PNGAF MAG ISSUE # 9 B - 5B4D3 Dr John Davidson Accompaniment "RAINBOW EUCALYPT MAN" Part 2 of 8.

RAINBOW EUCALYPT MAN A Parallel Journey John Davidson (Part 2)

CONTENTS PART 1 1967

2

Botanical exploration

2

Preparation of a paper for the Ninth Commonwealth Forestry Conference

29

Natural regeneration of E. deglupta

30

Artificial regeneration of E. deglupta from seed

32

Grafting E. deglupta

33

Growing cuttings of E. deglupta

37

Confirmation of E. deglupta as a major native tree species for continuing study

44

I enrol in a Master of Forestry Degree at the ANU

46

PART 2 1968

47

Ninth Commonwealth Forestry Conference

47

Duty travel to Canberra and ANU for initial wood properties examination on E. deglupta

47

Density of the wood of E. deglupta

48

Structure of the wood of E. deglupta

51

Fibre length

51

Statistical design required for the proposed tree improvement programme for E. deglupta

56

Continuation of research on E. deglupta at Keravat

57

Planting trials of E. deglupta

58

Preparation of a paper for a Conference of the Institute of Foresters of Australia

59

Selection and assessment of “candidate” and “breeding” populations of E. deglupta

59

Grafting and cuttings of E. deglupta

60

First visit to Keravat by Professor Pryor of the Department of Botany ANU

62

Helicopter reconnaissance of the Gazelle Peninsula and Central New Britain

66

Flowering and fruiting studies on E. deglupta

69

Emasculation technique developed for flowers of E. deglupta

71

Vegetative propagation of E. deglupta using epicormic and coppice shoots

71

Second visit to the Territory by Professor Pryor of the Department of Botany ANU

75

Wood density variation in E. deglupta

85

Silvicultural Research Conference Bulolo 26 to 30 August 1968

87 ii

PART 3 1968

96

Writing up a report on the fibre length and density work to date

96

Ordering of a rifle for seed and scion collection

96

Experimental proposals

96

E. deglupta fertilizer trial

98

Professor Pryor visits Mindanao in the Philippines to examine E. deglupta

98

Vegetative propagation of epicormic and coppice shoots of E. deglupta

99

Lopping and tying down branches of E. deglupta to control height growth

99

Establishment of a trial grafted seed orchard for E. deglupta

100

Ordering of a machine for removing wood samples from standing trees

102

Arrival of rifle for collection of seed and vegetative material from tall trees

103

1969

105

Collection of seed and vegetative material of E. deglupta

105

Grafting and cuttings experiments

105

Move to Canberra to take up PhD studies at the ANU

105

Aims of the experimental work to be undertaken during my PhD

104

FAO – Forestry and Timber Bureau E. deglupta collecting expedition to Indonesia

106

Fibre wall thickness and lumen diameter

107

Arithmetic ratios of fibre length, wall thickness and lumen diameter

108

Cross-sectional area of fibres, fibre wall material and lumens

109

Arithmetic ratios of fibre cross-sectional dimensions

109

Percentage of tissue types by volume

109

Number of vessels per unit area of cross-section

110

Mean cross-sectional area of vessels

110

Co-authored paper for 2nd World Consultation on Forest Tree Breeding

110

Formal transfer to a Commonwealth Postgraduate Scholarship

111

Chemistry of the wood of E. deglupta

112

End of wood parameter measurement

113

Associations among the primary and derived variables

114

The association between wood density and other measured wood parameters

116

Baku Forest Station

117

1970

117

Wood sampling in standing trees at Keravat

117 iii

Growing plants in controlled environments

119

Experimental methods for growing plants of E. deglupta in the phytotron

120

Provenance studies in the phytotron

122

Fieldwork in Keravat

123

1971

126

Species trial at Baku, Gogol

126

Membership of Forest Research Working Group No 1 Forest Genetics

127

Wood density among random and candidate populations of trees

127

Environmental and genetic variation in wood density

127

Heritability of wood density in E. deglupta

129

Selection differential

130

Selection of propagation population trees

131

Establishment of provenance trials of E. deglupta in Keravat, Dami and Gogol

132

Investigation of heart rot in E. deglupta

133

Volume tables for E. deglupta

135

1972

139

Site Quality computation for E. deglupta

139

Growth and Yield for E. deglupta

139

Completion of my PhD studies at ANU

141

Bridging Report for Silvicultural Research in PNG

142

E. deglupta seed orchard establishment in Bulolo

142

Hybrid eucalypts

145

Hybridization of E. deglupta and E. “decaisneana”

147

Provenance trials of E. “decaisneana” at Bulolo

148

Simulated pulpwood logging at Baku

151

Confirmation of the award of PhD for my work on E. deglupta

153

First species trial at Baku at age two years

154

PART 4 1972 continued

161

Exploration of E. deglupta in the Garaina area, Morobe Province

161

Latest (1972), more detailed description of E. deglupta

164

Second provenance trial of E. deglupta at Baku

168

Reconnaissance on New Britain

170 iv

PART 5 1973

183

JANT start up in the Gogol 1973

183

Membership of Appita

187

JANT harvesting and reforestation activities

187

E. deglupta spacing trial, Baku

190

Thinning trials Keravat 1973

192

Heartwood decay in E. deglupta

192

Continuing harvesting plots at Keravat for pulpwood volume table compilation

194

Overseas duty travel to east Africa and New Zealand

195

Travel Bulolo PNG to Johannesburg, South Africa

196

IUFRO Division 5 Forest Products meeting 24 September to 12 October 1973, Cape Town and Pretoria, South Africa

198

PART 6 1973 continued

203

Rhodesia

203

Malawi

203

Zambia

207

Kenya

208

Back to Australia and on to New Zealand

210

Recreation leave in Australia from 18 November 1973 to 3 January 1974

212

1974

212

Visit by Christian Cossalter of the Centre Technique Forestier Tropical – Congo

212

Visit to Bulolo by Edgard Campinhos Jr of Aracruz Florestal S A

221

Papua New Guinea Tropical Forestry Research Note series

221

Assessment of E. deglupta provenance trial No 2, Keravat, New Britain in 1974 at age 2 years 223 Second provenance trial of E. “decaisneana” at Bulolo

226

July – Oct 1974: Vegetative propagation of epicormic and coppice shoots of E. deglupta

229

Bole shape of E. deglupta

234

Provenances

235

Initial spacing or initial stocking

235 v

Site quality

236

Tree age and average size

237

Practical implications of bole shape

238

An updated general volume table for E. deglupta

241

Further provenance trial of E. “urophylla”

241

Forest Tree Series Leaflet on E. deglupta

243

Progress report on tree introduction and improvement

244

1975

244

Provenance seed collection of E. deglupta in the Celebes and Ceram Islands May 1975

244

Provenance seed collection of E. deglupta in Irian Jaya, Indonesia, 3–17 June 1975

247

Natural distribution of E. deglupta

261

Taxonomy of E. deglupta

268

Taxonomic characters

270

Measurements on E. deglupta leaves preserved as herbarium specimens

272

Measurements on leaves from a provenance trial at Keravat

277

Single taxon at the level of species for E. deglupta

281

Reprinting of my PhD thesis

282

Permission to publish my work on E. deglupta

284

PART 7 1976

285

E. deglupta progeny trial two years old, Kunjingini

285

XVI IUFRO World Congress, Oslo, Norway

287

1977

289

FAO/IUFRO Third World Consultation on Forest Tree Breeding

289

Post Consultation tour to Coffs Harbour

291

IUFRO Workshop, Brisbane

293

1978

293

Visit to Ulamona to investigate damage to a plantation of E. deglupta owned by the Catholic Mission and impacted by a recent volcanic eruption

293

Industrial Timber Corporation of Indonesia

300

Paper Industries Corporation of the Philippines

302

Jari Brazil

306 vi

Seed production from the E. deglupta seed orchard, Bulolo

308

1979

310

Conference on Forest Land Assessment and Management for Sustainable Uses

310

1980

310

Visiting Scientist at the CSIRO Division of Forest Research, Canberra

310

IUFRO Symposium and Workshop on Genetic Improvement and Productivity of Fast Growing Tree Species, Brazil

311

Case studies on forest and watershed development in Asia and the Pacific

325

Reprint of Forest Tree Leaflets

326

1983

326

Publication of the papers given at the IUFRO Symposium on Genetic Improvement and Productivity of Fast Growing Tree Species in Brazil, August 1980

326

IUFRO meeting on frost-resistant eucalypts

327

1984

328

Award of the Marcus Wallenberg Prize to the research team at Aracruz working on Vegetative propagation

328

PICOP at its peak in 1984

329

1986

329

Data book on endangered tree species

329

1993

330

PICOP revisited in 1993

330

PART 8 1993 continued

333

UNDP/FAO Regional Project – FORTIP

333

Publication of Eucalypt Domestication and Breeding

333

1994

334

Second edition of Eucalypt Domestication and Breeding

334

1995

335

Genetic variation in height growth and leaf colour in E. deglupta

335

1997

336

Pulping and papermaking potential of plantation-grown E. deglupta from PNG

336 vii

2003

336

Molecular studies on Eucalyptus subgenus Minutifructus

336

2007

342

Breeding programme for E. deglupta in the Solomon Islands

342

2008

342

E. deglupta Seed Orchard Bulolo

342

2011

342

Herbarium collections of E. deglupta by K Damas

342

Open Bay Timber Company

344

2013

345

E. deglupta chloroplast genome sequenced

345

ACIAR Project “Facilitating the availability and use of improved germplasm for forestry and agroforestry in Papua New Guinea”

347

2014

348

Reference genome of E. grandis released

348

2015

349

Article on E. deglupta in “The Forester”

349

2017

350

E. deglupta in the Adelaide Botanic Garden

350

2018

350

E. deglupta in “Trees for Life in Oceania”

350

2019

353

Our book on “Eucalypt Domestication and Breeding”

353

Geological time line with reference to some eucalypts

354

Open Bay Timber Plantations

359

Keravat 2019

360

ABBREVIATIONS AND ACRONYMS

361

viii

1968

Sampling underway in the 1948 plantation of E. deglupta. Felled trees are visible just inside the stand in the bottom right hand corner of the photograph.

Ninth Commonwealth Forestry Conference The Ninth Commonwealth Forestry Conference was held in New Delhi India 3 – 27 January 1968. Sixty delegates from 15 overseas countries and 97 delegates from India attended the Conference.

There were representatives from FAO and Nepal and 20 other invitees. I did not attend in person. Alan Cameron had prepared a paper on Teak1, to accompany my paper on Kamarere (left). Don McIntosh and Eric Hammermaster submitted a joint paper with the title “Forest Resource Assessment Using Helicopter Transport”.

One thousand copies of each paper were printed by the Government Printer in Port Moresby in December 1967 and a proportion sent on to the organisers in India prior to the Conference and the remainder distributed widely. This paper on Kamarere was my first published work in my forestry career.

Duty travel to Canberra and ANU for initial wood properties examination on E. deglupta Gloria and I left Keravat for Canberra early on Thursday morning 4 January 1968, arriving in Canberra on Friday evening 6 January. As I was a Public Servant we were accommodated in the then Acton Commonwealth Hostel. Meals were taken at small round tables seating up to four persons. At our

1

Cameron A L 1968 Forest Tree Improvement in New Guinea I. Teak. Prepared for the Ninth Commonwealth Forestry Conference, 1968. Department of Forests, Port Moresby.

