Achievements in forest tree improvement in Australia and New Zealand 6: Genetic improvement and conservation of Araucaria cunninghamii in Queensland

Araucaria cunninghamii (hoop pine) typically occurs as an emergent tree over subtropical and tropical rainforests, in a discontinuous distribution that extends from West Irian Jaya at about 0°30'S, through the highlands of Indonesian New Guinea and Papua New Guinea, along the east coast of Australia from 11°39'S in Queensland to 30°35'S in northern New South Wales. Plantations established in Queensland since the 1920s now total about 44000 ha, and constitute the primary source for the continuing supply of hoop pine quality timber and pulpwood, with a sustainable harvest exceeding 440 000 m3 y-1. Establishment of these managed plantations allowed logging of all native forests of Araucaria species (hoop pine and bunya pine, A. bidwillii) on state-owned lands to cease in the late 1980s, and the preservation of large areas of araucarian forest types within a system of state-owned and managed reserves. The successful plantation program with this species has been strongly supported by genetic improvement activities since the late 1940s - through knowledge of provenance variation and reproductive biology, the provision of reliable sources of improved seed, and the capture of substantial genetic gains in traits of economic importance (for example growth, stem straightness, internode length and spiral grain). As such, hoop pine is one of the few tropical tree species that, for more than half a century, has been the subject of continuous genetic improvement. The history of commercialisation and genetic improvement of hoop pine provides an excellent example of the dual economic and conservation benefits that may be obtained in tropical tree species through the integration of gene conservation and genetic improvement with commercial plantation development. This paper outlines the natural distribution and reproductive biology of hoop pine, describes the major achievements of the genetic improvement program in Queensland over the past 50+ y, summarises current understanding of the genetic variation and control of key selection traits, and outlines the means by which genetic diversity in the species is being conserved.

Quantitative and molecular genetics of juvenile wood traits in radiata and slash/Caribbean pines.

The Juvenile Wood Initiative (JWI) project has been running successfully since July 2003 under a Research Agreement with FWPA and Letters of Association with the consortium partners STBA (Southern Tree Breeding Association), ArborGen and FPQ (Forestry Plantations Queensland). Over the last five and half years, JWI scientists in CSIRO, FPQ, and STBA have completed all 12 major milestones and 28 component milestones according to the project schedule. We have made benchmark progress in understanding the genetic control of wood formation and interrelationships among wood traits. The project has made 15 primary scientific findings and several results have been adopted by industry as summarized below. This progress was detailed in 10 technical reports to funding organizations and industry clients. Team scientists produced 16 scientific manuscripts (8 published, 1 in press, 2 submitted, and several others in the process of submission) and 15 conference papers or presentations. Primary Scientific Findings. The 15 major scientific findings related to wood science, inheritance and the genetic basis of juvenile wood traits are: 1. An optimal method to predict stiffness of standing trees in slash/Caribbean pine is to combine gravimetric basic density from 12 mm increment cores with a standing tree prediction of MoE using a time of flight acoustic tool. This was the most accurate and cheapest way to rank trees for breeding selection for slash/Caribbean hybrid pine. This method was also recommended for radiata pine. 2. Wood density breeding values were predicted for the first time in the STBA breeding population using a large sample of 7,078 trees (increment cores) and it was estimated that selection of the best 250 trees for deployment will produce wood density gains of 12.4%. 3. Large genetic variation for a suite of wood quality traits including density, MFA, spiral grain, shrinkage, acoustic and non-acoustic stiffness (MoE) for clear wood and standing trees were observed. Genetic gains of between 8 and 49% were predicted for these wood quality traits with selection intensity between 1 to 10% for radiata pine. 4. Site had a major effect on juvenile-mature wood transition age and the effect of selective breeding for a shorter juvenile wood formation phase was only moderate (about 10% genetic gain with 10% selection intensity, equivalent to about 2 years reduction of juvenile wood). 5. The study found no usable site by genotype interactions for the wood quality traits of density, MFA and MoE for both radiata and slash/Caribbean pines, suggesting that assessment of wood properties on one or two sites will provide reliable estimates of the genetic worth of individuals for use in future breeding. 6. There were significant and sizable genotype by environment interactions between the mainland and Tasmanian regions and within Tasmania for DBH and branch size. 7. Strong genetic correlations between rings for density, MFA and MoE for both radiata and slash/Caribbean pines were observed. This suggests that selection for improved wood properties in the innermost rings would also result in improvement of wood properties in the subsequent rings, as well as improved average performance of the entire core. 8. Strong genetic correlations between pure species and hybrid performance for each of the wood quality traits were observed in the hybrid pines. Parental performance can be used to identify the hybrid families which are most likely to have superior juvenile wood properties of the slash/Caribbean F1 hybrid in southeast Queensland. 9. Large unfavourable genetic correlations between growth and wood quality traits were a prominent feature in radiata pine, indicating that overcoming this unfavourable genetic correlation will be a major technical issue in progressing radiata pine breeding. 10. The project created the first radiata pine 18 k cDNA microarray and generated 5,952 radiata pine xylogenesis expressed sequence tags (ESTs) which assembled into 3,304 unigenes. 11. A total of 348 genes were identified as preferentially expressed genes in earlywood or latewood while a total of 168 genes were identified as preferentially expressed genes in either juvenile or mature wood. 12. Juvenile earlywood has a distinct transcriptome relative to other stages of wood development. 13. Discovered rapid decay of linkage disequilibrium (LD) in radiata pine with LD decaying to approximately 50% within 1,700 base pairs (within a typical gene). A total of 913 SNPS from sequencing 177,380 base pairs were identified for association genetic studies. 14. 149 SNPs from 44 genes and 255 SNPs from a further 51 genes (total 95 genes) were selected for association analysis with 62 wood traits, and 30 SNPs were shortlisted for their significant association with variation of wood quality traits (density, MFA and MoE) with individual significant SNPs accounting for between 1.9 and 9.7% of the total genetic variation in traits. 15. Index selection using breeding objectives was the most profitable selection method for radiata pine, but in the long term it may not be the most effective in dealing with negative genetic correlations between wood volume and quality traits. A combination of economic and biological approaches may be needed to deal with the strong adverse correlation.

