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

RAINBOW EUCALYPT MAN A Parallel Journey John Davidson (Part 8)

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

1993 continued UNDP/FAO Regional Project – FORTIP I shared the responsibility for the design and carried out a number of consultancies in implementing the UNDP/FAO Regional Project on Improved Productivity of Man-made Forests Through Application of Technological Advances in Tree Breeding and Propagation, otherwise known as the FORTIP Project.

Ten countries were involved – Bangladesh, Bhutan, India, Indonesia, Malaysia, Nepal, Pakistan, Philippines, Sri Lanka and Thailand. At a pre-project planning workshop, country representatives identified a small number of eucalypt species to which priority should be given. E. deglupta was one of those species. (The others were E. camaldulensis, E. globulus, E. grandis/E. saligna, E. pellita and E. tereticornis.)

Over the course of a couple of visits to the Project Headquarters in Los Baños, Philippines and after travel around the participating countries, in 1993 I produced a Domestication and Breeding Programme for Eucalyptus in the Asia-Pacific Region. E. deglupta was one of the chapters (on pages134 – 146). This document was published by FAO five years later and towards the end of the Project (January 1998).1

Publication of Eucalypt Domestication and Breeding In October 1993, Clarendon Press in Oxford published our book on Eucalypt Domestication and Breeding, at last realising our goal set out at the Third World Consultation on Forest Tree Breeding held in Canberra in 1977.

1

Davidson J 1998 Domestication and Breeding Programme for Eucalyptus in the Asia – Pacific Region. UNDP/FAO Regional Project on Improved Productivity of Man-Made Forests Through Application of Technological Advances in Tree Breeding and Propagation – FORTIP, RAS/91/004, Field Document No. 25, January 1998, FAO Los Baños Philippines. 252pp.

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On pages 73 – 80, E. deglupta was one of twelve species covered in detail under sub-headings of natural forests, plantations, genecology, provenance variation, base populations for selection and conservation.

The first print run was a hard cover edition of one thousand, targeted at libraries, university student reading lists and other places where durability was seen as a requirement and was standard for the Oxford press at the time. This edition was priced at £55.00.

1994 Second edition of Eucalypt Domestication and Breeding As a result of sending out flyers to the nearly 300 members of the Breeding Eucalypts Working Party of IUFRO, our eucalypt book sold well and the first printing ran out very quickly. The authors were contracted to produce a paperback edition at a lower cost. Vlad Mosmondor in the CSIRO Division of Forest Research Canberra designed the cover. This edition was first published on 4 August 1994 with only minor internal editorial changes and sold for £35.00. (In 1993-1994 £1.00 was equivalent to about A$2.00.)

This print run also sold out and a second print run of the paperback was undertaken in late September 1994.2The authors authorised future reprinting by digital means as required. Those copies have the

2

Since I was receiving 40% of the royalty on each copy sold, I could keep track of the sales. However, there were a number of pirated facsimile copies noticed in offices across Asia during my travels as a consultant for which neither the Oxford Press nor the authors received any remuneration.

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message “This book has been printed digitally in order to ensure its continuing availability” in the publication details section at the front.

1995 Genetic variation in height growth and leaf colour in E. deglupta High levels of additive genetic variance were found in height growth up to 16 months of age in three progeny tests of E. deglupta in Costa Rica.3Heritabilities were within the range typically found in forest trees. High levels of additive genetic variation and family heritabilities in leaf colour were also found.

Purple leaves were associated with slow growth, and green leaves with fast growth. The best families showed a high degree of stability across the three sites. Predicted genetic gain from directed seed collections of the best mother-trees ranged from 12.4 to 15.8% 3

Cornelius J P, Corea E A and Mesén J F 1995 Genetic variation in height growth and leaf colour of Eucalyptus deglupta Blume at ages up to 16 months in Costa Rica. Forest Ecology and Management 75(1-3):49-59.

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1997 Pulping and papermaking potential of plantation-grown E. deglupta from PNG A long-awaited synthesis of the pulping and papermaking potential of plantation-grown E. deglupta was published in 1997.4A quote of the abstract follows: “The papermaking properties of both unbleached and bleached sulphate as well as unbleached neutral sulphite semichemical (NSSC) pulps were evaluated to determine the effect of age on pulpwood quality in Eucalyptus deglupta. All age classes of pulpwood could be pulped readily by the sulphate process and good yields of pulp were obtained. With increasing age and constant alkali, higher screened pulp yields were produced at lower Kappa numbers. As the age increased, the volume of wood, size of plantation and amount of alkali required to produce one tonne of sulphate pulp decrease. Sulphate pulps had very good papermaking properties which will be adequate for a wide range of unbleached paper grades. High brightness sulphate pulps were readily produced using a CEHD [C: Chlorination; E: Extraction; H: Hypo; D: Chlorine Dioxide] bleaching sequence from unbleached pulps with Kappa numbers less than 35. These bleached pulps were suitable for the manufacture of a wide range of bleached paper products. NSSC pulps were suitable for use in corrugating medium and in a variety of other paper and paperboard products especially if pulped to a Kappa number of about 100. Eucalyptus deglupta pulpwood from plantations at any age between 3 and 20 years was very suitable for chemical and semichemical pulp production. The 10- to 12-year-old trees were the preferred age classes for sulphate pulping based on the combination of early harvesting for economic reasons and acceptable pulping and papermaking properties.”

2003 Molecular studies on Eucalyptus subgenus Minutifructis Eucalyptus subgenus Minutifructis (sensu Brooker 2000) (replacing the informal subgenus ‘Telocalyptus’, one of eight informal subgenera of Eucalyptus sensu Johnson 1976) comprises four ‘tropical box’ species and is characterised by compound terminal inflorescences (rare in eucalypts), very small buds and fruits and discolourous leaves with densely reticulate veins.5Minutifructis is divided into two sections: Equatoria, the name referring to the equatorial distribution in the Philippines, Indonesia and PNG of the lone species E. deglupta in the section, and Domesticae referring to the occurrence on the Australian mainland of all three species comprising the section, E. brachyandra in northern Western Australia, E. howittiana in north eastern Queensland and E. raveretiana in central-eastern Queensland.