47

assigned table for the duration was the most senior female police officer at the time in the Commonwealth Police Force.2 My enrolment in the external Master of Forestry Degree course gave me immediate on-site access to the facilities of the ANU. But first I had to learn to use Fortran IVG the compiled high level programming language used by the ANU’s IBM System 360 Model 50 Computer that had just been installed in the University’s Computer Centre. This small mainframe computer released by IBM in August 1965 at the time had a core memory of only 256K! I also had to learn how to use the standard 80-column Fortran Statement and Data punch cards for running jobs on the computer.

Density of the wood of E. deglupta The wood samples had arrived by air from Keravat. Part of each of the large samples was thinned down to produce cambium to pith samples to the precise thickness dimension of 6.9 mm in the longitudinal direction for x-raying. This was carried out using a Black and Decker router modified in the Forestry Department engineering workshop to work as a spindle moulder with a tungsten carbide tipped cutter.

Before and after thicknessing, the wood samples were conditioned in a closed container over a saturated solution of sodium dichromate, which had a relatively constant aqueous vapour pressure at ambient temperatures leading to an equilibrium moisture content in the wood of 8±0.5%. Wood kept indoors in Canberra also was normally close to 8% equilibrium moisture content so there was virtually no change in the wood sample thickness during the machining operation or later during the short time the samples were removed from the sealed container to be placed on the x-ray film and exposed.

X-rays at first were taken in a lead-lined room at the Forestry School in Yarralumla because the new Forestry building at the ANU was not finished.

Full details on the measurement of wood density in E. deglupta have been presented in my PhD thesis.3 The unanalyzed hard-copy graphical output from the microdensitometer was carried back to Keravat for analysis later, to free up time for other work, given the limited time available in Canberra.

2

The Commonwealth Police and Australian Capital Territory Police were separate until 1979 when they were merged to become the Australian Federal Police.

3 Chapter 5, pages 87 – 106 In Davidson J 1972 Variation, Association and Inheritance of Morphological and Wood Characters in an Improvement Programme for Eucalyptus deglupta Blume. A thesis submitted in partial fulfilment of the requirements for the Degree of Doctor of Philosophy in the Australian National University, Canberra.

48

Typical block of wood about 10 cm thick as received from Keravat. It had been cut with a chainsaw from a full cross-sectional disk at Keravat to reduce its weight, while still retaining a substantial sample from bark to pith (nearest camera), originally oriented in the North cardinal direction (or four samples from the four cardinal directions for the breast height disk).

Modified router used for preparing bark to pith samples of a precise thickness in the longitudinal direction (6.9 mm for E. deglupta). Note: For the photograph, the semicircular safety guard has been detached and moved to the right to show the tungsten carbide tipped cutter. The sample was held tangentially in the long clamp on the left, which travelled past the cutter from left to right, precisely guided by the rail at the rear. The sample was machined on both the upper and lower sides with respect to the original orientation in the living tree. Thickness was checked after each pass using a G-shaped micrometer screw gauge.

49

Right: Operating the Joyce Loebl double-beam recording microdensitometer scanning the x-ray films of wood samples on a moving table at the front to determine the density across the image and recording this scan on graph paper on a second oppositely moving table at the rear.

A positive black and white photographic print taken from an x-ray film of the wood sample for “KT5” (Kamarere Tree 5) at “B” (Breast Height). The sample is from one North oriented horizontal radius with the sapwood (“S”) direction to the left and the pith (“P”) direction to the right, but has been cut in half to fit on the x-ray film. Here the cambium is at the left hand end of the top image and the pith is at the right hand tip of the lower image. The darker the tone on this print, the higher is the wood density, whereas on the x-ray film from which this print was made lighter less exposed areas indicated higher wood density. “Growth rings” are evident, but in E. deglupta these are not seasonal or annual growth rings in the usual sense but rather indicate the tree’s varying rate of growth in response to the prevailing environmental conditions (for example, soil moisture) at the time the wood was being laid down. The image here is about natural size (tree radius 27 cm, diameter about 54 cm, at breast height).

50

Structure of the wood of E. deglupta The physical and mechanical properties of wood are related to the composition and arrangement of the various types of tissues and the shape, size, wall thickness and structure of the cellular components. An early description of the wood of E. deglupta was in the form of a card key based on features that could be seen on a transverse surface with a 10X hand lens.

Compared to wood of most other eucalypts, that of E. deglupta has a relatively complicated structure with considerable diversity in types and spatial arrangement of cells. The macro arrangement is fairly constant and is used in the description of the wood of this species (see card key and photographs on the next two pages). However, at the microscopic level, the dimensions, shape, and relative proportions of component cells and tissues may vary, leading to a range of physical and mechanical properties within the species. In E. deglupta preliminary investigation revealed much of this variation may be random but systematic trends existed in some features depending on from where in the tree the observation was made. If the wood properties of an individual tree had to be characterised by a single or low number of relatively small dimension samples, this systematic variation would have a bearing on the representativeness of the sample or samples to characterize the whole tree. This aspect would be especially relevant in non-destructive sampling to determine the wood properties of individual trees in a tree-breeding programme for E. deglupta.

Fibre length A number of investigations with other tree species had indicated that cell length was genetically controlled and known to be a factor affecting properties and utilization of both wood and pulp.4 In the wood of E. deglupta the libriform5 or true fibres are short (0.6 to 1.4 mm approximately), minute in cross-section (10 to 15 μ approximately) and systematically arranged in files only in the radial direction when viewed on the transverse surface.

4

Back then two excellent reviews had been published: Dinwoodie J M 1961 Tracheid and fibre length in timber – review of the literature.

Forestry 34 (2):125 – 144; and Dinwoodie J M 1965 The relationship between fibre morphology and paper properties; a review of the literature. TAPPI 48(8):440 – 447. However, much of the work reviewed in those publications had been carried out on softwoods.

Insufficient data were available at that time on hardwoods, and especially on eucalypts.

5 “Libriform” fibres are greatly elongated wood (xylem) cells with tapered ends, and with relatively thick walls that usually have small relatively simple pits.

51

This card for E. deglupta was included in CSIRO’s 1958 Card Sorting Key for the Timbers of New Guinea and Neighbouring Islands. The key was prepared in a small number of sets using a printed base card format with details laboriously typed on each card individually, and all were hand-punched with reference to a master set. This card had a red bar against “Highlands” indicating a lack of surety, and anyway the card format did not have a category called “Lowlands” which had to be separately noted on the rear, because this species is predominantly found in lowlands. Probably also “Foothills” and “Coastal” should have been punched, also No 33, since Ceram and Sulawesi would fall under the category “ETC”.

52

The wood of E. deglupta, showing the diversity in types and arrangement of cells. A, Transverse section; B, tangential longitudinal section; C, radial longitudinal section; all approximately x 30. D, Macerated wood showing the numerous true libriform fibres and the other less numerous individual elements x 60: f = libriform fibre, v = vessel, r = uniseriate ray, vp = vertical parenchyma, cp = chambered parenchyma, some cells with crystals, ft = fibre tracheid.

53

Interspersed among the fibres are thin-walled parenchyma cells, chambered parenchyma cells and fibre tracheids, the latter not unexpected in this “primitive”6 eucalypt. The remaining significant proportion of the wood volume in E. deglupta is occupied by the vessels, which vary in shape, size and distribution pattern. All these vertically oriented elements are interrupted by the transverse rays, which also usually comprise a significant proportion of the wood volume.

For the investigation of fibre length in E. deglupta, wood maceration involved taking small chips from each the designated sample points (9 heights and several intervals from pith to bark at each height in each of 9 trees, totaling some 450 samples overall) and heating them in a 1:1 (v/v) mixture of glacial acetic acid and 100 volume hydrogen peroxide at 95°C for four hours in a water bath. The corrosive digestion mixture was flushed from the delignified chip sample with water and final fibre separation achieved by vigorous shaking of the sample in water. Microscope slides were prepared by pouring the fibre suspension over the slide under water and then allowing the water to drain off leaving fibres settled on the slide in a random fashion. For viewing under the microscope a temporary mount was made by applying a small drop of glycerine and a glass coverslip.

Left: Gillet and Sibert Conference Projecting Microscope. Magnified images of the fibres of E. deglupta were projected from the slide on the microscope stage on to the paper on the desktop to the right of the microscope at a known magnification, where their length was measured on the image in batches of 25 with a mapping wheel.

6

The first or “primitive” eucalypts (‘proto-Eucalyptus’) were probably trees of the tropical rainforest or rainforest margins. E. deglupta has many features of both a proto-eucalypt and some rainforest trees and appears to have originated from an ancestral proto-eucalypt stock in the rainforests of Antarctic Gondwana. Fibre-tracheids are tracheids having pointed ends, a relatively thick wall, and a narrow lumen as in a fibre, but with small bordered pits, probably functioning in support rather than conduction, and normally the major component of the wood of gymnosperms (softwoods like pines), which evolved before the angiosperms (hardwoods like eucalypts).

54

Fibre images were projected on to a sheet of white paper on the adjacent desktop at 60 – 100 magnifications using a Gillet and Sibert Conference Projecting Microscope. Only whole undamaged true fibres were measured successively in four groups of 25 (100 per sample), with a mapping wheel.7 Full details on the measurement of fibre length in E. deglupta have been presented in my PhD thesis.8 There are shorter and longer fibres present that contribute collectively to the whole tree weighted average fibre length. The very short fibres would be lost in the pulp screening process but the longer fibres would help the bonding qualities of the paper web. A whole tree average of 1 mm or more is satisfactory for pulp production from short-fibred eucalypts.

What was important was the difference in fibre length between breast height and 30% height and at heights closer to the ground and between 2.5 cms from the pith and 15 cms from the pith. This within tree variation affects the decision on where to extract a limited sample from the tree (sample representativeness). Whereabouts must the sample be taken to give the best estimate of weighted whole tree fibre length?9 In E. deglupta the best height to obtain a single representative value for weighted whole tree fibre length was at 30% of total tree height.

Weighted fibre length in the breast height sample was only weakly correlated with weighted whole tree fibre length at 30% of total height (correlation coefficient R = 0.39 and not significant). This was disappointing as breast height is a most convenient sampling position. To obtain a correlation significant at the 1% level or better would have necessitated sampling at ground level where there would be difficulties operating the sampling machine and where buttressing might be an issue, or, to sample at or above 20% of tree height. This would represent about 30 – 35 feet (about 9 – 11 m) above ground on the trees from which selection in the proposed tree breeding programme would be made. Operating the proposed heavy chainsaw-like mortising machine10 that far above the ground to take a wood sample would be difficult and probably too hazardous for the operator(s).

In addition, to produce a long-fibred pulp which would be comparable to that made from a softwood would require a much greater relative increase in fibre length than was currently being achieved in softwood breeding programmes. With E. deglupta I believed that no marked genetic improvement in fibre length could be achieved, as the natural variation in whole tree value from tree to tree was too 7

Preliminary investigations had determined that 100 fibres measured in two groups of 25 on each of two slides representing each sample point would result in less than a 3% error at the 95% confidence level.

8 Pages 38 – 44 In Davidson J 1972 loc. cit.

9

Sample representativeness is discussed in detail on pages 140 to 144 and fibre length on pages 144 to 146 In Davidson J 1972 loc. cit.

10

As described In Sulc V 1967 The extraction of wood samples from living trees. Journal of Forestry 65(11):804 -806.

