Native and restored forests are increasingly impacted by pests and diseases, including large herbivores. While community- and species-level impacts of these tree enemies are often well-documented, there is little understanding of their influence on finer-scale eco-evolutionary processes. We here study the influence of large-mammal herbivory on the survival and height growth of trees in a mixed species restoration planting of the Australian forest trees, Eucalyptus ovata and E. pauciflora, in Tasmania, Australia. Common-garden field trials mixing the two species were compared in adjacent unbrowsed (fenced) and browsed (unfenced) plantings. The browsed planting was exposed to mammal browsing by native marsupials, as well as feral introduced European fallow deer (Dama dama). Each tree species was represented by open-pollinated families from 22 paired geographic areas, allowing the assessment of the effects of browsing on the species and population differences, as well as on family variation within each species. In the browsed planting, a marked reduction in species and population differences, as well as in family variance, was observed for both height growth and survival. The pattern of height growth and survival of the populations of both species also differed between browsing regimes, with significant changes of climate relationships involving both focal tree attributes detected. Our results argue for a major disruption of the eco-evolutionary dynamics of restored forests in the presence of browsing by large mammalian herbivores, at the observed period of the tree life cycle. Importantly for forest restoration and conservation in the face of global change, our results challenge the choice of tree populations for translocation based solely on predicted or observed relationships of their home-site climate with current and predicted future climates of the restoration sites, while emphasising the need for genetic diversity to provide future resilience of restored forests to both biotic and abiotic stresses.

Myrtle rust caused by Austropuccinia psidii is an important disease of Myrtaceae species in tropical and subtropical regions of the world. The pathogen is of concern to the forest plantation industry in Brazil which is primarily based on introduced eucalypts from Australia. A. psidii has multiple biotypes, including a native Brazilian biotype, that causes damage to Eucalyptus spp. and Syzygium jambos, and a pandemic biotype. As an invader, the pandemic biotype affects numerous Myrtaceae hosts in countries around the world, including those in Australia. Although the pandemic biotype of A. psidii has had little impact on eucalypt plantations in Australia to date, this may change if the Brazilian or other biotypes/races invade, or if there is genetic change in the pandemic biotype in Australia. In Australian seedling inoculation studies, families from range-wide native stand seed collections of Australia’s main plantation eucalypt, Eucalyptus globulus, have shown considerable genetic variation in resistance to the pandemic biotype of A. psidii. In a seedling inoculation study undertaken in Brazil with a subset of these families, we similarly show significant genetic variation in host resistance to the common Brazilian race of A. psidii (race 1) at the family and population levels. The disease severity in the Brazilian and Australian studies was significantly correlated at the family level, ranging between 0.49 and 0.57, due mainly to genetic-based covariation in the immune response. Overall, the E. globulus families exhibited less resistance to the Brazilian than the pandemic biotype, but it is unclear whether this is due to greater pathogen virulence or different inoculation and growing conditions. Regardless, there is genetic variation in E. globulus susceptibility to both A. psidii biotypes, which would allow genetic improvement. Further, despite variation in the relative susceptibility of some families to the different biotypes, selection of E. globulus for reduced susceptibility to one biotype is expected to make positive gains in resistance to the other, and some families exhibited relatively ‘stable resistance’ with low susceptibility to both biotypes.

Long regarded as posing a low invasion risk, eucalypts are now increasingly recognized as invasive in certain contexts, leading to a surge of studies on this topic in recent years. We performed the first systematic literature review to analyse the invasion risk, the invasion performance and the drivers of eucalypt invasion, using information from 140 publications and 96 species. We also map the eucalypt area and the distribution of eucalypt research, worldwide. Eucalypts cover at least 30 million hectares globally, mostly in Brazil, India and China, but nearly half of the invasion studies were conducted in Portugal and South Africa, mostly on Eucalyptus globulus and E. camaldulensis, respectively. These two widely cultivated species, together with E. tereticornis, have the highest scores among eucalypt species, in invasion risk assessments. A dispersal model was built using data from 15 papers, leading to an estimated maximum dispersal distance of 110 m from mother plants. Soil disturbance, moisture and fire are important key drivers of invasion. Despite notable advances in the invasion ecology of eucalypts in recent years, research biases, context dependency, and persistent knowledge gaps remain challenges that must be addressed to achieve a comprehensive understanding of eucalypt invasion ecology.