Before 1976 Pryor and Johnson (1971) had divided these four species between two sections of subgenus Symphyomyrtus: section Equatoria, series Degluptae (E. deglupta, E. raveretiana and E. brachyandra) and section Howittianae (E. howittiana). In 1988 Chippendale followed this arrangement describing two 4

Phillips F H, Logan A F and Harries E D 1997 The pulping and papermaking potential of plantation-grown Eucalyptus deglupta from Papua New Guinea. Part 2. Sulphate and NSSC pulps from wood of various ages. Earlier articles included: Phillips F H, Logan A F and Balodis V 1979 Suitability of the tropical forest for pulpwood: mixed hardwood, residues and reforestation species. Tappi 62(3):77-81 and, Logan A F 1981 Pulping of tropical hardwood reforestation species. CSIRO Australia, Division of Chemical Technology, Research Review.

5 Johnson L A S 1976 Problems of species and genera in Eucalyptus (Myrtaceae). Plant Systematics and Evolution 125:155-167; Brooker M I H 2000 A new classification of the genus Eucalyptus L’Her. (Myrtaceae). Australian Systematic Botany 13: 79-148.

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series on the basis of differences in inflorescence and fruit morphology: Myrtiformes (corresponding to Pryor and Johnson’s 1971 section Equatoria series Degluptae) and Howittianae.6 Papers by Sale et al. (1993, 1996) and Steane et al. (2002) indicated that Minutifructus should not be ranked as high as in a separate subgenus and instead the species be nested within subgenus Symphyomyrtus.7 Molecular data presented by Steane et al. (2002) and Whittock et al. (2003) support the earlier opinion of Pryor and Johnson (1971) that species from Minutifructus evolved from within subgenus Symphyomyrtus8 leading to the hypothesis that E. deglupta evolved outside Australia from a symphyomyrt ancestor possibly during northwards and westwards migration of fragments of the Australian plate and long-distance dispersal by wind and updrafts from volcanoes along the western margin of the “Pacific rim of fire”.

This opinion is supported by some current hybridisation successes. Normally crosses between eucalypt species from different major subgenera are not successful. Generally, the more closely related eucalypt taxa are, the more likely they will make successful interspecific crosses. Manipulated interspecific crosses of E. deglupta from Minutifructus and two species of subgenus Symphyomyrtus section Latoangulatae (sensu Brooker 2000), E. deglupta♀ x E. pellita♂ and E. deglupta♀ x E. urophylla♂, were successful at PICOP in the Philippines.

Whittock et al. 20039produced a strict consensus cladogram of species from Minutifructus within subgenus Symphyomyrtus (figure on next page). In this figure the branch hierarchy supports E. deglupta (section Equatoria) and E. brachyandra (section Domesticae sensu Brooker) being more closely aligned with sections Exsertaria (SE), Latoangulatae (SL) and Maidenaria (SM) in clades B and C. E. howittiana and E. raveretiana (also both section Domesticae) appear to have an ancestor in common with sections Bisectae (SB part 1), Adnataria (SA) and Dumaria (SD) in Clade A.

6

Chippendale G M 1988 Eucalyptus, Angophora (Myrtaceae), Flora of Australia 19. Australian Government Publishing Service, Canberra; Pryor L D & Johnson L A S 1971 A Classification of the Eucalypts. Australian National University, Canberra.

7 Sale M M, Potts B M, West A K and Reid J B 1993 Relationships within Eucalyptus using chloroplast DNA. Australian Systematic Botany 6:127-138; Sale M M, Potts B M, West A K and Reid J B 1996 Relationships within Eucalyptus using PCR-amplification and southern hybridisation of chloroplast DNA. Australian Systematic Botany 9:273-282.

8 Steane D A, Nicolle D, McKinnon G E, Vaillancourt R E and Potts B M 2002 Higher level relationships among the eucalypts are resolved

by ITS-sequence data. Australian Systematic Botany 15:49-62.

9 Whittock S, Steane D A, Vaillancourt R E & Potts B M 2003 Molecular evidence shows that the tropical boxes (Eucalyptus subgenus Minutifructis) are over-ranked. Transactions of the Royal Society of South Australia 127(1):27-32.

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This figure from Whittock et al. 2003 shows the “Symphyomyrtus” part of a strict consensus cladogram from cladistic analysis of internal transcribed spacer (ITS) data showing positions of species from Minutifructus within subgenus Symphyomyrtus. Bootstrap percentages over 50% and branch lengths (from a single tree) are shown respectively above and below the internal branches. A, B, C and D are clades of subgenus Symphyomyrtus.

Clade A comprises sections Adnataria (SA), Bisectae (SB) (part I), Platysperma (SP), Dumaria (SD), as well as E. howittiana and E. raveretiana from subgenus Minutifructus; Clade B sections Latoangulatae (SL) and Exsertaria (SE); Clade C sections Maidenaria (SM) and Racemus (SR); Clade D sections Inclusae (SI) and Bisectae (SB) (part II). ME = Minutifructis section Equatoria; MD = Minutifructis section Domesticae.

Origins: E. howittiana 1 MS1999 Atherton Queensland, E. howittiana 2 DN2526 Northern Ranges, Queensland, E. raveretiana 852255 Australian Botanic Garden Mount Annan NSW, E. deglupta 1 Philippines CSIRO seed lot 19492-1, E. deglupta 2 New Britain?

(Bulolo PNG CSIRO seed lot 19790B), E. deglupta 3 New Britain? (Bulolo CSIRO seed lot 19791B), E. deglupta 4 A cultivated specimen from the Botanical Gardens in Darwin (origin unknown).

The seed lots of E. deglupta attributed to Bulolo, whether collected there from the Seed Orchard, or from within and around the town of Bulolo, are most likely of New Britain origin. If from the Seed Orchard, the origin is more specifically from Keravat plantations of local provenance in East New Britain.