55

narrow (1.08, 1.12, 1.03, 1.08, 0.99, 1.05, 1.02, 1.07, 1.16 mm for trees KT1 – 9 a range of 0.99 to 1.16 mm). Thus the combination of a non-representative breast height sample and the lack of any possibility of significant improvement of fibre length through breeding caused me to relegate selection of individual trees based on fibre length to a secondary role.

Of the nine trees in the preliminary investigation, and if they had been candidates for selection for a breeding programme (they were not), I would have been inclined to reject only the one tree that had a weighted whole tree fibre length of slightly less than 1 mm (0.99 mm).

Statistical design required for the proposed tree improvement programme for E. deglupta While in Canberra the opportunity was taken to discuss with several experts the statistical design required for selection of superior trees in the Keravat plantations for breeding purposes.

The outcome was that three populations would be required to be identified and assessed: 1.

A random population of 125 or more trees representing the routine unimproved plantation trees involved at Keravat. This would be the lowest level in the hierarchy of populations. Also called a base population by some workers. It would serve also for gene conservation, storing genes of the unimproved population which may be lost during later generations of tree breeding, but which may be required to fall back on in later if breeding is required for a different purpose. It could be a suitable backcross source destination to reverse any severe inbreeding depression that might develop in succeeding generations of the breeding population.

2.

A candidate tree population of 150 to 200 trees for the current tree improvement programme based (i) on external morphological appearance of a number of characters of relevance to selection of superior individuals (including bole straightness, bole cylindricity, minimal buttressing and bole fluting), and (ii) on superior growth based on total height and diameter at breast height, determined by comparison with total height and diameter at breast height of the 5 nearest neighbor trees.

3.

A breeding population of not less than 150 trees resulting from selection of the best trees from among the candidates in regard to rate of growth and external morphological appearance.

No. 1 was also used to establish that individual stand and tree variables in the Keravat plantations were normally distributed and not skewed and to serve as a reference to which the selected populations could be compared. A program called SCREEN was written by me for screening data under pre-set conditions, taking the sub-set satisfying the conditions and calculating several basic statistics such as the mean, median, standard deviation, confidence limits, skewness, kurtosis and the Kolmogorov-Smirnov

56

statistic.11 This was important since statistical procedures for calculating parameters like probability and heritability are based on statistics of the normal distribution. Before leaving Canberra I wrote program CORRE to compute product-moment coefficients between pairs of variables, WEIGHT CORRE to compute weighted disk values and weighted whole tree values and produce a correlation matrix of all combinations of disk values against whole tree values, ARITH CORRE a similar program to the former except for application to arithmetic average values, WEIGHT CORREG similar to WEIGHT CORRE except that an optional regression analysis subroutine was available, and ARITH CORREG similar to ARITH CORRE except that an optional regression analysis subroutine was available.12 Continuation of research on E. deglupta at Keravat We left Canberra at 5.20 PM on Wednesday 3 April 1968 and spent the night in Sydney.

After an early start we left Sydney Airport at 7.00 AM and, via a brief stop in Brisbane, arrived in Port Moresby at 12.00 noon on Thursday 4 April 1968. This was on one of the first round-trip daily daytime flights to operate between Sydney and Port Moresby and serviced by a TAA Boeing 727 (right).

We were accommodated overnight in the Hotel Papua. From 8.00 AM to 11.30 AM the next day Friday 5 April I spent in a briefing session with Alan Cameron and Kevin White at the Department of Forests Headquarters in Konedobu. Then Kevin invited us to have an early lunch with him at the Hotel Papua, before a rush to the airport to depart at 1.30 PM via Lae for Rabaul, where we arrived at 5 PM. We were met and taken by road to Keravat, arriving home at 6 PM. From 7.45 AM on Monday 8 April 1968 we were back at work in our respective offices.

It took nearly two days to inspect the nursery, bring the cuttings equipment back into service, attend to a pile of correspondence and fill in the compulsory blue FD9s and green boot allowance13 forms.

11

In statistics, the Kolmogorov–Smirnov test is a nonparametric test of the equality of continuous, one-dimensional probability distributions that can be used to compare a sample with a reference probability distribution, or to compare two samples. It is named after Andrey Kolmogorov and Nikolai Smirnov.

12 Appendix 5, pages 262 and 263 In Davidson J 1972 loc. cit.

13

The FD9 was a monthly diary with two lines (AM and PM) for each day of the month where an officer filled in the one or two major activities that he had undertaken. The boot allowance form was a monthly form in which field officers (Forestry Officers, Patrol Officers, Agricultural Officers and the like) recorded the number of miles walked each day. The miles earned money to be paid to the officer to buy

57

From Wednesday 10 April the number one priority was calibration of the density curves that had been carried back from Canberra, measuring the area under the curves at intervals along the trace and converting the result to wood density by relating the curve to traces made of wood blocks of known density and acetate wedges that were x-rayed on each plate along with the samples of E. deglupta. This work was fitted in between the other research tasks undertaken at Keravat.14 A positive print of an entire x-ray plate shows the images of five wood blocks of known density (down the right hand side) and two acetate step wedges that were scanned to calibrate the density curves produced by the densitometer of the six wood samples of E. deglupta (in this case bark to pith samples of three trees, each divided into two parts to fit on the size of xray film routinely used).

Planting trials of E. deglupta Planting trial No. 1 was established on Monday 22 April (Treatments A to F) and Tuesday 23 April 1968 (Treatments G to N). There was heavy rain immediately following planting on the Monday and a light shower in the late afternoon on the Tuesday. A site was inspected in Bridge LA for a second planting trial. Planting trial No 1 was assessed for survival on Tuesday 2 July and a report prepared on 3 July (including the Table at left).

The benchmark was “Tubed stock control”, which was stock raised in rigid metal tubes that were removed just before planting. Only the routine stock in plastic tubes with the tubes removed before planting was as successful. Leaving the plastic tube on the seedling at planting time reduced survival. Stock that was too small did not survive as well as normal routine stock. boots. There was a big windfall for Forest Officers on the resource surveys who were walking 10 to 16 miles (16 to 26 km) or more each day out and back along assessment strip lines. (They also received a daily camping allowance, from which was deducted the pooled expenses of running the camp mess and provision of food that was carried into the filed.)

14

Mainly carried out on 11, 18, 23, 24 and 26 April 13, 15, 16, 17, 20, 21and 22 May and 17, 18, 21, 24 June. From 24 to 26 May the first batch of density data was transferred to ANU Fortran coding forms and airmailed to the Department of Forestry ANU for punched card preparation and computing. From 20 to 24 June a second and final batch of density data was entered on coding forms and sent to the ANU for punched card preparation and computing. With data for density in all nine of the sample trees complete, the series of programs that I had written and left in Canberra (see earlier) could be applied. Printouts of the results were received at Keravat by airmail later.

58

Other treatments gave unsatisfactory results or failed. (Note: Peat pots were not included in these early trials.)

Preparation of a paper for a Conference of the Institute of Foresters of Australia.

I prepared a paper with the title “Improving Production from Eucalyptus deglupta in the Territory of Papua New Guinea” for the Fifth Conference of the Institute of Foresters of Australia in Perth Western Australia in 1968. It was typed in Forest Department Headquarters and forwarded from there to the conference organisers. I did not attend in person.

Selection and assessment of “candidate” and “breeding” populations of E. deglupta Research technical staff led by Alan Williams started a systematic examination of all the standing E. deglupta trees at Keravat from Compartment 1 Fryar LA planted 1950-51 to Compartment 15 Fryar LA planted 1955-56 (total = 137 ha). The aim was to eventually select 150 trees for their superiority in growth rate and having as well favourable morphological characteristics judged subjectively (straight bole, none or minimal bole fluting, minimal buttressing, a healthy deep cylindrical crown, small branches close to horizontal, no un-extruded branch stubs to the height of the first green branch).

The first pass identified over 200 trees for possible inclusion without recording any data. These were marked with a single yellow paint band near breast height so that they could be found easily again later.

These trees were the “candidate population” I joined Alan for a second pass in which 150 of the 200 provisionally marked (one yellow band) trees were chosen for written assessment. To improve consistency in the subjective scoring of these trees for bole straightness and bole fluting a card with sample photographs was consulted the field to provide an indication of the magnitude of the score to be applied to what was being observed.

For vigour, each tree assessed had points awarded at the rate of one for every foot (about 0.3 m) of height advantage over the height of the stand in that age class and compartment and similarly one point for each 0.2 inches (about 5 mm) of diameter advantage. The nearest five neighbor trees to the assessed tree provided the benchmark, or alternatively growth plot data if the selected tree(s) occurred within a growth plot. Collectively the growth plot trees and the nearest neighbour trees became the “random population” (can also be called a “base population”, a term preferred by some geneticists).

59

The 150 best trees had a second yellow band applied after they were assessed to indicate that they were then designated as part of the “breeding population”. A metal plate with the candidate number (for example C81) painted on it was nailed to a creosote treated sawn timber stake about 1.5 m tall that was driven into the ground near the base of the tree.

Grafting and cuttings of E. deglupta Top cleft and patch grafts were made at intervals when breaks from other work permitted. New techniques tried were cuttings made from the shoots that had developed from top cleft grafts, the scions of which had been taken from the upper crown of candidate trees by climbers. The aim was to see if grafting had led to any rejuvenation of the original adult material. Cuttings were taken also from trees felled in plots for the purpose of constructing a volume table. This kind of experimentation was recorded as occurring on 29 April, 1, 9, 10, 14, 29, and 30 May. On 14 May the cuttings and grafting scions were collected from Candidate trees 67, 68, 70, 72, 73, 74, 75 and 76 in Compartment 14 Fryar LA using sectional aluminium ladders and climbing. Cuttings were placed under mist and grafts carried out on the same day. For these the first new shoot buds appeared on both the cuttings and grafts on 22 May, that is, after only about one week.

The cuttings plantation of E. deglupta at Vudal was inspected at age 4 months.

Right: Cutting of E. deglupta four months after planting, photographed in early May 1968 in the cuttings plantation in Compartment 2 Vudal.

60

Left: Card used in the field to improve the consistency of subjective assessment and scoring of straightness and fluting of the bole of E. deglupta. Below: Buttresses were measured using a 5-foot (about 1.5 m) long graduated height stick.

Photographs of an example of the two-page assessment sheet that was used in the field for characterizing trees of E. deglupta.

61

The all-cuttings plantation area at Vudal in early May 1968, photographed four months after planting. This plantation resulted from the first mass clonal propagation of E. deglupta by cuttings at Keravat. 1. The arrow indicates a line of one of the better clones that at the same time was represented by a relatively high number of genetically identical saplings. 2. Better growth was evident on portions of the site that were relatively still clear of weeds. 3. Growth was poorer on weedy locations, especially where there was already grass competition at the time of planting. 4. Vudal River with a much older routine plantation of E. deglupta on the left and logged over rainforest on the far bank.

First visit to Keravat by Professor Pryor of the Department of Botany ANU, 31 May – 1 June 1968 After our discussions in Canberra, Professor Lindsay Pryor15 of the Botany Department ANU was keen to inspect forests and plantations of E. deglupta at first hand. It was one of only a couple of species of Eucalyptus that up until that time he had not seen in the field. In Canberra he was posing the question to me that “was it a ‘true eucalypt’ in all its features?” He was eager to examine the systematic position of this species in the taxonomy of eucalypts. Maiden in 1910 had pointed out that it was not easy to establish the affinities of E. deglupta. Maiden noted the anthers were anomalous in that although their shape was “renantherous”, the lobes dehisced by separate slits.16 Blakely in 1934 placed E. deglupta and E. 15

Lindsay Dixon Pryor (26 October 1915 – 17 August 1998) was an Australian botanist noted for work on the taxonomy and vegetative propagation of Eucalyptus. He graduated from the Australian Forestry School with a Diploma in Forestry in 1936 and with the 1935 Schlich Medal. In September 1960 he became the first appointee to the Chair of Botany at the ANU. He retired in 1976, but remained at the ANU in several honorary roles until 1990. He was made an Officer of the Order of Australia in 1983 for services to botany.