Understanding how functional traits affect plant performance and fitness is a key step in unravelling the role of natural selection in shaping the evolutionary trajectory of populations. We examined early-age selection acting on leaf traits via their effects on growth performance and fitness, measured in Eucalyptus ovata trees planted in a common-garden field trial embedded in a reforestation planting in Tasmania, Australia. We focused on two important leaf traits - stomatal length and specific leaf area (SLA) - measured two years after planting, and compared interplanted E. ovata groups originating from dry and wet home-site climates, with the trial site having intermediate long-term mean annual rainfall. Two-year height growth was used as the performance attribute, and the time-averaged tree survival over the subsequent six years as the fitness component. There was evidence for performance-based selection on the leaf traits, with the strength and form of selection depending on the trait and climate group being considered. In this sense, selection in the dry group operated mainly on stomatal length where a combination of directional (favouring longer stomata) and stabilizing selection was detected, whereas selection in the wet group acted only on SLA and was purely stabilizing. Estimates of performance-based correlational selection were not statistically significant. For both climate groups, estimates of fitness-based selection gradients provided evidence for significant directional (but not quadratic) selection on height performance, favouring individuals with faster growth, but did not indicate statistical support for direct effects of the leaf traits on tree survival, conditional on measured performance. These results validated qualitative inferences of selection from the performance-based analysis, and suggested that selection on the leaf traits appeared to be mediated by their effects on early-age height performance, which in turn directly influenced later-age survival. We discuss the mechanisms by which the focal traits may have affected height performance, and likely factors contributing to the different patterns of phenotypic selection observed in the two groups experiencing the same environment. We also provide expressions of analytical derivatives that were developed for the estimation of selection gradients based on a logistic regression model relating a binary fitness response to linear and nonlinear covariate terms for the target regressor variables.

Internal checking is a wood-drying defect that affects many tree species, including eucalypts, and can adversely affect the recovery of high-quality timber. We studied the patterns of within- and between-tree variation in wood checking from three Eucalyptus nitens silvicultural trials and determined the predictability of wood checking from wood properties. We hypothesised that changed wind exposure and inter-tree competition due to thinning would increase wood checking. In total, 144 trees from two silvicultural treatments (thinned/unthinned) and three social status classes per site (dominant/subdominant/suppressed) were sampled. Regardless of site, social status and silvicultural treatment, longitudinal and radial variation within trees was the major source of variation in checking. Checking was concentrated longitudinally in bottom logs and radially in the middle of wedges, with a shift in checking towards the cambium in the thinned treatment. On average, dominant trees in the thinned treatment had higher levels of checking than dominant trees in unthinned treatments at all sites, but the opposite trend was found for suppressed trees. Wedge checking was poorly predicted using non-destructive traits assessed on standing trees. Our models accounted for at most 22% of the variation in wedge checking and the best predictors were the destructive measures of wedge collapse. With most checking occurring in lower logs, future studies should concentrate on reducing checking in bottom logs, which are typically pruned for clearwood products.

The genes ECERIFERUM1 (CER1) and ECERIFERUM3 (CER3) encode the biosynthesis of alkane waxes, a key component of the plant cuticle. To study the evolution of CER1 and CER3 in a highly diverse group of eucalypts, we performed a genome-wide survey using recently released genome assemblies of 28 Myrtaceae species, with 22 species from the main eucalypt lineage and 6 non-eucalypt Myrtaceae tree species. We manually annotated 250 genes and pseudogenes, identifying a near-ubiquitous single copy of CER3 and 2 to 10 CER1 gene copies per Myrtaceae species. Phylogenetic analysis suggested that copy number variation in eucalypts is due to multiple tandem duplication events, both ancient (shared by all Myrtaceae species studied) and relatively recent (present only in eucalypts). Inter-chromosomal translocations were discovered for both CER1 and CER3, along with recurrent loss of often the same CER1 introns in the WAX2 domain, the domain that is essential for wax production. Despite the varied environments occupied by the eucalypt species in this study, we did not find statistically significant associations between intra-genic structural changes or CER1 copy number and aspects of the environment they occupy (including aridity). The challenge is now to explain the species-specific evolutionary histories that contributed to the observed variation in CER1 and the extent to which it may contribute to the adaptability of eucalypts.