There is an anomaly in that E. deglupta 2 and 3 probably collected from the same locality in Bulolo, given their consecutive CSIRO seed lot numbers, branch at different levels. That E. deglupta 3 branches at the same level as 1 and 4 suggests the anomalous one is 2. GenBank sequences of ITS indicate E. deglupta 2 differs from 3 by four substitutions and one insertion. Section Equatoria (ME), with its three E. deglupta (1,3 and 4, excluding the apparently anomalous 2), falls neatly and separately in the middle 338

of the very large subgenus Symphyomyrtus. E. raveretiana and E. howittiana 1 and 2 of section Domesticae (MD) however are imbedded in Clade A of subgenus Symphyomyrtus. E. howittiana 1 and 2 from two different localities at Atherton and Northern Ranges respectively in Queensland branch at different levels. E. brachyandra branches earlier than E. deglupta 1, 3 and 4 and, like the latter, does not fall within any of the four clades A, B, C and D of subgenus Symphyomyrtus.

Among Eucalyptus only five species have been documented to have a coralline calyptra with externally visible, weakly fused petals with overlapping edges that do not form a single structure. Four suture lines appear as a cross like the cross on a hot cross bun. Only three of these species, E. brachyandra, E. guilfoylei and E. microcorys, are within or near subgenus Symphyomyrtus. (Brooker (2000) placed E. guilfoylei by itself in Eucalyptus subgenus Cruciformes because of the prominent cross-like suture on the calyptra.) This morphological character in E. brachyandra is sufficient to distinguish it from the other three species in Minutifructus and may indicate it is older in the evolutionary scale than E. deglupta, E. howittiana and E. raveretiana. In the evolutionary context it is also worth noting that the petal edges are obvious in the Patagonian fossil flower buds dated from the early Eocene at about 51.9 million years ago.10 There are many uncertainties in dating eucalypt evolution using molecular phylogenies. The exact geological age relationship of all four species in Minutifructis has yet to be determined. In the Whittock et al. 2003 fully-labelled ITS tree given above, E. brachyandra is the taxon shown arising from node C of the following Figure 3 from Crisp et al. 2004.11If all four species are added to the branch from node C, then node C becomes an estimate of the age of the Minutifructis clade. Node C is estimated by Crisp et al. 2004 in their Table 4 at 38-26 million years old (their ‘most recent’ column) and 45-26 million years old (their ‘earliest’ column) but in their Figure 3 the node is shown at about 38 million years of age. Ladiges and Udovicic 200512 estimate the E. deglupta node 2 to have a geological age of about 8 million years. This agrees with the geological model for the geographic region in which the species occurs. Isolation of E. brachyandra in the Kimberley in northwest Australia is a much older (earlier) event if it is assumed to have the same age as node C (38 million years).

10

Gandolpho M A, Hermsen E J, Zamaloa M C, Nixon K C, González C C, Wilf P, Cúneo N R and Johnson K R 2011, Oldest known Eucalyptus fossils are from South America, PloS ONE 6(6):e21084 doi:10.1371/journal.pone.0021084. Epub. June 28, 2011.

11 Crisp M, Cook L and Steane D 2004 Radiation of the Australian flora: what can comparisons of molecular phylogenies across multiple taxa tell us about the evolution of diversity in present-day communities? Philosophical Transactions of the Royal Society of London B 359:1551-1571.

12 Ladiges P Y and Udovicic F 2005 Comment on the molecular dating of the age of eucalypts. Australian Systematic Botany 18:291-293.

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Chronogram of eucalypts sourced from Crisp et al. 2004 loc cit. that compares estimates of divergence times (million years ago (Mya), scale on the right) among eucalypt lineages based on climatic and tectonic events, and assuming the basal divergence between the eucalypts sensu lato (arrow at node A) and Arillastrum at 70 Mya based on calibration from a vicariance event. The grey bars over the nodes indicate there is a range in the estimates because of variation in topology and choice of method and calibration points. The dating of the Patagonian fossils at about 51.9 Mya and their placement in Symphyomyrtus could be accommodated in this figure without altering the structure at the younger nodes. The dashed lineages are extra-Australian. Symbols indicate the biome in which each terminal taxon occurs: Ω, aseasonal wet; Δ, monsoonal; Ο, southwest temperate; +, southeast temperate;§, eremean; * other.

Node B: Allosyncarpia (monsoonal) versus Eucalyptopsis (wet tropics); Node C: Ancestor of E. deglupta clade (Southeast Asia) diverges from Australian sister taxon; Node E: divergence of E. urophylla (Timor) clade from Australian sister taxon. The four asterisks are equivalent E. deglupta numbers 1, 3 4 and 2 (left to right) from Whittock’s figure given earlier. The triangle immediately to the left of the asterisks is equivalent to E. brachyandra from the same source.

Relationships of the four tropical boxes within subgenus Symphyomyrtus. Nodes 1 and 2 are supported by both morphology and cpDNA data.

Node C is as in Figure 3 in Crisp et al. 2004 given above and possibly also supported by morphological data (for example the cross-like suture on the calyptra of E. brachyandra). Q, Queensland; NG, New Guinea; P, Philippines; K. Kimberley. Node C is shown by Crisp et al. 2004 in their Figure 3 at about 38 million years of age The E. deglupta node 2 is estimated by Ladiges and Udovicic 2005 to have a geological age of about 8 million years. Isolation of E. brachyandra in the Kimberley in northwest Australia is a much older (earlier) event if it has the same age as node C (38 million years).

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A more recent paper by Grattapaglia et al. 201213employed a Splitstree4 analysis14from genome-wide genotyping of 94 species in Eucalyptus sensu stricto to present a different way of viewing the phylogenetic network. Most of the industrial eucalypt plantations of the world are based on just a few eucalypt species. Nine are listed in the Splitstree figure below in red. They occur in only two main poorly-differentiated clades and in only three sections - Latoangulatae and Exsertaria (“Clade B”) and Maidenaria (“Clade C”). Of those nine species, the three in bold red and with an asterisk, one from each section, have been the subject of detailed genomic studies.

A Splitstree4 summary analysis of 94 species in Eucalyptus sensu stricta is shown. 8,354 DArT markers from genome-wide genotyping were used. The DArT phylogeny provided more resolution within major clades than had been obtained previously. The results largely agreed with traditional taxonomy and ITS-based phylogenies presented earlier, but I have removed from this figure reference to Minutifructis as a sub-genus on the basis of the earlier discussion about it being over-ranked. Instead, the individual species have been named and nested within subgenus Symphyomyrtus in the locations previously occupied by sections Domesticae and Equatoria of Minutifructis. More data are required to determine whether the association of E. raveretiana and E. howittiana (section Domesticae) with sections Dumaria and Adnataria is real or an artifact (dashed oval around “Clade A” in the figure). E. brachyandra was not included in this analysis. The remainder of the figure was sourced unchanged from Grattapaglia et al. 2012 loc cit.