Association genetics in Corymbia citriodora subsp variegata identifies single nucleotide polymorphisms affecting wood growth and cellulosic pulp yield

Wood is an important biological resource which contributes to nutrient and hydrology cycles through ecosystems, and provides structural support at the plant level. Thousands of genes are involved in wood development, yet their effects on phenotype are not well understood. We have exploited the low genomic linkage disequilibrium (LD) and abundant phenotypic variation of forest trees to explore allelic diversity underlying wood traits in an association study. Candidate gene allelic diversity was modelled against quantitative variation to identify SNPs influencing wood properties, growth and disease resistance across three populations of Corymbia citriodora subsp. variegata, a forest tree of eastern Australia. Nine single nucleotide polymorphism (SNP) associations from six genes were identified in a discovery population (833 individuals). Associations were subsequently tested in two smaller populations (130160 individuals), validating our findings in three cases for actin 7 (ACT7) and COP1 interacting protein 7 (CIP7). The results imply a functional role for these genes in mediating wood chemical composition and growth, respectively. A flip in the effect of ACT7 on pulp yield between populations suggests gene by environment interactions are at play. Existing evidence of gene function lends strength to the observed associations, and in the case of CIP7 supports a role in cortical photosynthesis.

Nocturnal Perching of Scarabaeine Dung Beetles (Coleoptera, Scarabaeidae) in an Australian Tropical Rain Forest

Wongabel, a northeastern Queensland tropical, wet, evergreen forest, contains 22 species of Scarabaeinae dung beetles. Five of these species were observed to perch commonly on leaves at night. Length of the beetle and the height of its perch were recorded for each of 56 1 specimens. Unlike the New World tropical dung beetle perchers, no clear evidence was found that small species perched closer to the ground than larger species. The evidence gathered, at least for the four most common perchers, supports the hypothesis that perching is one type of foraging strategy. The similarities and differences between the Australian and New World perchers are discussed.