16 Maiden J H 1910 A Critical Revision of the Genus Eucalyptus Volume II, Part 2 (Part XII of the whole work):80-81, Government Printer, Sydney.

62

schlechteri with E. raveretiana F Muell and E. brachyandra F Muell in series Myrtiformes of Section Renantherae.17 By 1953 Blake’s uncertainty about E. deglupta was expressed in his statement that the species seemed to be only loosely allied to Australian species of Eucalyptus.18 Gauba and Pryor, in 1958, described a testa (seed coat) structure different from that of the Renantherae.19 Carr and Carr, in 1959, found the calyptra structure also not to be the same as that of the Renantherae.20 All of these observations had been made on dry herbarium specimens where the shape of these organs may have altered during the drying, treatment and preservation processes.

Following my return to Keravat from Canberra, Lindsay Pryor decided to visit Rabaul and Keravat after he had made a scheduled visit to the Solomons. Before Lindsay’s arrival, I obtained some fresh bud and flower material and took it to the LAES where I was permitted to use a microscope with a camera attachment for a detailed examination and take photographs of early flower bud development and morphology of the anthers.

Early flower bud development in E. deglupta. Left: Outer bud bract whorl enclosing inner bud bracts (x15). Right: Developing flower buds inside the whorl of inner bracts, most of the outer bracts having been shed (x 10).

17

Blakely W F 1934 A Key to the Eucalypts, The Workers Trustees, Sydney. Reprinted by the Forestry and Timber Bureau, Canberra, 1955. Section Renantherae later became equivalent to Subgenus Monocalyptus.

18 Blake S T 1953 Studies on Northern Australian Species of Eucalyptus, Australian Journal of Botany 1(2):185-352.

19

Page 27 In Gauba E and Pryor L D 1958 Seed coat anatomy and taxonomy in Eucalyptus. Proceedings of the Linnaean Society of NSW 83:20-31.

20 Carr D J and Carr S G M 1959 Floral morphology and taxonomy of Eucalyptus, Nature 184:1549-1552.

63

Left: Flower bud primordia developing inside the whorl of inner bracts (x 10) Right: Outer calyptra being shed from very young flower buds, leaving scar tissue in a ring around the bud. Inner bracts have also all shed by this stage (x 5). Typically from three to seven of these buds would be retained to form the flower umbellaster in this species.

Anthers of E. deglupta are shown under a microscope at high magnification (x 70). Shape is a rounded oblong, almost earlike when seen from the side. Appearance is a translucent pale yellow. Actual average size is about 0.25 mm wide by 1 mm long. The flexible attachment point near the middle of the back of the anther and an adjacent glossy terminal gland are illustrated at left, and one of the two longitudinal slits that occur along opposite sides of the anther, through which pollen has been shed, is shown on the right. The slits remain separate throughout their length. These images confirm the anthers of E. deglupta are not like the distinctly kidney-shaped examples typical of the Renantherae.

64

Whether a species has one or two calyptra during flower bud development is a diagnostic character used in the classification of eucalypts. E. deglupta definitely has two, an inner and an outer calyptra. The outer calyptra is discarded very early in the life of the developing bud, leaving an annular scar around its circumference.

Anther morphology also is an important diagnostic character in eucalypts. The anthers in E. deglupta are dorsifixed. That is the filament is attached loosely to the back of the anther, such that the anther is free to swivel (also referred to as “versatile”). Longitudinal slits open on each side through which the pollen is shed. The fresh material examined revealed the anthers were not really the definite kidney shape of the Renantherae at all. The slits were mainly parallel to each other on each side of the anther and did not converge to meet at one end as in the Renantherae.

Fresh bark was also examined. The bark of E. deglupta was found to be non-aromatic. That is, it does not contain oil glands and it does not have the typical eucalyptus smell when crushed. The leaves however do have oil glands and the typical eucalypt smell, but the oil content is low.

On Friday afternoon 31 May 1968 Gloria and I travelled by road in to Rabaul to meet Professor Pryor at the Rabaul airport and take him to his Hotel. We stayed in Rabaul overnight and set out for Keravat with Professor Pryor at 7.30 AM on the Saturday.

The whole day was spent at Keravat where Lindsay was briefed on my grafting and cuttings work on E. deglupta and he inspected grafts and cuttings in the Keravat nursery, trees in the 1948 plantation, and plantations of E. deglupta along the Kalabus Road. After lunch with us at our place, Lindsay and I made the trip to Vudal Compartment 2 to inspect the E. deglupta cuttings plantation, then just five months old. Professor Pryor was taken back to Rabaul by road, arriving at 8PM. He spent the night of 1 June 1968 in Rabaul and took the plane out the next morning.

Left: Professor Pryor inspects a cutting of one of the best E. deglupta clones in the, by then, five-month-old clonal cuttings plantation at Vudal on 1 June 1968.

65

Helicopter reconnaissance of the Gazelle Peninsula and Central New Britain As a prelude to seed collection from natural populations of E. deglupta for provenance studies, a helicopter reconnaissance was organized to map the stands then observable from the air on the Gazelle Peninsula and Central New Britain.

An all-day charter was agreed with Helicopter Utilities who had a machine in Rabaul but could only make it available on a weekend in between other work for another client already underway on weekdays.

Drums of fuel were delivered to the grassed area in front of the Davidson’s house, which was the preferred landing place in Keravat. At 7.00 AM on Saturday 9 June 1968, the helicopter with an engineer on board arrived. The fuel was topped up and the first sortie to the limit of endurance on full tanks was undertaken. The engineer remained on the ground near the landing site to refuel the aircraft after each flight and before returning to Rabaul. The sole passenger was myself, equipped with a camera, some out-of-date air photographs and a sketch map of the area showing the main rivers and some of the smaller tributaries on which the distribution of the main observable pure stands of E. deglupta had been marked in the past and now could be could updated to the then present situation.

The first sortie was to fly southeast from Keravat to pick up the headwaters of the Toriu River and follow it to the coast then turn back for Keravat this time following the Asrogi River from its junction with the Toriu north east to its headwaters.

The second sortie was to pick up the headwaters of the Mevelo River and follow it to its mouth in Henry Reid Bay. Also some other streams in that vicinity and another draining into Wide Bay21 were explored, before again returning to Keravat.

The third sortie was to explore the Sai River and other rivers flowing into Open Bay such as the Mavelu.

On arrival back at Keravat at 2 PM it was decided the weather in Central New Britain was deteriorating too much to allow another sortie. The helicopter was refueled at Keravat and it returned to Rabaul with the engineer on board.

21

Called “Spacious Bay” on some old maps and on the labels of some early botanical specimens of E. deglupta.

66

Helicopter reconnaissance 1968. Top: From the air, the ease of separation of E. deglupta with its lighter green coloured canopies (arrows) from the surrounding rainforest vegetation was easiest when the light was overhead near midday and the viewing angle was oblique. Bottom: In the morning light and at close range the emergent canopies were obvious (arrows) and the distinctively coloured trunks stood out when viewed from the side, here helped by the open transect made by a logging road near Open Bay.

67

Helicopter reconnaissance 1968. Top: With the morning light behind the aircraft, the colour difference was less, but the height of the emergent canopies of E. deglupta (arrows) ensured they stood out from the surrounding vegetation. Bottom: In the late seral stage shown here, the rainforest element had almost completely taken over. The slightly lighter and marginally taller canopies of the remaining E. deglupta (arrows) dispersed through the rainforest elements were harder to spot. For this situation, being able to slow the speed over the ground and to descend much lower in a helicopter than in a fixed wing aircraft were particularly helpful.

68

Stands of naturally occurring E. deglupta in the North Bainings – Open Bay – Wide Bay region of New Britain (black areas) confirmed and/or updated during a reconnaissance by helicopter on 9 June 1968.

Flowering and fruiting studies on E. deglupta A planned long-running flowering study was started on 24 June 1968. Several of the marked candidate trees were observed regularly using binoculars to record the development of flowers and fruits over time.

Also the amount of seed harvested from each tree was recorded. The periodicity of significant fruiting

69

events was also noted (some trees at Keravat had at least a few flowers on them all year round). Climbers collected flowering panicles in varying stages of development for detailed study.

This is a summary of the results from these studies: •

90 – 200 panicle clusters per tree on six-year-old trees

•

Pistil22 6 mm long by 4 mm wide

•

Style 2 mm long, lengthening to 2.5 – 3.0 mm at the time the stigma is receptive

•

Anthers oblong, dorsifixed (versatile), 0.25 mm wide x 1.0mm long, dehiscing by longitudinal slits

•

Pollen triplete (triangular) with trilete (tri-radiate) scars, tapering to pointed tips, sticky, adhering in clumps (found on parrots’ beaks and on insects, especially on the “Green Tree Ant”23, among the principal pollination vectors of E. deglupta)

•

Inner calyptra green to light green for 5 – 6 weeks, greenish white for weeks 7 – 8 then straw coloured, indicating it will fall off in a few days to one week

•

Anthesis takes one day from calyptra fall. Filaments

Flowers of E. deglupta: 1. Straw coloured inner calyptra being shed, 2.

Stamens inflexed, about to unfurl, 3.

Filaments spreading, 4. Flowers fully open, stamens pale yellow, all fertile, anthers dehiscing releasing viable pollen, stigmas still not lengthened or receptive. (About actual size.)

spread outwards radially after being reflexed (folded inwards) in the bud, anthers dehisce exposing pollen. Pollen is already mature, as determined by germination tests, but stigma still not receptive at this stage. Stamens after spreading are numerous, prominent, showy and attractive to insects and birds. Anthers begin to turn brown and shrivel 2 - 3 days after they dehisce.

•

Stigma receptivity starts 1 – 2 days after anther anthesis, that is 2 – 3 days after the inner calyptra detaches, and continues for 2 – 3 days

•

Secretion of nectar does not start first from the stigma, though it does become visibly shiny and sticky at the onset of receptivity. Receptivity coincides with the inner walls of the staminal ring beginning to “perspire”, producing minute droplets that eventually fill the cupshaped void up to the level of the rim. Inside and below the staminal ring and above the ovary

22

Term used for the collective female reproductive parts of the flower. In E. deglupta the centrally located pistil has a swollen base (the ovary), containing potential seeds (ovules), a stalk (style) arising from the top of the ovary and a pollen-receptive tip (stigma).

23 The usually red to orange coloured Oecophylla smaragdina is a species of tree-inhabiting ant found in tropical Asia and Australia. These ants form linked colonies with multiple nests in the canopy of one or more trees, each nest being made of live green leaves stitched together using the silk produced by the ant larvae. They were especially prevalent in the Kamarere plantations at Keravat and when disturbed capable of inflicting a painful bite with their fangs because they could inject concentrated formic acid through any broken skin.

70

are also numerous nectaries, which begin to produce a copius nectar flow. The nectar attracts even greater numbers of birds and insects that serve as pollen vectors •

Within a single flower no transfer of viable pollen takes place on to the unreceptive stigma.