In plants where seed dispersal is limited compared with pollen dispersal, hybridisation may enhance gene exchange and species dispersal. We provide genetic evidence of hybridisation contributing to the expansion of the rare Eucalyptus risdonii into the range of the widespread Eucalyptus amygdalina. These closely related tree species are morphologically distinct, and observations suggest that natural hybrids occur along their distribution boundaries and as isolated trees or in small patches within the range of E. amygdalina. Hybrid phenotypes occur outside the range of normal dispersal for E. risdonii seed, yet in some hybrid patches small individuals resembling E. risdonii occur and are hypothesised to be a result of backcrossing. Using 3362 genome-wide SNPs assessed from 97 individuals of E. risdonii and E. amygdalina and 171 hybrid trees, we show that (i) isolated hybrids match the genotypes expected of F1 /F2 hybrids, (ii) there is a continuum in the genetic composition among the isolated hybrid patches from patches dominated by F1 /F2 -like genotypes to those dominated by E. risdonii-backcross genotypes, and (iii) the E. risdonii-like phenotypes in the isolated hybrid patches are most-closely related to proximal larger hybrids. These results suggest that the E. risdonii phenotype has been resurrected in isolated hybrid patches established from pollen dispersal, providing the first steps in its invasion of suitable habitat by long-distance pollen dispersal and complete introgressive displacement of E. amygdalina. Such expansion accords with the population demographics, common garden performance data, and climate modelling which favours E. risdonii and highlights a role of interspecific hybridisation in climate change adaptation and species expansion.

Climatic changes during the Pleistocene were responsible for dramatic redistributions of plant species worldwide. On the rugged southern hemisphere island of Tasmania, temperature increases following the last glaciation saw upslope migration of climatically suitable species from lowland refugia and the expansion of eucalypt-dominated forests and woodlands in the Central Highlands. We integrate multiple lines of evidence (chloroplast and nuclear DNA markers, seedling morphology, and survival in common garden experiments) from a group of closely related endemic eucalypts (the alpine white gums) to argue that (i) the Central Highlands of the island were colonised by multiple glacial refugia with hybridisation among species and previously separated populations, and (ii) natural selection has filtered the admixed populations, resulting in local adaptation to the harsh sub-alpine environment. Chloroplast haplotype diversity decreased and nuclear microsatellite diversity increased with altitude, chloroplast sharing among taxa was common, and nuclear DNA differentiation of morphologically distinct taxa was lower in the Central Highlands compared with lowland regions. Local adaptation in the highlands was signalled by evidence from (i) a glasshouse trial in which directional selection (Q(ST) > F-ST) had shaped seedling morphological trait variation and (ii) population survival differences in 35-year-old reciprocal plantings along the major environmental gradients. We conclude that the evolutionary response of these island endemic trees to past climate change has involved the interplay of both hybridisation and natural selection, highlighting the importance of maintaining species interactions under future climate change.