The risk of pollen-mediated gene flow from exotic Corymbia plantations into native Corymbia populations in Australia

Sectors of the forest plantation industry in Australia are set to expand in the near future using species or hybrids of the spotted gums (Corymbia, Section Politaria). Plantations of these taxa have already been introduced across temperate and subtropical Australia, representing locally exotic introductions from native stands in Queensland and New South Wales. A literature review was undertaken to provide insights into the potential for pollen-mediated gene flow from these plantations into native populations. Three factors suggest that such gene flow is likely; (1) interspecific hybridisation within the genus has frequently been recorded, including between distantly related species from different sections, (2) apparent high levels of vertebrate pollinator activity may result in plantation pollen being moved over hundreds of kilometres, (3) much of the plantation estate is being established among closely related taxa and therefore few barriers to gene flow are expected. Across Australia, 20 of the 100 native Corymbia taxa were found to have regional level co-occurrence with plantations. These were located most notably within regions of north-east New South Wales and south-east Queensland, however, co-occurrence was also found in south-west Western Australia and eastern Victoria. The native species found to have co-occurrence were then assessed for the presence of reproductive barriers at each step in the process of gene flow that may reduce the number of species at risk even further. The available data suggest three risk categories exist for Corymbia. The highest risk was for gene flow from plantations of spotted gums to native populations of spotted gums. This was based on the expected limited existence of pre- and post-zygotic barriers, substantial long-distance pollen dispersal and an apparent broad period of flowering in Corymbia citriodora subsp. variegata plantations. The following risk category focussed on gene flow from Corymbia torelliana × C. c. variegata hybrid plantations into native C. c. variegata, as the barriers associated with the production and establishment of F1 hybrids have been circumvented. For the lowest risk category, Corymbia plantations may present a risk to other non-spotted gum species, however, further investigation of the particular cross-combinations is required. A list of research directions is provided to better quantify these risks. Empirical data will need to be combined within a risk assessment framework that will not only estimate the likelihood of exotic gene flow, but also consider the conservation status/value of the native populations. In addition, the potential impacts of pollen flow from plantations will need to be weighed up against their various economic and environmental benefits.

Developing African mahogany (Khaya senegalensis) germplasm and its management for a sustainable forest plantation industry in northern Australia: Progress and needs.

The demonstrated wide adaptability, substantial yield potential and proven timber quality of African mahogany (Khaya senegalensis) from plantings of the late 1960s and early 1970s in northern Australia have led to a resurgence of interest in this high-value species. New plantations or trials have been established in several regions since the early 1990s -in four regions in north Queensland, two in the Northern Territory and one in Western Australia. Overall, more than 1500 ha had been planted by early 2007, and the national annual planting from 2007-2008 as currently planned will exceed 2400 ha. Proceedings of two workshops have summarised information available on the species in northern Australia, and suggested research and development (R&D) needs and directions. After an unsustained first phase of domestication of K. senegalensis in the late 1960s to the early 1970s, a second phase began in northern Australia in 2001 focused on conservation and tree improvement that is expected to provide improved planting stock by 2010. Work on other aspects of domestication is also described in this paper: the current estate and plans for extension; site suitability, soils and nutrition; silviculture and management; productivity; pests and diseases; and log and wood properties of a sample of superior trees from two mature plantations of unselected material near Darwin. Some constraints on sustainable plantation development in all these fields are identified and R&D needs proposed. A sustained R&D effort will require a strategic coordinated approach, cooperative implementation and extra funding. Large gains in plantation profitability can be expected to flow from such inputs.

Runoff, sediment loss and water quality from forest roads in a southeast Queensland coastal plain Pinus plantation