As mentioned, the stigma only becomes receptive 2 – 3 days after the stamens have expanded outwards, by then all the pollen has gone or has lost viability. (This protandry appears to be an inherited morphological trait to avoid self-pollination; also fruit set is very low to nonexistent from forced self pollination, indicating at least one other inherited physiological or chemical blocking mechanism is operating)

•

Bagged un-pollinated flowers fall off

•

Capsules per panicle 20 – 200

•

Viable seeds per capsule 5 – 28 (average 15 – 16), plus copius “chaff” (aborted ovules)

•

Valves usually 4, sometimes 3, rare extremes 5 - 9

•

Seeds set in an average of 25% of capsules, extremes of 0 – 50% of capsules.

Emasculation technique developed for flowers of E. deglupta For artificial cross pollination and hybridisation work in the tree breeding programme for E. deglupta it was necessary to devise a method for removing the anthers (emasculation or removal of the male parts) from a flower and leaving the stigma (female part) intact. This turned out easier than expected, since the inner calyptra and the strongly inflexed stamens inside could be removed as a complete unit by running an incision around and just through the capsule wall with a sharp curved scalpel using as a guide the circumferential scar left where the outer calyptra had abscised. This could be timed just before there was a change from a green to a straw colour that indicated the calyptra was about to be shed normally. At this time, and for a couple of days after, the stigma would not develop to a receptive stage. When the stigma had become receptive, artificial pollination could be undertaken by brushing on the donor pollen and the pollinated flower heads bagged in unwoven terylene bags to prevent contamination by unwanted foreign pollen.

Vegetative propagation of E. deglupta using epicormic and coppice shoots Epicormic shoots that grow from the stems of eucalypts have their origin in the dormant bud strands, as has been explained earlier in the section on patch grafting.24 There are more dormant bud strands at the bases of eucalypts that develop lignotubers than at the bases of those that do not have them; however, 24

See also Jacobs M R 1955 Growth Habits of the Eucalypts. Forestry and Timber Bureau, Canberra. 262 pp. (Sections on dormant bud strands and lignotubers in eucalypts.)

71

all eucalypts produce about the same number of dormant bud strands on the trunk and branches and all these strands retain the ability to form epicormic buds throughout the life of the tree.

Buds that develop on the higher reaches of the trunk and branches are termed “epicormic branches” while those arising further down the trunk are called “epicormic shoots” and those arising at or near the root collar are called “coppice shoots” or, simply, “coppice”.

E. deglupta and other eucalypts such as E. regnans, E. diversicolor, E. gigantea, E. fastigata, E. grandis and E. nitens have several characteristics in common. They have very thin, smooth bark, do not develop lignotubers and are very prone to damage by fire. These eucalypts are often called “non-coppicing species” because of the absence or poor development of epicormic shoots following damage by fire or other agents. This reputation has grown up largely because of the lack of detailed information and in most respects is erroneous. If any of these species is defoliated, without injury to the bark, epicormic shoots will develop, although often in a haphazard fashion and sometimes under apparently fortuitous environmental conditions.

There was ample evidence that epicormic shoot development occurred in E. deglupta.

The long raking crowns of the veteran trees at Keravat were composed almost entirely of epicormic branches that had developed following severe damage or loss of the original crown, almost certainly resulting from one or more volcanic eruption events in the case of trees in the vicinity of the Rabaul volcanoes.

Left: A veteran tree near Keravat that has a replacement canopy of epicormic branches.

These replacement branches had developed following severe damage to and possible decapitation of the original canopy, perhaps some decades previously as a result of an eruption or eruptions among the volcanoes in the vicinity of Rabaul.

In the Keravat plantations, large trees (30 – 40 cm diameter, 50 m high), which had been broken off by wind 6 – 10 m above ground, produced numerous groups of epicormic shoots under favourable conditions of humidity and sunlight.

72

Coppice and epicormic shoots in some other eucalypts were considered to have potential for vegetative propagation, as dormant bud strands low down in the bole retain a degree of juvenility and sometimes strike from cuttings, according to discussions I had just had with Professor Pryor during his recent visit to Keravat. It was decided to attempt to strike cuttings taken from epicormic and coppice shoots of E. deglupta.

Epicormic shoots often did not develop on every tree at the one location even though the microenvironment of each stem appeared to be the same (in the absence of any quantitative data for comparison). Thinning of a dense stand of E. deglupta in one location had allowed in enough light to stimulate epicormic shoot development on some trees. Shoots had developed often from cut logs of E. deglupta lying on the forest floor in damp shaded situations. These all died when the logs dried out.

They never attained more than a few centimetres in length.

Epicormic shoots on a 20-year-old plantation tree of E. deglupta from about 1 - 3 metres above the ground. This tree had lost its upper crown in a storm. Several dormant buds from a number of bud strands had developed. These kinds of shoots were used for trials of vegetative propagation by cuttings.

Epicormic shoots developing on the sunny side of the bole of a 20-year-old plantation tree of E. deglupta, which had been decapitated by wind about 5 m above ground level.

73

In one case at Keravat, a pole was cut and immediately erected in the ground to serve as a support for an electric cable. The green bark was left on. Within a few days, several epicormic shoots had developed.

These were short-lived (two weeks) and no callus or roots formed at the base of the pole.

Coppice shoots appeared to occur haphazardly also. In more than twenty plots, each about one-twentieth hectare in several age classes that were felled during volume table compilation, the stumps of only one age class (10 years) produced coppice shoots. The main differences noted among plots were age, time of sampling, site and tree size-class distribution. No special circumstances observed could subjectively explain why only one plot produced coppice. Even within that plot, only some individuals (40 percent) had coppiced. On these individuals coppice shoots varied in number and size, with generally more shoots on stumps of larger diameter (there are more dormant buds on large stumps). Some of the coppice shoots were very short lived, the node of the callus falling out of the bark before becoming firmly fused to the wood of the trunk. Callus was always slow to develop in a centripetal direction and coppice shoots high on the stump were attached most tenuously. Young coppice saplings that reached to three metres in height were commonly attached to the stump over an area of only about 100 cm2. Lack of rigid attachment, poor ability of the callus growth to cover the cut surface of the stump and susceptibility of the stump meanwhile to virulent decay fungi in the moist tropical environment all contributed to reduced longevity of E. deglupta coppice compared to what would be the norm in a “coppicing eucalypt species”.

Coppice shoots developing from the root collar region (just below ground level) of E. deglupta seemed more persistent, because some support was given by the surrounding soil. Cutting the stump very short favoured the development of “ground coppice” in E. deglupta.

On large rootstocks (that is large diameter stumps), coppice of E. deglupta that did survive grew much faster (and straighter) than seedling material of the same calendar age probably because of the contribution of a super-abundance of feeding roots left behind after the original stem was removed.

A small number of single node cuttings was harvested from coppice emerging low down on a cut stump and a number of multiple node tip cuttings were harvested from a cluster of epicormic shoots found on a 20-year old tree about 2 m above the ground. Both kinds produced roots under the procedures then in use at Keravat.

74

Above: Single node cutting from coppice. Right: Multinode cutting from epicormic growth. Both were the same age after striking in vermiculite and sand under mist, with hormone assistance. The less vigorous rooting in the cutting from an epicormic shoot may have been because of the older ontogenetic age of the bud strands 2 m up the trunk. (Both near natural size.)

Second visit to the Territory by Professor Pryor of the Department of Botany ANU On Thursday 8 August 1968, Professor Lindsay Pryor returned to Keravat after having transited Port Moresby and Lae on the previous day and spending the night in Rabaul. On this trip he was funded by the Department of Forests with the aim of visiting and reporting on as many of the accessible natural populations of E. deglupta as possible. I was assigned to accompany him for his time in the Territory.

Most of Thursday and the Friday morning were spent on the banks of the Keravat and Vudal Rivers. In some places wading in the shallows was necessary to collect botanical specimens from a number of different sites. Lindsay and I spent Friday afternoon at the Regional Office in Rabaul for the collection of air tickets and to finalize air bookings and air charters and to collect accommodation warrants for the remainder of the Professor’s travel in the Territory. The Professor spent the night in Rabaul while I returned to Keravat.

The next day Saturday 10 August I collected Lindsay from his hotel in Rabaul and drove him to Keravat where more specimens and photographs were obtained and herbarium material placed in plant presses for drying. Several duplicate specimens were collected destined for the Gauba Herbarium in the Department of Botany ANU and the Herbarium in Lae, and through the latter to selected overseas herbaria.

75

Left: E. deglupta specimen collected from Site 2 Keravat by “L Pryor and J Davidson” and later shown here as deposited in the Herbarium at Lae.

Professor Pryor, Gloria and I travelled in to Rabaul in the evening. We took Lindsay out to dinner at one of Rabaul’s Chinese restaurants, then returned to Keravat while Lindsay spent the night in Rabaul.

On Sunday 11 August, I travelled in to Rabaul to join Lindsay at the airport to board a charter flight to Cape Hoskins. On the way the flight was directed over several natural stands of E. deglupta on the Toriu, Asrogi and Sai Rivers in the Open Bay Timber Area for Lindsay to see how the species typically occurred in the landscape (see next page). In the Hoskins area, a number of sites was visited including at Dami and numerous herbarium samples were collected. On 12 August Lindsay and I flew from Cape Hoskins to Lae on a charter, then on a scheduled flight to Goroka.

Specimen of E. deglupta collected from Site 2 of the “L Pryor and J Davidson” survey at Cape Hoskins on 11 August 1968, seen later mounted and deposited in the Lae Herbarium. Duplicates were sent to several overseas herbaria. Professor Pryor took a set of specimens back to Canberra for depositing in the Gauba Herbarium of the Department of Botany at the ANU.

76

Lindsay Pryor and I flew over several stands of E. deglupta on the Toriu, Asrogi and Sai Rivers on our way from Rabaul to Cape Hoskins in a chartered aircraft, 11 August 1968.

77

Left: Flying into the Highlands in a DC3 aircraft in 1968.

View forward out of the cockpit windows. If the weather became too severe for continuing to fly safely, common practice of the time was to land at the nearest suitable airstrip and for everyone to get out and stand around under the wing of the aircraft until the weather cleared enough to continue on!

Over Goroka in a TAA DC3, while coming in to land on 12 August 1968. Some of the tall street trees in the centre and left side of the photograph at the time were E. deglupta.

On 13 August, and Lindsay and I first turned our attention to an investigation of the E deglupta trees that had been planted along some of the streets in Goroka. We then moved on to the Sing-Sing Grounds in Kerowagi to see, and collect specimens from, the relatively large trees that were growing there.

Several commentators had remarked on these trees of E. deglupta as probably carried in from the north coast as seedlings for trade by visiting tribes in ancient times.25

25

This idea was not supported by the morphological evidence gathered by Lindsay and I on this trip. The morphological affinities were with E. deglupta growing in the Jimmi Valley and other highland valleys, and those in turn with the inland trees visited in Pagei and Ossima further to the west. There is no evidence that the species in the past ever occurred naturally on most of the north coast of the mainland.

Also it is doubtful that any seedlings would have been carried in from the north coast as trade goods since the highlanders did not interact peacefully with the coastal lowlanders there, according to historical accounts.