Eucalypts are of great ecological and economic importance. They dominate many woodland and forest landscapes in Australia and are planted in temperate to tropical areas worldwide for forestry. Population genomics research in eucalypts builds on a long history of genetics research, from quantitative trait analysis of field provenance trials to modern molecular genetics studies exploring neutral variation and population structure. It is backed by a high-quality reference genome and, whilst this rapidly expanding field is still in its infancy, it is already providing new insights into both ecological and industry-focused applications. This chapter reviews population genomics research in eucalypts, in the context of past genetics research. After describing the genomic resources available for eucalypts, we show that genomics is enabling greater resolution than ever before of species boundaries and patterns of hybridisation. Landscape genomics studies have revealed various patterns of population structure in natural populations, cryptic lineages within species and low divergence between closely related species. Genomic signatures of adaptation have been identified in several species, highlighting both climatic and edaphic factors as potential drivers of adaptation. In investigating the genetic architecture underlying complex traits, genomic tools are revealing not only associations with genes but also associations with differences in gene expression. With their worldwide importance, eucalypts are at the forefront of developments for integrating genomics into models for the genetic evaluation of forest tree breeding populations. Into the future, an increasing number of population genomics studies of eucalypts will help develop our understanding of the evolution of the genus; how species adapted to varied and changing environments; and the genetic architecture underlying adaptation and complex traits. Such knowledge will help guide species and provenance choices for conservation and ecological restoration under environmental change and will provide a valuable resource to breeding for forestry.

To understand the potential of forests to adapt to wildfire, we studied the genetic architecture of fire-related structural, damage and recovery traits in a globally important Australian forest tree species, Eucalyptus globulus. Fourteen traits were evaluated in an outcrossed F-2 population in a field trial in Tasmania, Australia, which was burnt by a wildfire 14 years after planting. The trial also included open-pollinated families of the grandparental dwarf and tall ecotypes used to produce the F-2 population. We studied the phenotypic correlations within the F-2 population and performed quantitative trait loci (QTL) analyses using a linkage map comprised of 472 markers. Ecotype comparisons revealed that almost all traits were under genetic control, with trees of the dwarf ecotype significantly more damaged and mainly recovering from lignotubers, whereas tall ecotype trees mainly recovered from epicormic resprouts extending for a variable height up the stem. Within the F-2, tree size was negatively correlated with fire damage and positively correlated with recovery. Genetic control of fire-related traits was confirmed by the detection of 38 QTL in the F-2 population. These QTL accounted for 4 to 43% of the phenotypic variation in these traits. Several QTL co-located and likely reflect pleiotropic effects. However, many independent QTL were detected, including QTL for crown consumption and trunk scorch, epicormic resprouting, resprout herbivory, and seedling establishment. The QTL detected argue that many genetically controlled mechanisms are responsible for variation in fire damage and recovery.

Selection on plant functional traits may occur through their direct effects on fitness (or a fitness component), or may be mediated by attributes of plant performance which have a direct impact on fitness. Understanding this link is particularly challenging for long-lived organisms, such as forest trees, where lifetime fitness assessments are rarely achievable, and performance features and fitness components are usually quantified from early-life history stages. Accordingly, we studied a cohort of trees from multiple populations of Eucalyptus pauciflora grown in a common-garden field trial established at the hot and dry end of the species distribution on the island of Tasmania, Australia. We related the within-population variation in leaf economic (leaf thickness, leaf area and leaf density) and hydraulic (stomatal density, stomatal length and vein density) traits, measured from two-year-old plants, to two-year growth performance (height and stem diameter) and to a fitness component (seven-year survival). When performance-trait relationships were modelled for all traits simultaneously, statistical support for direct effects on growth performance was only observed for leaf thickness and leaf density. Performance-based estimators of directional selection indicated that individuals with reduced leaf thickness and increased leaf density were favoured. Survival-performance relationships were consistent with size-dependent mortality, with fitness-based selection gradients estimated for performance measures providing evidence for directional selection favouring individuals with faster growth. There was no statistical support for an effect associated with the fitness-based quadratic selection gradient estimated for growth performance. Conditional on a performance measure, fitness-based directional selection gradients estimated for the leaf traits did not provide statistical support for direct effects of the focal traits on tree survival. This suggested that, under the environmental conditions of the trial site and time period covered in the current study, early-stage selection on the studied leaf traits may be mediated by their effects on growth performance, which in turn has a positive direct influence on later-age survival. We discuss the potential mechanistic basis of the direct effects of the focal leaf traits on tree growth, and the relevance of a putative causal pathway of trait effects on fitness through mediation by growth performance in the studied hot and dry environment.