13 Grattapaglia D, Vaillancourt R E, Shepherd M, Thumma B R, Foley W, Külheim C, Potts B M and Myburg A A 2012 Progress in Myrtaceae genetics and genomics: Eucalyptus as the pivotal genus. Tree Genetics and Genomes 8(3):463-508.

14 Hudson D H and Bryant D 2006 Application of phylogenetic networks in evolutionary studies. Molecular Biology and Evolution 23(2):254-267.

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2007 Breeding programme for E. deglupta in the Solomon Islands In the Solomon Islands, an active breeding program for E. deglupta in the 2002-2007 period resulted in the selection of 40 plus trees which were progeny tested and the trial culled to a Seedling Seed Orchard with 20 families in 4 replications.

2008 E. deglupta Seed Orchard Bulolo This orchard had been used for over 35 years. Through fires and vandalism, the number of trees has diminished over time (photographs on the next page). An attempt under an ACIAR project to establish a new clonal seed orchard at Bulolo in 2006 comprising 30 clones with four ramets per clone was unsuccessful.

2011 Herbarium collections of E. deglupta by K Damas From 20 April to 5 May 2011, K Damas from the National Herbarium in Lae travelled in West New Britain in the vicinity of Kimbe, Biala and Talasea, collecting over 60 specimens from the Dagi, Ru, Berima, Evula, Savula, Kuludagi and Kapaluk Rivers and Moyou Creek which have been deposited in Lae with duplicates where available sent to herbaria in Bogor and Melbourne.

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Above: Recent seed collection in the E. deglupta seed orchard Bulolo. (Photograph: D Spencer) Right and below: Perfectly grafted trees at Bulolo about 35 years old in 2008. (Photographs: via Steve Midgley, CSIRO, Canberra)

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Open Bay Timber Company In September 2011, Open Bay Timber Limited (OBT) was granted Forest Stewardship Council (FSC) certification of 11,770 ha of plantation forests under management at Open Bay, New Britain, mainly E. deglupta (photograph below).

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2013 E. deglupta chloroplast genome sequenced In November 2013 came the announcement that the complete chloroplast genome of E. deglupta had been sequenced.15Comprising 160,177 base pairs, the genome is in the public domain at the National Centre for Biotechnology Information (NCBI) (Reference Sequence NC_022399).16

Parts of the record available at the NCBI for the complete chloroplast genome of E. deglupta. These are the instructions in a gene that tell the cell how to make a specific protein. A, C, G, and T are the “letters” of the DNA (Deoxyribonucleic Acid) code, also known as “bases”. They stand for the chemicals adenine (A), cytosine (C), guanine (G), and thymine (T) that make up the nucleotide bases of DNA. Adenine pairs with thymine, and cytosine pairs with guanine. There are 160177 bases in the complete chloroplast genome for E. deglupta. It would take 46 pages like the one on the right to print out the whole genome!

15

Bayly M J, Rigault P, Spokevicius A, Ladiges P Y, Ades P K, Anderson C, Bossinger G, Merchant A, Udovicic F, Woodrow I E and Tibbits J 2013 Chloroplast genome analysis of Australian eucalypts – Eucalyptus, Corymbia, Angophora, Allosyncarpia and Stockwellia (Myrtaceae). Molecular Phylogenetics and Evolution 69(3):704-716.

16 Available at: NCBI; https://www.ncbi.nlm.nih.gov/genomes/GenomesGroup.cgi?opt=plastid&taxid=2759

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The GenBank sequence database is an open access, annotated collection of all publicly available nucleotide sequences and their protein translations. This database is produced and maintained by the National Center for Biotechnology Information as part of the International Nucleotide Sequence Database Collaboration.

The eucalypts are predominantly out-crossers with hermaphroditic animal-pollinated flowers, eucalypts are highly heterozygous and display pre- and post-zygotic barriers to selfing to reduce inbreeding depression for fitness and survival.

The evolutionary history of the Eucalyptus genome is marked by a lineage-specific palaeo-tetraploidy event newly revealed by genomic analysis, superimposed on the earlier palaeo-hexaploidy event shared by all eudicots.17The whole-genome duplication (WGD) is estimated by Bayly and co-authors to have occurred about 109.9 (105.9–113.9) million years (Myr) ago in a Gondwanan ancestor around the time when Australia and Antarctica began to separate from East Gondwana. This WGD event is considerably older than those typically detected in other rosids.18 In 1971, Pryor and Johnson proposed an informal taxonomic classification of all eucalypts, which has become widely accepted as portraying the natural groupings within the genus. However, there have been subsequent modifications, notably the upgrading of Corymbia to the level of genus by Hill and Johnson in 1995 and subsequent downgrading it back to a subgenus of Eucalyptus by Brooker in 2000. E. cloeziana did not fit easily into established subgenera due to its unique operculum structure and thus remained the monotypic member of the subgenus Idiogenes. However, E. cloeziana is recognized by several authors as having a close affinity with the subgenus Monocalyptus. Phylogenetic assessment of the genus has supported E. cloeziana's position as basal to Monocalyptus19 and its subgeneric status was maintained in the formal classification proposed by Brooker in 2000. In a revision of white mahoganies (Monocalyptus, series Acmenoideae), Hill in 1999 proposed to split E. acmenioides into several species.

Reports of both natural and manipulated hybridization are common between eucalypts (Eucalyptus, Corymbia and Angophora), with 289 of the 528 species (55 %) reviewed by Griffin et al. reported to hybridize with at least one other species.20Subgeneric classification is thought to delimit the extent of 17

Bayly et al. loc. cit.

18

The “rosids” are a large clade of flowering plants. It includes about 70,000 species, more than a quarter of all angiosperms. The rosids are divided into 17 orders. These orders together make up about 140 families.

19 For example: Sale M M, Potts B M, West A K, and Reid J B 1993 Relationships within Eucalyptus using chloroplast DNA. Australian Systematic Botany 6:127-138.