Runoff and sediment loss from forest roads were monitored for a two-year period in a Pinus plantation in southeast Queensland. Two classes of road were investigated: a gravelled road, which is used as a primary daily haulage route for the logging area, and an ungravelled road, which provides the main access route for individual logging compartments and is intensively used as a haulage route only during the harvest of these areas (approximately every 30 years). Both roads were subjected to routine traffic loads and maintenance during the study. Surface runoff in response to natural rainfall was measured and samples taken for the determination of sediment and nutrient (total nitrogen, total phosphorus, dissolved organic carbon and total iron) loads from each road. Results revealed that the mean runoff coefficient (runoff depth/rainfall depth) was consistently higher from the gravelled road plot with 0.57, as compared to the ungravelled road with 0.38. Total sediment loss over the two-year period was greatest from the gravelled road plot at 5.7 t km−1 compared to the ungravelled road plot with 3.9 t km−1. Suspended solids contributed 86% of the total sediment loss from the gravelled road, and 72% from the ungravelled road over the two years. Nitrogen loads from the two roads were both relatively constant throughout the study, and averaged 5.2 and 2.9 kg km−1 from the gravelled and ungravelled road, respectively. Mean annual phosphorus loads were 0.6 kg km−1 from the gravelled road and 0.2 kg km−1 from the ungravelled road. Organic carbon and total iron loads increased in the second year of the study, which was a much wetter year, and are thought to reflect the breakdown of organic matter in roadside drains and increased sediment generation, respectively. When road and drain maintenance (grading) was performed runoff and sediment loss were increased from both road types. Additionally, the breakdown of the gravel road base due to high traffic intensity during wet conditions resulted in the formation of deep (10 cm) ruts which increased erosion. The Water Erosion Prediction Project (WEPP):Road model was used to compare predicted to observed runoff and sediment loss from the two road classes investigated. For individual rainfall events, WEPP:Road predicted output showed strong agreement with observed values of runoff and sediment loss. WEPP:Road predictions for annual sediment loss from the entire forestry road network in the study area also showed reasonable agreement with the extrapolated observed values.

Construction of microsatellite linkage maps for Corymbia

The genus Corymbia is closely related to the genus Eucalyptus, and like Eucalyptus contains tree species that are important for sub-tropical forestry. Corymbia's close relationship with Eucalyptus suggests genetic studies in Corymbia should benefit from transfer of genetic information from its more intensively studied relatives. Here we report a genetic map for Corymbia spp. based on microsatellite markers identified de novo in Corymbia sp or transferred from Eucalyptus. A framework consensus map was generated from an outbred F 2 population (n = 90) created by crossing two unrelated Corymbia torelliana x C. citriodora subsp. variegata F1 trees. The map had a total length of 367 cM (Kosambi) and was composed of 46 microsatellite markers distributed across 13 linkage groups (LOD 3). A high proportion of Eucalyptus microsatellites (90%) transferred to Corymbia. Comparative analysis between the Corymbia map and a published Eucalyptus map identified eight homeologous linkage groups in Corymbia with 13 markers mapping on one or both maps. Further comparative analysis was limited by low power to detect linkage due to low genome coverage in Corymbia, however, there was no convincing evidence for chromosomal structural differences because instances of non-synteny were associated with large distances on the Eucalyptus map. Segregation distortion was primarily restricted to a single linkage group and due to a deficit of hybrid genotypes, suggesting that hybrid inviability was one factor shaping the genetic composition of the F2 population in this inter-subgeneric hybrid. The conservation of microsatellite loci and synteny between Corymbia and Eucalyptus suggests there will be substantial value in exchanging information between the two groups.

Genetic control of propagation traits in a single Corymbia torelliana x Corymbia variegata family

Genetic control of vegetative propagation traits was described for a second-generation, outbred, intersectional hybrid family (N = 208) derived from two species, Corymbia torelliana (F. Muell.) K.D. Hill & L.A.S. Johnson and Corymbia variegata (F. Muell.) K.D. Hill & L.A.S. Johnson, which contrast for propagation characteristics and in their capacity to develop lignotubers. Large phenotypic variances were evident for rooting and most other propagation traits, with significant proportions attributable to differences between clones (broad-sense heritabilities 0.2-0.5). Bare root assessment of rooting rate and root quality parameters tended to have the highest heritabilities, whereas rooting percentage based on root emergence from pots and shoot production were intermediate. Root biomass and root initiation had the lowest heritabilities. Strong favourable genetic correlations were found between rooting percentage and root quality traits such as root biomass, volume, and length. Lignotuber development on a seedling was associated with low rooting and a tendency to poor root quality in cuttings and was in accord with the persistence of species parent types due to gametic phase disequilibrium. On average, nodal cuttings rooted more frequently and with higher quality root systems, but significant cutting type x genotype interaction indicated that for some clones, higher rooting rates were obtained from tips. Low germination, survival of seedlings, and rooting rates suggested strong hybrid breakdown in this family.