BACKGROUND AND AIMS:Hybridization is increasingly recognized as an integral part of the dynamics of species range expansion and contraction. Thus, it is important to understand the reproductive barriers between co-occurring species. Extending previous studies that argued that the rare Eucalyptus risdonii was expanding into the range of the surrounding E. amygdalina by both seed and pollen dispersal, we here investigate the long-term fitness of both species and their hybrids and whether expansion is continuing.METHODS:We assessed the survival of phenotypes representing a continuum between the two pure species in a natural hybrid swarm after 29 years, along with seedling recruitment. The performance of pure species as well as of artificial and natural hybrids was also assessed over 28 years in a common garden trial.KEY RESULTS:In the hybrid zone, E. amygdalina adults showed greater mortality than E. risdonii, and the current seedling cohort is still dominated by E. risdonii phenotypes. Morphologically intermediate individuals appeared to be the least fit. Similar results were observed after growing artificial first-generation and natural hybrids alongside pure species families in a common garden trial. Here, the survival, reproduction, health and growth of the intermediate hybrids were significantly less than those of either pure species, consistent with hybrid inferiority, although this did not manifest until later reproductive ages. Among the variable progeny of natural intermediate hybrids, the most E. risdonii-like phenotypes were the most fit.CONCLUSIONS:This study contributes to the increasing number of reports of hybrid inferiority in Eucalyptus, suggesting that post-zygotic barriers contribute to the maintenance of species integrity even between closely related species. However, with fitness rapidly recovered following backcrossing, it is argued that hybridization can still be an important evolutionary process, in the present case appearing to contribute to the range expansion of the rare E. risdonii in response to climate change.

The evolutionary trajectory of a population both influences and is influenced by characteristics of its genome. A disjunct population, for example is likely to exhibit genomic features distinct from those of continuous populations, reflecting its specific evolutionary history and influencing future recombination outcomes. We examined genetic diversity, population differentiation and linkage disequilibrium (LD) across the highly disjunct native range of the Australian forest tree Eucalyptus globulus , using 203,337 SNPs genotyped in 136 trees spanning seven races. We found support for four broad genetic groups, with moderate F ST , high allelic diversity and genome-wide LD decaying to an r 2 of 0.2 within 4 kb on average. These results are broadly similar to those reported previously in Eucalyptus species and support the ‘ring’ model of migration proposed for E. globulus . However, two of the races (Otways and South-eastern Tasmania) exhibited a much slower decay of LD with physical distance than the others and were also the most differentiated and least diverse, which may reflect the effects of selective sweeps and/or genetic bottlenecks experienced in their evolutionary history. We also show that F ST and rates of LD vary within and between chromosomes across all races, suggestive of recombination outcomes influenced by genomic features, hybridization or selection. The results obtained from studying this species serve to illustrate the genomic effects of population disjunction and further contribute to the characterisation of genomes of woody genera.

The integration of genomic data into genetic evaluations can facilitate the rapid selection of superior genotypes and accelerate the breeding cycle in trees. In this study, 390 trees from 74 control-pollinated families were genotyped using a 36K Axiom SNP array. A total of 15,624 high-quality SNPs were used to develop genomic prediction models for mammalian bark stripping, tree height, and selected primary and secondary chemical compounds in the bark. Genetic parameters from different genomic prediction methods-single-trait best linear unbiased prediction based on a marker-based relationship matrix (genomic best linear unbiased prediction), multitrait single-step genomic best linear unbiased prediction, which integrated the marker-based and pedigree-based relationship matrices (single-step genomic best linear unbiased prediction) and the single-trait generalized ridge regression-were compared to equivalent single- or multitrait pedigree-based approaches (ABLUP). The influence of the statistical distribution of data on the genetic parameters was assessed. Results indicated that the heritability estimates were increased nearly 2-fold with genomic models compared to the equivalent pedigree-based models. Predictive accuracy of the single-step genomic best linear unbiased prediction was higher than the ABLUP for most traits. Allowing for heterogeneity in marker effects through the use of generalized ridge regression did not markedly improve predictive ability over genomic best linear unbiased prediction, arguing that most of the chemical traits are modulated by many genes with small effects. Overall, the traits with low pedigree-based heritability benefited more from genomic models compared to the traits with high pedigree-based heritability. There was no evidence that data skewness or the presence of outliers affected the genomic or pedigree-based genetic estimates.