78

E. deglupta street trees in Goroka in August 1968.

E. deglupta at the Sing-Sing Grounds in Kerowagi in 1968. Other trees present included casuarinas and pines.

79

14 and 15 August were spent travelling by road and bush tracks between Goroka and Mount Hagen to inspect streamside E. deglupta, particularly in the Jimmi River Valley, and in the areas photographed by R G Vial. Collections had been made previously by J S Womersley April 1951 from Nondugl, L J Brass 29 July 1959 from east slopes of Mt Wilhelm, A N Millar 3 February 1964 from Keglsugl (in “moss forest”), and L K Wade 21 October 1966 from Gembogl. Highland streams are high-energy streams compared with the rivers of New Britain. Also their beds comprise relatively large rocks and stones and coarse sandy sediments compared to the fine, mainly pumice-based sediments in New Britain. When in flood the boulders hurtling along in the fast flowing water cause considerable damage to regenerating seedlings and saplings of E. deglupta. At these elevations (up to 2,000 m) the adjacent forest comprised various mixtures of Casuarina, Nothofagus, Castanopsis, Podocarpus, Dacrycarpus, Papuacedrus and Libocedrus. This kind of forest was similar to the Conifer-Nothofagus forest that P van Royen described in the Mt Nettoti region of the Vogelkop Peninsula, Netherlands New Guinea (now West Papua, Indonesia) during his collection of E. deglupta on 30 November 1961.

Specimens of E. deglupta collected from the Sing-Sing Grounds at Kerowagi. Left: One of my specimens for the Herbarium in Lae. Right: Professor Pryor’s specimen lodged in the Gauba Herbarium, Department of Botany, ANU.

80

Travelling anywhere off the main track to visit the highlands streams to inspect occurrences of E. deglupta was fraught with difficulty if one wanted to use a vehicle instead of walking to save time, as Lindsay and I found on several occasions!

The typical scattered occurrence of E. deglupta in the highland river valleys is shown here on the left and on the next page. Highland streams are high-energy streams compared with the rivers of New Britain. Also their beds comprise relatively large rocks and stones and coarse sandy sediments compared to the fine, mainly pumice-based sediments in New Britain. When in flood the boulders hurtling along in the fast flowing water cause considerable damage to regenerating seedlings and saplings of E. deglupta (right hand photograph). So only a few survive to be quickly overcome by the other streamside vegetation.

81

At elevations of 1,800 to 2,000 m, the adjacent forest comprised various mixtures of Casuarina, Nothofagus, Castanopsis, Podocarpus, Dacrycarpus, Papuacedrus and Libocedrus that compete with the few E. deglupta that reach sapling and tree size. There were no bands of streamside regeneration of a number of age cohorts comprising pure E. deglupta as typically found next to the rivers on New Britain. Seed and botanical specimens are being collected here. The locations shown here were near the origin of the Mingende seedlots used in the early provenance trials of E. deglupta.

On 17 August Lindsay and I flew out of Mount Hagen for Lae, then from Lae to Vanimo. Sunday 18 August was a rest day spent in Vanimo. There we were able to gain access to drafts of some of the then recent (July 1968) work done by CSIRO during a land resources survey of the Vanimo area26, which included areas around Pagei and Ossima that were on the schedule to be visited. We learned that, on the proposed Pual Land System in the upper reaches of the Puwani and Bewani Rivers, Casuarina cunninghamia – E. deglupta forest, or its seral stages in which Casuarina became more dominant, was to be found. The largest trees were less than 5 ft (1.5 m) girth but still up to 100 ft (30 m) tall.

26

Much later published as: Löffler E, Haantjens H A, Heyligers P C, Saunders J C and Short K 1972 Land Resources of the Vanimo Area, Papua New Guinea, Land Research Series No. 31, CSIRO, Australia.

82

On Monday 19 Lindsay and I, accompanied by Peter Britton of Vanimo Timbers, boarded an aircraft chartered by the Department of Forests at Vanimo airport. This was a single-engined Cessna and I was surprised when the pilot in the left hand front seat invited Lindsay seated in the right hand front seat to take us off and set course on the way to Pagei airstrip (575 ft (175 m) elevation). The takeoff was quite expertly performed. (I was not aware until then that Lindsay had a private pilot’s licence and regularly flew this model of Cessna in Australia!27) The weather was clear and one could see into the headwater valleys of several Sepik River tributaries en route.

At Pagei, specimens of E. deglupta were collected from a streamside location near the airstrip.

Left: One of my specimens from Pagei deposited in the Herbarium in Lae.

27 “Lindsay founded the school of ‘Aviation Botany’ so that he could check critical species in his classification in person - the more remote a species occurred, the more critical it was for personal observation! During long flights to Central and Northern Australia, Lindsay developed a love for flying, taking over manual control at all opportunities without the help of the autopilot or his co-pilot, usually Dugald Paton, who enjoyed the accompanying commentary on the eucalypt species below. The Professor revelled in the challenge of keeping on track with all instruments “spot on”, using the same attention to detail that he brought to bear in his classification and other publications.

His sense of purpose is illustrated through achievement of his unrestricted pilot’s licence when he was in his mid 50’s. On occasions, Lindsay persuaded his wife Wilma to accompany him on such field trips, and taught her how to hold the plane level and straight while he attended to the navigation chores. Wilma’s recurrent nightmare was the Lindsay would succumb to a stroke and she would be left holding the plane level and straight until it eventually ran out of fuel. Lindsay’s temperament exhibited itself on occasions when flying through rough weather, a glint would come into his eyes, but rarely did he indulge in rough words. His composure under trying conditions was the same as he displayed with verbal turbulence at taxonomy meetings.” Source: Quoted from the Inaugural Lindsay Pryor Memorial Lecture address by Allan Hawke, Chancellor of the Australian National University, Coombs Lecture Theatre, ANU, 26 September 2006.

83

Right: Young E. deglupta adjacent to the school grounds at Ossima. On the left in dark shorts is Peter Britton. On his right is the schoolmaster, with a number of curious school pupils.

The survey party then flew the short distance to land at Ossima airstrip (275 ft (85m) elevation). From there it was a short walk to the local school ground to inspect some adjacent E. deglupta saplings on the bank of a stream. These trees were said to have grown as natural regeneration. The species was reported growing some distance upstream but there was not time to walk to inspect this, and it was not obvious from the air on the flight out towards Vanimo. Before the specimens obtained on this visit, there was only the example collected by C D Sayers from “Ossima Village” in 1964 and located in the Herbarium in Lae.

84

Bottom of previous page and left: Specimens from three different E. deglupta trees at Ossima collected by me on 19 August 1968 and deposited in the Herbarium in Lae. The generally broader leaves of these specimens, and the ones from Pagei, compared to the New Britain sources were noteworthy.

The team flew back to Vanimo to spend the night there. On Tuesday 20 August Lindsay and I travelled to Lae by air. On 21 August we spent the day at the Herbarium in Lae inspecting all the eucalypt specimens held there. There was an opportunity to divide up the large number of specimens of E. deglupta that we had collected between those for the local Herbarium and those to be sent to Canberra for Professor Pryor to lodge in the Gauba Herbarium of the Botany Department at ANU. Extra duplicates, where available, were sent to other herbaria, in the following order: Leiden, Queensland Herbarium, Canberra (now the Australian National Herbarium, CSIRO), Harvard University, Kew, Bogor, Singapore, NSW National Herbarium Sydney, Bishop Museum, the Smithsonian Institution and the Royal Botanic Garden Edinburgh.

Lindsay flew from Lae to Port Moresby on Thursday 22 August and spent Friday in the Department of Forests at Konedobu reporting on our activities. On Thursday 22 August I flew from Lae to Rabaul where I was met by a vehicle from Keravat and taken home. I spent Friday 23 August at Keravat putting finishing touches to my presentation for the Silvicultural Research Conference at Bulolo. While I had been away, the final computer printouts for wood density had arrived in Keravat. This enabled me to add more information on the wood density of E. deglupta to my forthcoming presentation.

Wood density variation in E. deglupta The results showed a phenotypic range of density of over 20% (0.45 – 0.57 gms/cc at 8% moisture content) among the nine Keravat trees. The patterns of variation in density in each of the nine sampled trees were complex.

85

Wood density variation in two of the nine sample trees of E. deglupta (KT 1 and 2, pith to bark in the north cardinal direction). For all nine sample trees, in very general terms, there was an increase in density from pith to bark and an increase from ground level to about 30% of tree height, then a decrease from there to the 70% height level (and probably continuing to decrease above that level). For the nine sample trees KT 1 – 9, weighted whole tree densities were 0.51, 0.51 0.49, 0.54, 0.45, 0.46, 0.56, 0.57, and 0.50 gms/cc respectively. Trees KT 7 – 9 from natural forest, of undetermined age but similar to slightly larger size, had an average density about 9% greater than the 19-year-old plantation trees (KT 1 – 6).

86

There were discrete areas of high and low density.28 The correlations of weighted whole tree density with weighted disk density at ground level and weighted disk density at breast height were not significant. However the weighted disk value for density at 10% height was significantly correlated to weighted whole tree density at the 1% level of probability, indicating that would be the height at which sampling for whole tree wood density would need to take place. Sampling at 10% height was feasible.

Silvicultural Research Conference Bulolo 26 to 30 August 1968.

On Saturday 24 August I flew from Rabaul to Lae, stayed there overnight, then flew to Bulolo on Sunday 25 August. The Silvicultural Research Conference was held at the Forestry School in Bulolo from Monday 26 August to Friday 30 August 1968 inclusive. Most of the senior officers of the Department were in attendance. On the Programme, chaired by K J White, were: Experimental Procedure and Communication Research Co-ordination Mycology and Forest Pathology Fire Protection Exotic Softwoods Forest Hardwoods Genetic Improvement of Forest Trees in New Guinea Forest Tree Improvement – Kamarere Tree Improvement – Hoop and Klinkii Pines

A L Cameron K J White P Wright J K Riley J E N Smith K J White A L Cameron J Davidson L T Clifford

Formal presentations29 were scheduled and interspersed with local field visits.

My presentation on Kamarere was important at the time because it included worked up results from my official Tour of Duty at the ANU in Canberra that had ended on 4 April 1968. I presented a much more detailed account, especially on wood density, over two hours on the day than what was set out in the

28

Another PhD student, Mark Higgs, a year later, found similar patterns for wood density in E. regnans: See Higgs M L 1969 Genetic and environmental factors influencing commercially important wood properties of Eucalyptus regnans, a thesis submitted in partial fulfilment of the requirements for the Degree of Doctor of Philosophy in the Australian National University, Canberra. A later principal components analysis by me indicated the variation in wood of E. deglupta did vary systematically with height and distance from the pith, but this was difficult to visualize at the time from the patterns of two of the nine trees shown here.

29

Kevin White had the Department of Forests publish the Proceedings of this Conference eight years later (in 1976 and priced at K1.50!). He considered the information still had value for a wider circulation, as the original papers, which were made available only to attendees at the Conference, had become hard to source. The paper that was reproduced therein in 1976 as my contribution was written well in advance of the Conference and was similar to one prepared also in 1968 for a Conference of the Institute of Foresters Australia. My actual two-hour presentation in Bulolo in August 1968 was in greater detail and more up to date on the day.

87

paper prepared earlier that was handed out. I included illustrations on overhead transparencies and exhibits such as x-ray films, density traces and wood samples that were passed around the audience.

TREE IMPROVEMENT KAMARERE I. INTRODUCTION The genus Eucalyptus is poorly represented by naturally occurring species in the Territory of Papua and New Guinea. Of the six species present five belong to the dry savannah woodland. Eucalyptus deglupta (Kamarere) is the exception and forms part of the early seral forest in rainforest communities. E. deglupta is unusual in that it does not occur in Australia.