Light environment and tree development of young Acacia melanoxylon in mixed-species regrowth forest, Tasmania, Australia

Blackwood (Acacia melanoxylon R. Br.) is a valuable leguminous cabinetwood species which is commonly found as a canopy or subcanopy tree in a broad range of mixed-species moist forests on tablelands and coastal escarpments in eastern Australia. This paper reports on the competitive light environment of a commercially valuable multi-species regrowth forest in NW Tasmania, in order to define some of the functional interactions and competitive dynamics of these stands. Comparative observations were made of the internal forest light environment in response to small-gap silvicultural treatments, in a young regenerative mix of three codominant tree species. Light measurements were made during periods of maximum external irradiance of the regrowth Eucalyptus obliqua/A. melanoxylon forest canopy at age 10.5 years. This was at a time of vigourous stand development, 4.5 years following the application of three experimental silvicultural treatments whose effects were observed in comparison with an untreated canopy sample designed as a control. Minimal irradiance was observed within and beneath the dense subcanopy of the native nurse species (Pomaderris apetala) which closely surrounds young blackwood regeneration. Unlike current plantation nurse systems, the dense foliage of the native broadleaved Pomaderris all but eliminated direct side-light and low-angle illumination of the young blackwood, from the beginning of tree establishment. The results demonstrated that retention of these densely stocked native codominants effectively suppressed both size and frequency of blackwood branches on the lower bole, through effective and persistent interception of sunlight. Vigorous young blackwood crowns later overtopped the codominant nurse species, achieving a predictable height of branch-free bole. This competitive outcome offers a valuable tool for management of blackwood crown dynamics, stem form and branch habit through manipulation of light environment in young native regrowth systems. Results demonstrate that effective self-pruning in the lower bole of blackwood is achieved through a marked reduction in direct and diffuse sunlight incident on the lower crown, notably to less than 10-15% of full sunlight intensity during conditions of maximum insolation. The results also contain insights for the improved design of mixed-species plantation nurse systems using these or functionally similar species' combinations. Based on evidence presented here for native regrowth forest, plantation nurse systems for blackwood will need to achieve 85-90% interception of external side-light during early years of tree development if self-pruning is to emulate the results achieved in the native nurse system.

Spatial model of site index based on Y-ray spectrometry and a digital elevation model for two Pinus species in Tuan Toolara State Forest, Queensland, Australia

Site index prediction models are an important aid for forest management and planning activities. This paper introduces a multiple regression model for spatially mapping and comparing site indices for two Pinus species (Pinus elliottii Engelm. and Queensland hybrid, a P. elliottii x Pinus caribaea Morelet hybrid) based on independent variables derived from two major sources: g-ray spectrometry (potassium (K), thorium (Th), and uranium (U)) and a digital elevation model (elevation, slope, curvature, hillshade, flow accumulation, and distance to streams). In addition, interpolated rainfall was tested. Species were coded as a dichotomous dummy variable; interaction effects between species and the g-ray spectrometric and geomorphologic variables were considered. The model explained up to 60% of the variance of site index and the standard error of estimate was 1.9 m. Uranium, elevation, distance to streams, thorium, and flow accumulation significantly correlate to the spatial variation of the site index of both species, and hillshade, curvature, elevation and slope accounted for the extra variability of one species over the other. The predicted site indices varied between 20.0 and 27.3 m for P. elliottii, and between 23.1 and 33.1 m for Queensland hybrid; the advantage of Queensland hybrid over P. elliottii ranged from 1.8 to 6.8 m, with the mean at 4.0 m. This compartment-based prediction and comparison study provides not only an overview of forest productivity of the whole plantation area studied but also a management tool at compartment scale.