Leptospermum scoparium is emerging as an economically important plant for the commercial production of mānuka honey and essential oils, both exhibiting unique antibacterial attributes. To support its domestication this is the first quantitative genetic study of variation for L. scoparium traits. It utilised plants from 200 open-pollinated families derived from 40 native populations, from across the species range in Tasmania, grown in a common garden field trial. The traits studied were survival, growth, and the flowering traits precocity, the timing of seasonal peak flowering, flowering duration, and flowering intensity. Significant genetic variation was evident at the population level for all traits studied and at the family level for three traits-growth, flowering precocity, and time to peak flowering. These three traits had moderate to high narrow-sense heritability estimates ranging from 0.27 to 0.69. For six of the traits studied, population differences were associated with climate attributes at the locations where seed was collected, suggesting adaptation to the local climate may have contributed to the observed population differentiation. Population level geographical trends suggest that genotypes to focus on for domestication originate from the eastern half of Tasmania for precociousness and the western half of Tasmania for earlier time to peak flowering and extended flowering duration.

There is increasing interest in the phenotypic plasticity of tree species to variation in climate as it may affect the economic value of current and future plantations. This study applied ensemble learning methods to explore the plastic response of Eucalyptus nitens pulpwood selection traits of growth (DBH, diameter at 1.3 m height), wood density and Kraft pulp yield to variation in elevation, geography and associated climate variables. To help explain the pulp yield response, we also modelled underlying biological traits—cellulose, lignin and extractives. The study was based on data from 84 harvest-age plots of common genetic origin across the pulpwood plantation estate in north-western Tasmania, Australia. DBH and wood density were obtained from resistance profile traces, collected on standing trees using a drilling resistance tool. In addition, outerwood cores were taken at 1.3 m stem height for (i) calibrating the resistance measures for wood density, and (ii) for assessment of pulp yield and wood chemistry based on near infrared spectroscopy. Modelling of the variation in plot means using the random forest algorithms showed growth and wood properties were influenced by the growing period climate of the plot. Of the climate variables studied, growth was mainly influenced by temperature, while wood density was mainly affected by rainfall-related variables. Wood density varied independently of growth and decreased with increasing annual rainfall and elevation of the plots. Pulp yield had the poorest fit statistics, it was influenced by a mix of climatic and geographic variables, and appeared independent of variation in growth and wood density. The plot variation in pulp yield was best explained by the modelled trends in the underlying biological traits of cellulose and lignin. Using these models, the plastic response of the key pulpwood traits to climate was mapped across the E. nitens plantation estate to help predict current plantation attributes and guide future choices of sites for plantation establishment.

With climate change impacting trees worldwide, enhancing adaptation capacity has become an important goal of provenance translocation strategies for forestry, ecological renovation, and biodiversity conservation. Given that not every species can be studied in detail, it is important to understand the extent to which climate adaptation patterns can be generalised across species, in terms of the selective agents and traits involved. We here compare patterns of genetic-based population (co)variation in leaf economic and hydraulic traits, climate–trait associations, and genomic differentiation of two widespread tree species (Eucalyptus pauciflora and E. ovata). We studied 2-year-old trees growing in a common-garden trial established with progeny from populations of both species, pair-sampled from 22 localities across their overlapping native distribution in Tasmania, Australia. Despite originating from the same climatic gradients, the species differed in their levels of population variance and trait covariance, patterns of population variation within each species were uncorrelated, and the species had different climate–trait associations. Further, the pattern of genomic differentiation among populations was uncorrelated between species, and population differentiation in leaf traits was mostly uncorrelated with genomic differentiation. We discuss hypotheses to explain this decoupling of patterns and propose that the choice of seed provenances for climate-based plantings needs to account for multiple dimensions of climate change unless species-specific information is available.