20 Griffin A R, Burgess I P and Wolf L 1988 Patterns of natural and manipulated hybridisation in the genus Eucalyptus L’Hérit. – a review.

Australian Journal of Botany 36:41-66.

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hybridization in eucalypts. Hybridizing taxa largely occur between parapatric or sympatric populations of closely related species found within the same series or section. The only substantiated case of hybridization between subgenera occurs from artificial crosses between E. deglupta (subgenus Telocalyptus = Minutifructis) and taxa from the subgenus Symphyomyrtus (see earlier). These hybrids were reported by Griffin et al., to produce weak seedlings that were unlikely to survive to sexual maturity. However, molecular phylogenetic analysis of the chloroplast genome and the nuclear ITS region in eucalypts has placed the subgenus Telocalyptus = Minutifructis within the Symphyomyrtus clade, hence suggesting Telocalyptus = Minutifructis, and thus E. deglupta, be incorporated into Symphyomyrtus.21 This highlights the existence of inconsistencies that still reside in the taxonomic classification within the Eucalyptus genus.

ACIAR Project “Facilitating the availability and use of improved germplasm for forestry and agroforestry in Papua New Guinea” The rationale for this Project was to support community tree planting programs in PNG by making available improved germplasm for a range of high value timber species as well as some non-timber species, and by training rural communities in the propagation and deployment of this material.

Additionally, new eucalypt hybrids with potential for plantation forestry in PNG and north Queensland also formed part of the strategy. The three-year project started in August 2005 involving Partner country institutions; Papua New Guinea Forest Research Institute, The Foundation for People and Community Development, Queensland Department of Primary Industry Agency for Food and Fibre Science and CSIRO. In PNG, the Project was to develop improved knowledge and establish germplasm sources for 11 target tree species; Dracontomelon dao (walnut), Calophyllum euryphyllum (kalophilum), Endospermum medullosum (basswood), Tectona grandis (teak), Pometia pinnata (taun), Eaglewood (Gyrinops ledermanii) (eaglewood), Santalum macgregorii (PG sandalwood), Santalum album (Indian sandalwood), Acacia crassicarpa (crassicarpa), Eucalyptus pellita (pellita) and E. deglupta (kamarere).

An attempt was to be made to develop a hybrid between E. deglupta and E. pellita. In Australia, a series of hybrid combinations was to be trialled in Queensland.

21

Sale et al. loc cit. and Steane D A, McKinnon G E, Vaillancourt R E and Potts B M 1999 ITS sequence data resolve higher-level relationships among the eucalypts. Molecular Phylogenetics and Evolution 12:215-223.

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A final report was prepared by Brian Gunn and published by ACIAR in November 2013.22 No significant progress was made at the time on advancement of germplasm supplies or the development of novel eucalypt hybrid combinations under Objective 3: “To develop Eucalyptus deglupta germplasm”.

The lessons from the almost 50 years of experiences with breeding E. deglupta in PNG, the Philippines and elsewhere are that not only in-situ stands containing valuable genetic resources continually at risk, but there may be even greater threats to economically-valuable ex-situ products of ongoing breeding programmes such as provenance block plantings, seed orchards, progeny trials, clone banks, hybrid trials and the like as well as vital associated records and documentation.

2014 Reference genome of E. grandis released On 19 June 2014, an international consortium of researchers, including US Department of Energy Joint Genome Institute (DOE JGI) scientists23, released the reference genome of E. grandis. The effort to sequence and analyse the 640 million base pair genome of the species employed more than 80 research scientists from 30 institutions in 18 countries. Results were that Eucalyptus has 36,376 genes, 10,049 clusters, and 30,341 of the genes in those clusters Among the more than 36,376 genes found in Eucalyptus (nearly twice as many as in the human genome), the researchers discovered many that influence the production of secondary cell wall material that can be processed for pulp, paper, biomaterials and bioenergy applications. Comparative analysis of the complex traits associated with the Eucalyptus genome offers new opportunities for accelerating breeding cycles for sustainable biomass productivity and optimal wood quality and for adaptation to climate change. The genome data provide a diagnostic tool for understanding the basis of extremely fast growth

22 Gunn B 2013 Facilitating the availability and use of improved germplasm for forestry and agroforestry in Papua New Guinea. Project FST/2004/009, Final Report No. FR2013-17. Australian Centre for International Agricultural Research (ACIAR), Canberra. 41pp.

23 Myburg A A, Grattapaglia D, Tuskan G A, Hellsten U, Hayes R D, Grimwood J, Jenkins J, Lindquist E, Tice H, Bauer D, Goodstein D M, Dubchak I, Poliakov A, Mizrachi E, Kullan A R K, Hussey S G, Pinard D, van der Merwe K, Singh P, van Jaarsveld I, Silva O B Junior, Togawa R C, Pappas M R, D Faria D A, Sansaloni C P, Petroli C D, Xiaohan Yang, Ranjan R, Tschaplinski T J, Chu-Yu Ye, Ting Li, Sterck L, Vanneste K, Murat F, Soler M, Clemente H S, Saidi N, Hua Cassan-Wang, Dunand C, Hefer C A, Bornberg-Bauer E, Kersting A R, Vining K, Amarasinghe V, Ranik M, Naithani S, Elser J, Boyd A E, Liston A, Spatafora J W, Dharmwardhana P, Raja R, Sullivan C, Romanel E, Alves-Ferreira M, Külheim C, Foley W, Carocha V, Paiva J, Kudrna D, Brommonschenkel S H, Pasquali G, Byrne M, Rigault P, Tibbits J, Spokevicius A, Jones R C, Steane D A, Vaillancourt R E, Potts B M, Joubert F, Barry K, Pappas G J, Strauss S H, Jaiswal P, Grima-Pettenati J, Salse J, van de Peer Y, Rokhsar D S and Schmutz J 2014 The genome of Eucalyptus grandis. Nature 510:356362. (19 June 2014)

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rates and optimum wood and fibre properties. Eucalypt trees are grown on over 40 million hectares in 100 countries across six continents.

The extensive catalogue of genes will allow eucalypt breeders to rapidly adapt genotypes for many purposes. The genome data are available publicly through the DOE JGI’s comparative plant genomics portal known as Phytozome (http://bit.ly/Phytozome-Eucalyptus).