Forest health surveillance methodology in hardwood plantations in Queensland, Australia

Forest health surveillance (FHS) of hardwood plantations commenced in Queensland in 1997 as plantations expanded following a state government planting initiative arising from the national 2020 forest policy vision. The estate was initially characterised by a large number of small plantations (10-50 ha), although this has changed more recently with the concentration of larger plantations in the central coast and South Burnett regions. Due to the disparate nature of the resource, drive- and walkthrough surveys of subsets of plantations have been undertaken in preference to aerial surveys. FHS has been effective in detecting a number of new hardwood pests in Queensland including erinose mites (Rhombacus and Acalox spp.), western white gum plate galler (Ophelimus sp.), Creiis psyllid and bronzing bug (Thaumastocoris sp.), in evaluating their potential impact and assisting in focussing future research efforts. Since 2003 there has been an increased emphasis on training operational staff to take a greater role in identifying and reporting on forest health issues. This has increased their awareness of forest health issues, but their limited time to specifically survey and report on pests and diseases, and high rates of staff turnover, necessitate frequent ongoing training. Consequently, common and widespread problems such as quambalaria shoot blight (Quambalaria pitereka), chrysomelid leaf beetles (mainly Paropsis atomaria) and erinose mites may be under-reported or not reported, and absence data may often not be recorded at all. Comment is made on the future directions that FHS may take in hardwood plantations in Queensland.

Genetic variation within two sympatric spotted gum eucalypts exceeds between taxa variation

Population substructure and hybridization, among other factors, have the potential to cause erroneous associations in linkage disequilibrium (LD) mapping. Two closely related spotted gum eucalypts, Corymbia variegata and C. henryi (Myrtaceae) occur in sympatry in the east coast of Australia and potentially interbreed. They are morphologically similar but are distinguished as separate species based on capsule and foliage size. To determine whether they hybridize in nature and its implications for LD mapping, we investigated the level of molecular divergence between the two species at two sympatric locations separated by 300 kilometres. Very few individuals of intermediate morphology were identified, despite the two species occurring only metres apart. Analysis of genetic structure using 12 microsatellite loci showed that genetic differentiation between populations of the same species at different locations (FST = 0.07 for both species; p = 0.0001) was significantly higher than that observed between species at each location (mean FST = 0.02 and 0.04 for Cherry tree and Bunyaville respectively; p = 0.0001; all Mann-Whitney U-test p ≤ 0.01). No species-specific alleles or significant allele frequency differences were detected within a site, suggesting recurrent local gene flow between the two species. The lack of significant allele frequency differences implies no population stratification along taxonomic lines. This suggested that there is little concern for cryptic hybridization when sampling from sites of sympatry for LD mapping.

Development of hazard site surveillance programs for forest invasive species: A case study from Brisbane, Australia

Hazard site surveillance is a system for post-border detection of new pest incursions, targeting sites that are considered potentially at high risk of such introductions. Globalisation, increased volumes of containerised freight and competition for space at domestic ports means that goods are increasingly being first opened at premises some distance from the port of entry, thus dispersing risk away from the main inspection point. Hazard site surveillance acts as a backstop to border control to ensure that new incursions are detected sufficiently early to allow the full range of management options, including eradication and containment, to be considered. This is particularly important for some of the more cryptic forest pests whose presence in a forest often is not discovered until populations are already high and the pest is well established. General requirements for a hazard site surveillance program are discussed using a program developed in Brisbane, Australia, in 2006 as a case study. Some early results from the Brisbane program are presented. In total 67 species and 5757 individuals of wood-boring beetles have been trapped and identified during the program to date. Scolytines are the most abundant taxa, making up 83% of the catch. No new exotics have been trapped but 19 of the species and 60% of all specimens caught are exotics that are already established in Australia.

Population genetics of the frog-killing fungus Batrachochytrium dendrobatidis

Global amphibian decline by chytridiomycosis is a major environmental disaster that has been attributed to either recent fungal spread or environmental change that promotes disease. Here, we present a population genetic comparison of Batrachochytrium dendrobatidis isolates from an intensively studied region of frog decline, the Sierra Nevada of California. In support of a novel pathogen, we find low diversity, no amphibian-host specificity, little correlation between fungal genotype and geography, local frog extirpation by a single fungal genotype, and evidence of human-assisted fungus migration. In support of endemism, at a local scale, we find some diverse, recombining populations. Therefore neither epidemic spread nor endemism alone explains this particular amphibian decline. Recombination raises the possibility of resistant sporangia and a mechanism for rapid spread as well as persistence that could greatly complicate global control of the pathogen.