Understanding the capacity of forest tree species to adapt to climate change is of increasing importance for managing forest genetic resources. Through a genomics approach, we modelled spatial variation in climate adaptation within the Australian temperate forest tree Eucalyptus globulus, identified putative climate drivers of this genomic variation, and predicted locations of future climate refugia and populations at-risk of future maladaptation. Using 812,158 SNPs across 130 individuals from 30 populations (i.e., localities) spanning the species’ natural range, a gradientForest algorithm found 1177 SNPs associated with locality variation in home-site climate (climate-SNPs), putatively linking them to climate adaptation. Very few climate-SNPs were associated with population-level variation in drought susceptibility, signalling the multi-faceted nature and complexity of climate adaptation. Redundancy analysis (RDA) showed 24% of the climate-SNP variation could be explained by annual precipitation, isothermality, and maximum temperature of the warmest month. Spatial predictions of the RDA climate vectors associated with climate-SNPs allowed mapping of genomically informed climate selective surfaces across the species’ range under contemporary and projected future climates. These surfaces suggest over 50% of the current distribution of E. globulus will be outside the modelled adaptive range by 2070 and at risk of climate maladaptation. Such surfaces present a new integrated approach for natural resource managers to capture adaptive genetic variation and plan translocations in the face of climate change.

Background and aims The petaline operculum that covers the inner whorls until anthesis and the woody capsule that develops after fertilization are reproductive structures of eucalypts that protect the flower and seeds. Although they are distinct organs, they both develop from flower buds and this common ontogeny suggests shared genetic control. In Eucalyptus globulus their morphology is variable and we aimed to identify the quantitative trait loci (QTL) underlying this variation and determine whether there is common genetic control of these ecologically and taxonomically important reproductive structures. Methods Samples of opercula and capsules were collected from 206 trees that belong to a large outcrossed F-2 E. globulus mapping population. The morphological variation in these structures was characterized by measuring six operculum and five capsule traits. QTL analysis was performed using these data and a linkage map consisting of 480 markers. Key results A total of 27 QTL were detected for operculum traits and 28 for capsule traits, with the logarithm of odds ranging from 2.8 to 11.8. There were many co-located QTL associated with operculum or capsule traits, generally reflecting allometric relationships. A key finding was five genomic regions where co-located QTL affected both operculum and capsule morphology, and the overall trend for these QTL was to affect elongation of both organs. Some of these QTL appear to have a significant effect on the phenotype, with the strongest QTL explaining 26.4 % of the variation in operculum shape and 16.4 % in capsule shape. Flower bud measurements suggest the expression of these QTL starts during bud development. Several candidate genes were found associated with the QTL and their putative function is discussed. Conclusions Variation in both operculum and capsule traits in E. globulus is under strong genetic control. Our results suggest that these reproductive structures share a common genetic pathway during flower bud development.

A major challenge in ecological restoration is assessing the success of restoration plantings in producing habitats that provide the desired ecosystem functions and services. Forest structural complexity and biomass accumulation are key measures used to monitor restoration success and are important factors determining animal habitat availability and carbon sequestration. Monitoring their development through time using traditional field measurements can be costly and impractical, particularly at the landscape-scale, which is a common requirement in ecological restoration. We explored the application of proximal sensing technology as an alternative to traditional field surveys to capture the development of key forest structural traits in a restoration planting in the Midlands of Tasmania, Australia. We report the use of a hand-held laser scanner (ZEB1) to measure annual changes in structural traits at the tree-level, in a mixed species common-garden experiment from seven- to nine-years after planting. Using very dense point clouds, we derived estimates of multiple structural traits, including above ground biomass, tree height, stem diameter, crown dimensions, and crown properties. We detected annual increases in most LiDAR-derived traits, with individual crowns becoming increasingly interconnected. Time by species interaction were detected, and were associated with differences in productivity between species. We show the potential for remote sensing technology to monitor temporal changes in forest structural traits, as well as to provide base-line measures from which to assess the restoration trajectory towards a desired state.