2015 Article on E. deglupta in “The Forester”

On page 11 of the June 2015 issue of The Forester (left), the Editor posed the question “Is this the only eucalypt

to

occur

naturally

in

the

northern

hemisphere?”

I submitted a detailed reply in the following issue (August 2015) (below).24

24

Davidson J 2015 Eucalyptus deglupta. The Forester, June 2015:28-31

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2017 E. deglupta in the Adelaide Botanic Garden The Botanic Gardens and State Herbarium of South Australia planted two trees in the Botanic Park arboretum in 2016. In 2017 four more were added to the Park and one in Adelaide Botanic Garden.

2018 E. deglupta in “Trees for Life in Oceania” E. deglupta was among 53 species, five of them eucalypts, that featured in this publication in 2018.

25

The following points

were made about the species.

25

Davidson J, Gunn B and Spencer D 2018 Eucalyptus deglupta. Pages 104 – 107 in Thomson L, Doran J and Clarke B (eds) 2018 Trees for life in Oceania: conservation and use of genetic diversity. ACIAR Monograph No. 201. Australian Centre for International Agricultural Research: Canberra. 278 pp.

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E. deglupta is a fast-growing hardwood suitable for planting in the lowland tropics where many other species of eucalypt do not thrive. It generally has excellent form, is easy to propagate from cuttings and is relatively free of many of the diseases that afflict eucalypts grown

in

the

tropics.

Disadvantages include its poor coppicing

ability,

its

requirement for fertile well-drained sites for optimum growth, and fire and cyclone susceptibility.

There is considerable diversity among trees from different regions in terms of morphology, stem form, wood properties and resistance to pests and diseases. Breeding programs have capitalised on selection for fast growth, more straight and cylindrical stems and higher and more uniform wood density in plantation-grown wood destined for pulping. Selection of particular provenances and hybridisation of these with other tropical eucalypts have improved resistance to foliar leaf spot diseases and stem borers.

Results of early provenance testing showed the best performing provenances to be the north coast of New Britain PNG and from southeast of Bislig Bay, Mindanao. There is considerable variation among regions in morphology, stem form, wood properties and resistance to pests and diseases. Indonesian provenances have not been sufficiently tested against Philippines and PNG provenances.

In the immediate future, the importance of E. deglupta lies in capturing its fast growth, excellent form, ease of propagation by cuttings and almost complete freedom from the leaf spot diseases that afflict most other eucalypts when grown in the lowland humid tropics, by developing hybrid combinations with such species as E. urophylla, E. wetarensis (tolerance of a more diverse range of site quality, higher wood density and some disease tolerance) and New Guinea sources of E. biterranea (higher wood density, good disease tolerance). E. deglupta x E. pellita was initially very successful in Mindanao, Philippines, 351

where the best hybrids were clonally propagated using cuttings. The following populations were expected to provide useful base/breeding populations: Geshes Clonal Seed Orchard, Bulolo, PNG subline (seed is available from 9 ramets of 3 surviving clones) North Coast, East New Britain, PNG subline. (New collections are required.)

North Coast, West New Britain, PNG subline. (New collections are required.)

West Sepik, PNG subline. (New collections are required.)

West Papua, Indonesia subline (New collections are required from surviving depleted stands in the two natural locations.)

Takalar Seedling Seed Orchard, South Sulawesi, Indonesia subline (present status needs to be determined, otherwise new collections are required).

Kenangan Seedling Seed Orchard, East Kalimantan, Indonesia subline (present status needs to be determined, otherwise new collections are required).

Mindanao, Philippines subline (new collections are required from natural stands surviving in New Bataan, Bislig, Monkayo and Pasian localities).

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The basic breeding strategy shown in the upper figure on the previous page is still relevant. If PNG wishes to start up the breeding programme again with E. deglupta, the basic strategy would need to expand into a more detailed breeding plan like the one given on the bottom of the previous page that would run over several generations (3 generations over 15 – 20 years are shown).

Knowledge of the E. grandis nuclear genome has already paved the way for detailed studies of genes associated with wood production, and for development of a new system for genotyping eucalypt individuals using genome-wide markers that could be widely applied to population studies and tree breeding programmes.

If full advantage is taken of the eucalypt genomes, marker assisted selection could be undertaken on juvenile seedling and clonal material grown in a glasshouse to rapidly turn over generations without recourse to field trials of adult material. Such a strategy would enable elements in the above breeding plan to be collapsed perhaps by a factor of ten in the time taken to produce results from the selections at each step.

2019 Our book on “Eucalypt Domestication and Breeding” has remained relevant. It often is given still as a reference in articles and publications on eucalypts. It remains available today from the publisher for £80.00. (In July 2019, £80 = US$100 = A$144 approximately.)

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Second hand paperback copies also turn up on ebay and Amazon from time to time and command a rather high price. One such advertisement in June 2019 is shown here for a used copy priced at A$163.95!

Geological time line with reference to some eucalypts With some new information it was possible to expand on the geological time line with reference to some eucalypts that I had presented in “The Forester” in 2015.26

26

Sources include: Ladiges P Y, Udovicic F and Nelson G 2003, Australian biogeographic connections and the phylogeny of large genera in the plant family Myrtaceae, Journal of Biogeography, 30(1), 989—998, Crisp M D, Cook L G and Steane D A 2004, Radiation of the Australian flora: what can comparisons of molecular phylogenies across multiple taxa tell us about the evolution of diversity in present day communities?, Philosophical Transactions of the Royal Society London, B 359, 1151-1571, Ladiges P Y and Udovicic F 2005, Comment on molecular dating of the age of eucalypts, Australian Systematic Botany, 18, 483-487, Boland D J, Brooker M I H, Chippendale G M, Hall N, Hyland B P M, Johnston R D, Kleinig D A, McDonald M W and Turner J D 2006, Forest Trees of Australia, CSIRO, Melbourne, Gandolpho M A, Hermsen E J, Zamaloa M C, Nixon K C, González C C, Wilf P, Cúneo N R and Johnson K R 2011, Oldest known Eucalyptus fossils are from South America, PloS ONE 6(6):e21084 doi:10.1371/journal.pone.0021084. Epub. June 28, 2011.