Kamarere is the major species of Eucalyptus planted in New Guinea up to the present time. Some years ago, it was decided that Kamarere might make a suitable pulping species when it was proved by Von Koeppen in 1958 that it would make a good pulp by the sulphate process.

The development of areas such as Vanimo and Wide Bay would require a large-scale reforestation movement on logged over areas to maintain operations on a sustained yield basis. A chip operation depends on a continuous supply of a large volume of wood to be economical. A two-ship export operation would require 120-150 million super feet [about 280 – 354 thousand m3] of wood and some 20-40 thousand acres [about 8 – 16 thousand hectares] of land would be required to sustain an operation of this magnitude.

If we are considering entering into a project of this size it is obvious that we should do our utmost to provide the best possible product for the buyer. This can best be done by a detailed investigation of our present growing stock to find out its good and bad points and to introduce a programme to improve the desirable properties and to eliminate as far as possible the undesirable ones. The actual tree improvement programme was envisaged about 1966 at a time when the establishment of E. deglupta plantations in the Territory was at a low level and mainly experimental.

In Table one, which is attached to your copies of the paper, I have listed the areas of Kamarere that have been planted at Keravat over the past nine years.

TABLE 1 EUCALYPTUS DEGLUPTA PLANTINGS 1959-1968 Year Compartment and Logging Area Station Area Established (acres) 1959/60 1. Little Vudal Keravat 133.5 1960/61 2. Little Vudal Keravat 83.6 1961/62 3. Little Vudal Keravat 25.0 1962/63 4. Little Vudal Keravat 45.5 1963/64 1. Vudal Keravat 1964/65 5. Little Vudal Keravat 25.0 1965/66 4. Kamarere Keravat 56.9 1966/67 5. Kamarere Keravat 5.0 1967/68 6. Kamarere, 2. Vudal Keravat Trial areas* * Planting trials, cuttings, prototype grafted seed orchard and other trials still to be planted in 1968.

Since 1948, when Kamarere was first established in a plantation at Keravat, only some 800 acres [about 325 ha] in total have been planted. This is only a drop in the ocean compared with what we now envisage. It is obvious that, in addition to the genetic programme, we need to take a critical look at the issues associated with the establishment of large areas of Kamarere.

Firstly, I am going to discuss how our genetic programme is coming along and later I’ll take a quick look at the production side of the issue.

II. THE GENETIC PROGRAMME The first step in the development of a genetic programme is to assess the natural variation to be found in plantation stands of the species being investigated.

Alan Cameron made some preliminary investigations of basic density and fibre length on 19 trees 13 years of age in 1966.

In this work samples were taken at a height of 3’ 6’’ [1.07 m], in a north-south direction through the pith. Only a single sample 2½’’ [about 6.4 cm] along the grain and 1’’[about 2.5 cm] across the grain was removed from each tree.

Basic density was found to be fairly low in these young, fast-grown trees. Considerable variation was found among trees, showing promise for selection and breeding.

88

In the preliminary study on fibre length, Alan found it somewhat variable but the variation was fairly small.

The overall pattern of fibre length and density in trees is well known but detailed investigations are essential to enable comparisons of these characters between trees. If one is not certain of the within tree variation of a particular character, the information obtained from a single sample, say from breast height (or from 3’ 6’’ height with Alan’s work) cannot be applied to a selection programme, as we shall see later on.

The most reliable method of comparing even-aged trees is to compare estimates of fibre length and density derived from a number of measurements throughout each tree. This technique requires that a large number of sampling points be scattered throughout each tree and this high sampling intensity normally means cutting the tree down. In practical selection, sampling must be non-destructive so we must restrict the number and size of samples taken from each tree. How are we going to manage this?

We can overcome the problem by making a detailed examination of a small number of trees to find out the natural pattern of the character within the tree then use this knowledge to predict whole tree values from a limited sample. Also, where this limited sample is located within the tree becomes of great importance.

I have made a detailed assessment of fibre length and density in nine Kamarere from Keravat. Six of these were from a 19year-old plantation and three from a natural stand of unknown age near the Keravat River.

Each tree was cut down and a disc removed from ground level, breast height, 10%, 20%, 30%, 40%, 50% and 70% of total tree height. Pith to bark sections of these discs were sent to Canberra. At the ANU small sub-samples of wood were taken at 5, 25, 45, 65, 85 and 95% distance from the pith at each height level. These were placed in a 1:1 mixture of glacial acetic acid and 100 volume hydrogen peroxide and cooked for 4½ hours to obtain fibre separation. The fibres were mounted on glass slides and measured from the image produced by a projection microscope at 80-120 magnifications depending on the average length of the fibres.

XXXXXXXXXXXXXXX TRANSPARENCY – FIBRE LENGTH XXXXXXXXXXX I have here a diagram showing fibre length distribution in two trees. A plane running vertically in the north axis and extending from pith to bark is represented. (Explain details of diagram.)

Fibre length increases from pith to bark in the normal way, but the longer fibre length values occur some way up the stem.

How do we compare say this tree with the other? How do we rank them with regard to selection?

In pulping the whole tree is used. The ideal way of obtaining a whole-tree fibre length estimate would be to pulp the whole tree and find out the fibre length in a well-mixed sample of its pulp. We cannot do this in practical selection without destroying the tree.

A pure arithmetic mean of all the results would give us a result approximating 1.15 mm. This is also unsatisfactory as undue emphasis is placed on high and low values. What is needed is some kind of weighting system.

Let us look at these two diagrams XXXXXXXXXXXXXXXXX

CO-ORDINATES TRANSPARENCY

XXXXXXXXXXX

These diagrams illustrate the weighting principle, which is the accepted method for calculating the average fibre length of the tree. If we take the jth disk and look at fibre length observations in that disk, the coordinates of FLij are Ri cms from the pith and Hj feet from the ground. Within the disk, FLij is assumed representative of the annular area Aij whose extremities correspond to the mid-point between successive observations. The weighted disk fibre length (WDFL) is calculated through the relationship: WDFLj = Σ(FLij x Aij)/Aj This weighted disk fibre length is assumed representative of the fibre length in volume Vj of a cylinder of cross-sectional area Aj and extremities corresponding to the midpoints between successive disks. The weighted tree fibre length (WTFL) is determined by this relationship WTFL = Σ(WDFLj x Vj)/V The information needed is fibre length (FL), total height of the tree, total radius of disks, height of the disk from the ground (H), and distance of the sample from the pith (R). These data were punched on cards and analysed on the ANU’s IBM 360 computer. For the nine trees these are the whole tree fibre lengths.

89

XXXXXXXXXXXXXXX

WHOLE TREE FIBRE LENGTH TRANSPARENCY

XXXXXXXXXXXXXXXXX

You might say that these fibres are short but anything over a mm is OK for a eucalypt. These values are derived from mean values of 100 fibres at each sampling position.

When we look at a frequency distribution for fibre length in a single sample the distribution is approximately normal and we have shorter and longer fibres present than the whole tree fibre length figures imply.

XXXXXXXXXXXXXX

FIBRE DISTRIBUTION TRANSPARENCY

XXXXXXXXXXXXXXXXX

The very short fibres will be lost in the screening process but the longer fibres will help the bonding qualities of the paper.

This diagram also shows the difference in fibre length between that at breast height and that at 30% height and between that at 2.5cms from the pith and 15 cms from the pith.

Next we turn to the issue of where to extract our limited sample from the tree (sample representativeness). Whereabouts must the sample be taken to give us our best estimate of whole tree fibre length?

Let’s go back to the previous transparency and look at some correlation coefficients – XXXXXXXXXXXXXXX

FIBRE CORRELATION TABLE TRANSPARENCY

XXXXXXXXXXXXXXXXX

We can see from this table that fibre length in the breast height sample is only weakly correlated with whole tree fibre length.

This as a sad state of affairs as breast height is a most convenient sampling position. To obtain a correlation significant at the 1% level we would have to go to ground level where we’ll have difficulties operating the sampling machine and where buttressing will be an issue, or, we will have to go up to about 20% of tree height. This would represent about 30 – 35 feet [about 9 – 11 m) above ground on the trees from which we are making our selection. Operating a chainsaw-like mortising machine that far from the ground would be no picnic!

We also come to a sticky end if we try to correlate the result at breast height with the result at 30% height (correlation coefficient R = 0.39 and not significant).

In addition, to produce a pulp which would be comparable to that made from a softwood would require a much greater increase in fibre length than is currently being achieved in softwood breeding programmes. With E. deglupta we cannot really believe that any marked improvement in fibre length would be achieved as the natural variation is too narrow.

Thus, the combination of a non-representative breast height sample and the lack of any possibility of significant improvement of fibre length through breeding caused me to relegate selection based on fibre length to a secondary role.

Of the nine trees in this table – XXXXXXXXXXXXXXX

WTFL TRANSPARENCY AGAIN

XXXXXXXXXXXXXXXXX

I would be inclined only to reject the one tree with a whole tree fibre length of less than 1 mm.

In the detailed study of wood density in the same nine trees I’ve used an X-ray technique, which is a relatively new idea for this type of work.

Small samples of wood extending from the pith to the bark were machined from the larger samples for each percentage height point. The end-grain surfaces were accurately machined using a specially modified router until they were 6.9 mm apart.

XXXXXXXXXXXXXX

PASS AROUND WOOD SAMPLE

XXXXXXXXXXXXXXXXX

These samples were placed on a sheet of X-ray film and irradiated with calibrated X-rays. If the thickness of the sample does not vary the intensity of radiation after passing through the sample solely depends on variation in density of the wood. Thus, an X-ray negative such as this is representative of wood density.

XXXXXXXXXXXXXXX

PASS AROUND X-RAY NEGATIVE

XXXXXXXXXXXXXXXXX

Being a negative the light-coloured parts represent higher density areas, that it the X-rays found it harder to penetrate the denser wood.

90

A linear relationship exists between the optical density of the X-ray negative and the density of the sample at the corresponding point. It is therefore sufficient to be able to record the variations in optic density across the X-ray negative of a radial wood sample to obtain, directly, the variation in density of the wood.

Having obtained the X-ray with its standards we analyse it with a recording microdensitometer. This machine gives a trace such as the one I have here.

XXXXXXXXXXXXXXX

PASS AROUND DENSITY TRACING

XXXXXXXXXXXXXXXXX

The height of the trace above a base line is proportional to wood density. By measuring the area under the trace for a short distance we can obtain an average value for that location. We can relate this value to the set of standard density values that we know accurately. I have done that for six points along this trace.

Using a number of these traces we can depict the pattern of density in the tree.

XXXXXXXXXXXXXXX

TRANSPARENCY OF DENSITY PATTERN

XXXXXXXXXXXXXXXXXX

As you can see the density pattern is not as straight forward as the fibre length pattern. (Explain diagram) We can rank these trees for density by calculating weighted whole tree density (WWTD) in the same manner that we calculated whole tree fibre length.

XXXXXXXXXXXXXXX

WEIGHTING PRINCIPAL TRANSPARENCY

XXXXXXXXXXXXXXXXXX

We substitute density for fibre length in the first equation and weighted disk density (WDD) for WDFL in the second formula.

Here are the figures for the nine trees – XXXXXXXXXXXXXXX

DENSITY FIGURES TRANSPARENCY

XXXXXXXXXXXXXXXXXX

Excluding the three non-plantation trees, phenotypic variation in densities is evident, covering a range of some 20% at 8% moisture content.

