brittle Heavy Blk. I1 4 488 170 16/23 Moderate 14/19 Heavy Blk. I, 5 641 160 Heavy Blk. Heavy Black I, 6 366 86 Moderate Blk. Clear 10 point Occ. I Limb 7 427 127 Brittle Heart Brittle Heart 5 pts. 5 pts. 8 366 127 Clear Heavy Black 9 732 92 Gum 1 per 3 Gum 1 per 3 10 427 153 Moderate Blk. Heavy Blk. 11 366 168 Slight Black Heavy Blk. No Registration ' 12 366 86 Clear 13 366 163 Slight Blk. Clear 1-? Gum Bllmp No Registration 14 488 137 Outer Gum Outer Gum 1 Occ. Limb 15 pts. 15 549 153 Clear Brittle Gum 1 per 2" 16 305 165 Clear Heavy Black 17 427 158 Gum 1 per 3 Moderate Black Epicormics 1 Scar. 12" 18 427 221 Heavy Black Heavy Blk. 1 Slight Swell (Excess) (Excess) (Not accounted for) 19 366 188 Ants 2W' Heavy Black Spiral Grain (Excess) Gum Bumps 20 549 153 Brittle 15 pts. Brittle 5 pts. Scar 15 pts. Gum 3 pts. Limb 5 pts. Epicormics 21 549 125 Black Excess Black (Excess) 1 Limb Ants 30 Epicormics 22 793 130 Brittle Brittle 2 Limbs 30 & 10 8 pts. Epics Top 10' 23 549 122 Black Brittle Clean Black Gum 24 366 275 Black Rot 15 pts. Clear Brittle 25 366 231 Black Black Green Epics. *Blk = Black stain or pencilling Occ = Occluded Ants = Termites 24 APPENDIX 2 Culture Media 1. Sawdust medium for sporophore production Sawdust (Pinus radiata D. Don sapwood) Sawdust (Brachychiton acerifolium F. Muel!. sapwood) 360 g Maize meal 240 g Bone meal 15 g Potato starch 4.5 g Dried yeast 1.2 g Malt extract 3.0 g I'~ Casein hydrolysate 1.05 g Tryptone 1.0 g Thiamine hydrochloride 0.02 g Tap water 720 ml 2. Malt/Yeast extract agar Malt extract 109 Glucose 20 g Yeast extract ("Vegemite") 10 g Agar ("Oxoid") 25 g Water 1000 ml 3. Oxidase tests for wood decays • Nobles gum guaiac solution Gum guaiac 0.5 g 95% Alcohol 30 ml • Bavendamn oxidase test agar for white and brown rots Malt extract 5g Agar (Oxoid) 20 g Gallic or tannic acid 5g Water 1000 ml I 4. Stains • Lactophenol aniline blue for detection offungi in wood Phenol 20 g Lactic acid 16 ml Glycerol 31 ml Water 20 ml Aniline blue 0.05 % 25 APPENDIX 3 Cultural and other characteristics of basidiomycetes isolated from Eucalyptus spp. Character 511 518 705 2841 3047 3417 1. Cultural appearance Advancing zone Even + + + + + + Bayed + Raised + + + + ~ Appressed + Growth rate Slow + Moderate + + Fast + + + Colony appearance Cottony + + + + + Downy + + + Felty + Floccose + Plumose + + Silky + Subfelty + Woolly + + + Chamois-like + Velvety + 2. Hyphal features Hyphae Contorted + Encrusted + Fibre-like + Helicoid + + Interlocking + + Clamps + + + + Multiple Clamps + Monomitic Dimitic + + + + Trimitic + + Hyaline + + + + + + Coloured + + Contorted/ irregular + + Contorted/ encrusted + Spores Conidia + + + Oidia + +? Chlamydospores + + + + Terminal + + + + Intercalary + 26 Character 511 518 705 2841 3047 3417 3. Biochemical Crystals present + + + + + Tannic Acid + + + + + Gallic Acid + + + + U.V. Melzers reagent 4. Odour 511 Sweet 518 Sour to citrus 'I 705 Nil 2841 Mushroom-like 3047 Banana-like 3417 Nil 5. Colour Edge Centre Reverse 511 White White Pale cream 518 Pale straw Pale yellow 705 " White Pale cream 2841 " 3047 " Pale yellow" " 3417 " Pale brown (saya1 zones) Dark "brown " 6. Pigments in media 3417 brown 7. Species from which isolated 511 E. obliqua 518 705 " 2841 E. pilularis" 3047 3417 " " 27
Pencil Streak, Stain & Decay in Eucalyptus & Syncarpia In