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Geological time line with reference to some eucalypts (Mya = million years ago; Kya = thousand years ago) 90 – 65 Mya: 70 Mya

65 – 55 Mya 60 Mya 51.9 Mya 65 – 35 Mya 40 – 45 Mya 40 Mya 45 – 20 Mya 30 - 20 Mya 30 – 13 Mya 25 – 10 Mya 10 – 5 Mya 5 – 2 Mya

2.58 Mya 2.6 – 0 Mya 150 - 30 Kya 60 Kya 25 Kya

10 - 8 Kya

Myrtaceae migrate into Australasian region of Gondwana from Antarctica (myrtaceous-like pollen found in sediments of this age from the Antarctic peninsula) Indo-Australian plate collides with the Eurasian plate, Gondwana sheds South America, Africa, New Zealand and India (Argentinian eucalypt fossils indicate a proto-eucalypt had already migrated into the South American part of Gondwana from Antarctica) Eucalypt lineage diverges early within the subfamily Lepidospermoideae Rapid radiation of myrtaceous genera begins Earliest fossil record of myrtaceous pollen in Australia and New Zealand Two major lineages – Angophora/Corymbia and Eucalyptus diverge Date of macrofossil of Patagonian symphyomyrt eucalypt, Chubut Province, Argentina — affinities to E. brachyandra (currently included in sub-genus Minutifructus with E. deglupta, E. howittiana and E. raveretiana), E. guilfolyei, and E. microcorys Oldest eucalypt macrofossils from southeast Queensland — affinities to Angophora/Corymbia Australian plate begins to move northwards Last separation of Australian plate from Antarctica New Guinea begins to form along the northern edge of the Australian plate Estimated age of subgenus Minutifructus (and probable ancient symphyomyrt ancestor of E. deglupta) Sclerophyllous eucalypts begin to appear in the fossil record Sections within subgenus Symphyomyrtus diverge Diversification within sections of eucalypts coincides with a drier and more seasonal climate on the Australian continent E. deglupta evolves from a symphyomyrt ancestor after differentiation from other symphyomyrt sub-genera and moves in association with tropical rainforest in the northern parts of New Guinea while the latter is still attached on the northern edge the Australian plate Fossil pollen records suggest modern eucalypt flora did not become widespread until during this time. Onset of severe aridity occurs on the Australian continent. Rapid uplift of the central New Guinea massif isolates the geologically older E. deglupta on its northern side still associated with rainforest. E. deglupta thus retains some primitive ancestral and rainforest-like characters compared to other eucalypts isolated to the south and evolving in in a drier climate Last ice age begins leading to glaciation on the central New Guinea massif Further speciation and diversification of eucalypts in the south in an unstable climate Sea level dramatically rises and falls, at times up to 200 m lower than today. Tasmania connected to the Australian mainland; existing Bass Straight Islands formed hills in the Bassean Plain.

Humans arrive in Australia (possibly earlier) and in New Guinea?

The sea level dropped to some 135 m lower than today in another significant fall in sea level between 30 and 15 Kya. Australia and New Guinea were connected in a fused landmass called Sahul. Sahul was still separated from Southeast Asia (called Sunda) by a deep channel in which many small islands developed (collectively called Wallacea), some of which are there today. Wallacea was a natural barrier to plant and land animal migration between Sahul and Sunda for many thousands of years, though when sea levels were low the widths of water gaps were considerably smaller. The Indonesian islands east as far as Borneo and Bali were connected to Sunda. Palawan was also part of Sunda, while the rest of the present day Philippine islands including Mindanao formed one large land mass separated from the continental plates north and south by the Sibutu Passage and Mindoro Strait. Recurring opportunities for E. deglupta to migrate from New Guinea west to Seram and Sulawesi and north to Mindanao on disturbed areas such as recent lava flows and older volcanic soils next to intact rainforest through long-distance seed dispersal aided by volcanic updrafts.

New Guinea disconnects from the Australian mainland

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25,000 years ago, the sea level was some 135 m lower than what it is today. Australia and New Guinea were connected in a fused landmass called Sahul. Sahul was still separated from Sunda (Southeast Asia)

Around 9,000 years ago New Guinea was disconnecting from the Australian mainland. (Both illustrations sourced from an animation of Sahul sea levels from 100 Kya to the present day at http://sahultime.monash.edu.au/explore.html (accessed 15 April 2019, but not available in 2021).

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The molecular phylogeny with best representation of species of Eucalyptus suggests that large stature species like E. deglupta have evolved at least four times within subgenus Symphyomyrtus (with independent evolution in E. diversicolor, E. deglupta and E. grandis).

The rainbow eucalypt appears to have originated from an ancestral eucalypt stock in the rainforests of Antarctic Gondwana. After the Australian continental plate began its northern migration (see timeline), New Guinea also began to form, initially in two parts, one on the northern rim of the Australian plate (today’s New Guinea mainland), the other part as string of islands off the northeast coast (today the Bismarck Archipelago, including New Britain). Molecular evidence suggests E. deglupta evolved from a symphyomyrt ancestor after differentiation from other sub-genera and moved into tropical rainforest in the northern parts of New Guinea while the latter was still attached on the northern edge the Australian plate. It was carried northwards, riding like a surfer on the northern rim of the Australian plate and staying ahead of the areas increasingly subjected to a drying climate to the south, with the rapid uplift of the central New Guinea massif creating the southern boundary.

Accelerated uplift of the central New Guinea massif isolated E. deglupta on its northern side still associated with rainforest. E. deglupta thus retained some primitive ancestral and rainforest-like characters compared to other eucalypts to the south evolving in in a drier climate. Left in its wake to the south were the progenitors of its related present-day species on the Australian mainland (E. brachyandra in north-eastern Western Australia (Kimberly region) and north-western Northern Territory, E. howittiana in north-eastern Queensland and E. raveretiana in central-eastern Queensland).

The migration of E. deglupta to the west and north of New Guinea probably was assisted by a combination of land bridges among the islands of Wallacea during periods of low sea level, by rafting on continental fragments and by dispersal of the very small winged seeds in volcanic updrafts along the western edge of the Pacific plate with its arc of active volcanoes.