For pulp manufacture, densities of around 27 to 30 lbs per cubic foot [432.5 to 480.6 kg/m3] have been advocated in the past.

However, the present opinion of the major pulp and paper companies using eucalypts, including APM, is that one should look for a low-density species and select for the higher densities within that species.

In the selection of “plus” trees of E. deglupta I advocate we do not specify a density and select for it. We should aim to segregate our candidate trees into two or at the most three density categories, for example, low and high or low, medium and high. I think it would be best to sort into low- and high-density groups, ignoring those in the middle. We could consider two other special classes: a low extreme and a high extreme.

XXXXXXXXXXXXXXXXX DENSITY CATEGORIES TRANSPARENCY

XXXXXXXXXXXXXXXXXX

Thus, we could get two possible outcomes, one with high density and one with low density. The resulting increased uniformity within a stand is likely to be of most importance in the future.

It seems likely that by looking at the wood properties of 100-150 plantation trees (the breeding population) we should end up with 30-40 good quality clones for our orchard (the propagation population).

Again, we come up against the issue of obtaining a representative sample from an individual tree without cutting it down.

Let’s look at the correlation coefficients for density.

XXXXXXXXXXXXXXXX

DENSITY CORRELATION TRANSPARENCY

XXXXXXXXXXXXXXXXXX

The relationships between whole tree density and density at ground level and at breast height are statistically not significant.

However, the correlation with 10% of total tree height is fairly good, being significant at the 1% level.

Therefore, our density sample must come from the 10% height level, that is, about 15 feet [4.6 m] from the ground. Although this will be more difficult than sampling at breast height, I think we can manage it using tall ladders Wood density is the result of a complex of characters including cell wall thickness and vessel volume in eucalypts, thus the criterion is not straightforward. Maximum efficiency in breeding for density cannot be achieved until one knows what

91

influence the components have in determining it. Only correctly designed clonal or seedling progeny trials can indicate the extent to which the genetic component is affecting wood density.

We are presently cruising plantations looking for suitable “Candidate” trees. We are not paying much attention to volume production.

If you examine the height and diameter growth data derived from growth plots at Keravat and which I have graphed in Figure 1 attached to your papers you will see that growth potential is quite adequate for our needs.

XXXXXXXXXXXXXX

TRANSPARENCY OF FIGURE 1

XXXXXXXXXXXXXXXXXX

The preliminary selection is being made for desirable external features, as these are easy to assess and also contribute to gross aspects of wood quality.

We are choosing trees that are straight and do not have marked fluting and spiral twists in the stem. Where possible we are selecting trees with minimum buttressing. It is likely that most of these features are largely genetically controlled and the form of the bole should be improved by selection.

Having selected 100-150 Candidate trees, we will apply our wood quality selection criteria. Samples will be removed from standing trees using a special machine incorporating a chainsaw motor power unit adapted to drive a mortising chain. This will give a pith to bark sample about 2’’ x 2’’ [5 cm x 5 cm] in tangential dimensions.

The wood property selection will be made firstly for density and a high selection differential will be applied. I would hope to discard at least two-thirds of the candidate trees in this initial stage. Density will be determined by the analysis of X-rays as I have described. A number of trees could be rejected on visual assessment of positive photographic prints of the X-rays, such as this one here.

XXXXXXXXXXXXX

PASS AROUND POSITIVE PRINT OF X-RAY PLATE

XXXXXXXXXXXXXXXXXX

The remainder will be analysed with the help of the recording microdensitometer to enable a decision to be made on the suitability of a particular tree.

A low selection differential will be applied to fibre length for reasons already discussed. The few trees with very short fibres will be rejected. We may have to eliminate another 5 – 10 trees at this stage.

By examination of X-rays from separate longitudinal and tangential irradiation of the wood sample, spiral grain may also be assessed.

XXXXXXXXXXXX

PASS AROUND TWO X-RAY PLATES (EXPLAIN)

XXXXXXXXXXXXXXXXXX

On this basis we may throw out a few more trees if we can afford to do so as spiral grain is supposedly strongly inherited and detrimental to most end uses of solid wood compared to use for pulp.

What else should we consider for the future genetic programme?

We must be very careful of just how much we include in our selection programme. If we try to extend our selection over too many characters our gains will be less and also the number of possible clones in the seed orchard would be reduced.

Cellulose content of the wood of E. deglupta may prove to be an important factor in pulping. Von Koeppen stated in 1958 that a sulphate pulp yield similar to that from other tropical species was obtained from 5-year-old plantation-grown E. deglupta; however, in 1965, Petroff, test pulping Kamarere grown in the Congo, found cellulose to be lower than that for all five other species of eucalypt tested at the same time.

A pulp industry operates on a large turnover and the profit margin per unit is relatively small. Thus, a very small increase in cellulose over lignin content could lead to a large increase in profit for the operating company. We haven’t yet looked at treeto-tree variation in cellulose content in E. deglupta grown in the Territory but I think we must do this. Perhaps it may pay to include it in the breeding plan in some way. This could lead to an increase in pulp yield from a given volume of wood but a general lack of heritability studies on chemical constituents of broadleaved species makes it difficult to predict any gain that might be achieved.

We hope to start off a seed orchard next planting season. We will probably use stock grafted by the “top-cleft” and “patch” methods and half-sib seed.

92

Kamarere has been routinely propagated by leaf and stem cuttings at Keravat from source material up to 6 months of age so far and several thousand cuttings of a number of unselected clones have been planted in a plantation near the Vudal River.

Experiments are continuing to try and find a method of routinely propagating cuttings from the crowns of mature trees. These involve variations of the mist spray regime and the application of different rooting hormones. We will see how we go!

Some experiments concerning seed orchard management are underway or proposed. Techniques to be used for breeding are being investigated. For example: the manner in which the inner operculum is shed, how the flowers open, and how and when to carry out emasculation.

A seed production study is underway at Keravat in an effort to ascertain how many trees will be needed in a seed orchard to produce a given amount of seed.

Seed collection from mature Kamarere is difficult because of its height growth and because it has a smooth branch-free trunk making it difficult to climb. We will be trying various combinations of lopping and tying down of branches to see if we can obtain a more compact tree crown and keep the seed crop low to the ground. The orchard will be laid out on a square or triangular grid so the possibility will exist in future to use hydraulic “cherry-pickers” like the ones used for picking fruit in Australia and elsewhere. I have seen one used by APM for pruning poplars, they are very handy, particularly the type that can be mounted on an ordinary wheeled tractor.

The collection of seed and scion material from select trees in the field has always been a problem in Australia and overseas.

Australian experience has shown that a high-powered rifle fitted with a telescopic sight is ideal for this type of work. We are negotiating purchase of a .222 Remington rifle and a 3-9 magnification zoom telescopic sight for use at Keravat.

We hope to get some type of provenance study underway also. Attempts are being made to obtain small quantities of seed from selected trees in various parts of the Territory. It is also hoped to include seed from Indonesia and the Philippines but we have not had much success in that direction yet.

III. PRODUCTION STUDIES In addition to the genetic program there is the need to determine appropriate economic procedures for the establishment and maintenance of large-scale plantations of E. deglupta with the primary objective of wood supply to the pulp industries. The use of clearing and burning for land preparation and the planting of tubed stock for plantation establishment should be reexamined. Away from the plantation area some type of planting on snig tracks and logged clearings will have to be used.

“Jiffy Pots”, a kind of pots made from compressed peat, have been used successfully for planting Eucalyptus species in many parts of Australia and overseas. Although not entirely appropriate in size, locally available Jiffy Pots have been used successfully to raise and plant out in the field thousands of cuttings at Keravat. It is hoped to obtain a quantity of peat pots of more appropriate dimensions and have a trial established in this year’s planting area at Keravat.

A planting trial was established some four months ago at Keravat to test various types of planting stock. Treatments include tubed stock controls, open rooted stock and stumped stock of various sizes. Some survival results are to hand for this trial. I have summarized these and they are given in Table 2 attached to your own copies of the paper.

XXXXXXXXXXXXX

TRANSPARENCY OF TABLE 2

XXXXXXXXXXXXXXXXXX

Apart from the tubed stock, survival was disappointing. Peat pots were not included in this trial.

We still have a lot to do in this part of the programme. I have mentioned several avenues of approach in the last couple of pages of my paper. It will probably be several years before everything is straightened out.

This planting season, we are establishing a second planting trial, which will be similar to the one mentioned here but is planned to include stock potted in tall jiffy pots. A spacing trial is planned with 8 x 8’ [about 2.5 x 2.5 m], 9 x 9’ [about 2.75 x 2.75 m], 12 x 12’ [about 3.66 x 3.66 m] and 15 x 15’ [about 4.6 x 4.6 m] (control) spacing treatments.

Studies will need to be made on length of planting season in various areas, plant size, tending, response of growth and yield to fertilizer application and fertility of soils converted to pure plantations.

Identification and control of nursery pathogens, wood destroying fungi and insect pests will assume more and more importance as the programme progresses.

Well, that’s about all I have to say. Twelve to eighteen months ago about five minutes would have been sufficient to relate our total knowledge of Kamarere. I’ve discussed Kamarere here for more than two hours today and have by no means covered

93

all of our present knowledge. We know enough at this stage to predict that our programme will be successful, but just how successful remains to be seen!

Above left: Specimen collected by Frans Arenz from the Kigara River (Milne Bay/Raba Raba) and deposited in the Lae Herbarium. Above right: Specimen collected by L D Pryor 4 miles (6 km) up the Kutu River (Milne Bay/Raba Raba) and deposited in the Gauba Herbarium, Department of Botany ANU. Left: Specimen collected by Frans Arenz from the Kigara River (Milne Bay/Raba Raba) and deposited in the Gauba Herbarium, Department of Botany ANU.

Meanwhile, on Friday 23 August Professor Pryor accompanied by Frans Arenz flew by charter to Milne Bay and visited Raba Raba.

Saturday and Sunday were spent inspecting and collecting specimens from natural stands of E. deglupta, on the Kigara River

94

and as far as four miles (6 km) upstream on the Kutu River. They returned to Port Moresby by charter early on Monday morning 26 August and Professor Pryor left for Australia later that afternoon.

I returned to Keravat on Sunday 1 September 1968. On arrival, I found a letter had been sent from the ANU offering me a Commonwealth Postgraduate Scholarship supplemented by the ANU out of research grant funds to study for a PhD30 in the Department of Forestry and building on the work already completed on E. deglupta towards my external Masters degree. However it was a requirement of study for a PhD under the Commonwealth Scholarship that the student attend ANU full time. This meant another prolonged absence from the Territory. Nevertheless I accepted the offer and advised the Department of Forests accordingly by memo dated 2 September.

Kevin White, then Chief, Division of Silviculture, wrote back on 9 September

on

behalf

of

the

Department, stating the Department had no objection, but raised issues on whether field work would be undertaken in Papua and New Guinea during the period and asking what supporting field work would be needed from staff in the Territory. He also pointed out the necessity that work continued on the volume tables and productivity investigations.

Following the receipt of Kevin’s letter on 21 September I advised the Registrar, ANU that I intended to transfer my enrolment from the external Masters Degree in Forestry to that for a PhD degree and that I intended to arrive in Canberra in early 1969.

30

Under the rules then in place in the ANU Faculty of Science in the School of General Studies, a student with an Honours first degree was not required to have a Masters Degree as a prerequisite to enrolment for a PhD.

95