The four present-day disjunct localities in the highlands of New Guinea are probably relictual, having been rapidly uplifted on the northern side of the central cordillera in Plio-Pleistocene times, surviving just below the edge of the ice sheet that formed during the last glacial maximum. E. deglupta occurs at about 2,000 metres elevation in the western highlands of Papua near Lake Paniai — this location is less than 200 km west of the receding remnant ice-age glaciers that still exist today in the mountainous region surrounding Puncak Jaya (4,884 m elevation). The morphology of the present-day highland occurrences is somewhat different to the others having wider leaves with less pronounced drip tips (more like the variety schlechteri suggested elsewhere), larger fruits with the valves not prominently exsert, ovary 357

inferior, not half inferior and number of valves four and occasionally five or six compared with the usual three or four, possibly indicating some change in the direction of evolution in response to a cooler, drier, montane environment. The species is now extinct in the total geographic range of all other extant eucalypts and is not found in the rainforests of the small remaining humid tropical area in northeast Queensland or in rainforests south of the central massif in New Guinea. The isolated northerly mainland New Guinea occurrences of E. deglupta in some places almost abut, but do not overlap anywhere, the southerly mainland New Guinea occurrences of all the other extra-Australian eucalypts which arrived there in more recent geological time.27 In a collaborative project undertaken by Royal Botanic Gardens Melbourne (RBGM), the University of Melbourne and Bogor Botanic Gardens Indonesia and supervised by Frank Udovicic (RGBM) a team has been undertaking analyses of the DNA of E. deglupta using a number of markers on 75 specimens from across its range. The conclusion so far is that the New Britain samples are quite distinct from the Seram, Sulawesi and Philippine samples and that E. deglupta from Seram, Sulawesi and Philippines will form one major clade and the New Britain samples will form a separate major clade.

ITS and ETS sequences have been generated from the same 75 specimens for phylogenetic analysis to clarify the position of E. deglupta within Eucalyptus. The team has been assembling alignments for the sequence markers and incorporating microsatellite data.

Publication of this work was originally expected towards the end of 201928 but is still awaited (October 2022).

27

Carr S G M 1972 Problems of the geography of tropical eucalypts. In Bridge and barrier: the natural and cultural history of Torres Strait. Walker D (ed.) ANU Publication no. BG/3, pp 153-181. Australian National University, Canberra; Payne K G, Dvorak W S and Myburg A 2007 Chloroplast DNA phylogeography reveals the island colonization route of Eucalyptus urophylla (Myrtaceae). Australian Journal of Botany 55:673-683.

28 Frank Udovicic personal communication June 2019.

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Eucalyptus deglupta plantations, Open Bay, New Britain, 2018. (© 2018, 2019 Google LLC, used with permission.)

359

Keravat township, “our” house is still there (near tip of arrow) 50 years after we lived in it, but the old forestry office, forest nursery and 1948 kamarere plantation that was behind the house are of course all long gone.

Keravat River

Keravat 2019. Forestry plantation and silvicultural activities by the PNG Forest Department ceased in 1973, after which the area west of the Keravat River and either side of the Kalabus Road that used to be teak and kamarere plantations has been planted up with oil palm. (Base image: 2019 Google LLC, used with permission. Annotations by J Davidson 2020)

Kalabus

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ABBREVIATIONS AND ACRONYMS A/OIC Acting Officer-in-Charge ACIAR Australian Council for International Agricultural Research (includes Forestry) AFC

Australian Forestry Council

ANU

Australian National University

APM

Australian Pulp and Paper Manufacturers

Appita Technical Association of the Australian and New Zealand Pulp and Paper Industry Inc. C

Centigrade

CERES Controlled Environment RESearch CFI

Commonwealth Forestry Institute (Oxford)

cm(s)

Centimeter(s)

cm2

Square centimetre(s)

CSIRO Commonwealth Scientific and Industrial Research Organization (Australia) CTFT

Centre Technique Forestier Tropicale (France and Congo)

D

Diameter

DBH

Diameter (of a tree) at Breast Height (or 1.3 metres)

DNA

Deoxyribonucleic Acid

DOE

Department of Energy (USA)

Dr

Doctor

EAPI

Environment and Policy Institute (of the EWC)

EWC

East-West Centre (Honolulu)

FA

Factor analysis

FAO

Food and Agriculture Organisation of the United Nations

FCNSW Forest Commission of NSW FRI

Forest Research Institute (Canberra Australia or Lae PNG)

FSC

Forest Stewardship Council

H

Height

ha

hectare(s)

HPH

Hak Pengus Hutan (Timber Concession, Indonesian)

HQ

Headquarters

IBRD

International Bank for Reconstruction and Development

IDA

International Development Agency (of the World Bank)

IPEF

(Instituto de Pesquisas e Estudos Florestais [Forest Science and Research Institute]

ITCI

Industrial Timber Corporation of Indonesia 361

ITS

Internal Transcribed Spacer

IUFRO International Union of Forest Research Organizations JANT

Japan New Guinea Timbers

JGI

Joint Genome Institute (of the DOE)

kg

Kilogram

LA

Logging Area

LAES

Lowlands Agriculture Experiment Station

M

Metre

MAI

Mean Annual Increment

mm

Millimetre

m3

Cubic metre(s)

Mya

Million years ago

Myr

Million years

NCBI

National Centre for Biotechnology Information (Bethesda, Maryland, USA)

NSSC

Neutral sulphite semichemical (wood pulp)

NSW

New South Wales

OBT

Open Bay Timber (limited)

OIC

Officer-in-Charge

PCA

Principal component analysis

PhD

Doctor of Philosophy Degree

PICOP Paper Industries Corporation of the Philippines PNG

Papua New Guinea

PNGUT The Papua New Guinea University of Technology Prof

Professor

RAAF Royal Australian Air Force RFO

Regional Forest Officer

RGBM Royal Botanic Gardens Melbourne RH

Rijks Herbarium

RWG

Research Working Group (of the AFC)

SADF South African Department of Forestry SE

Southeast

TAA

Trans Australia Airlines

TRP

Timber Rights Purchase

UK

United Kingdom

UNDP United Nations Development Programme 362

USA

United States of America

V

Volume

v/v

Volume to Volume

WA

Western Australia

WGD

Whole-genome duplication

WW I

World War One

WW II World War Two yr

Year

♀

Female

♂

Male

363