Role of insect vectors in epidemiology and invasion risk of <Emphasis Type="Italic">Fusarium circinatum</Emphasis>, and risk assessment of biological control of invasive <Emphasis Type="Italic">Pinus contorta</Emphasis>

Pitch canker, caused by the pathogen Fusarium circinatum, is a serious disease of pines, Pinus species. It is a threat to natural and planted pine forests, and to date it has invaded countries across five continents. Pine-feeding insects can play a key role in the epidemiology of the disease, as wounding agents allowing pathogen access or as vectors transmitting the pathogen from infected to healthy trees. We reviewed the role of insects in the epidemiology of pitch canker worldwide and assessed which insects are present in New Zealand that may act as wounding agents or vectors to determine whether pathogen invasion could adversely affect Pinus radiata plantation forests and urban trees. We also evaluated whether cone or seed insects of pines could be introduced as biological control agents of invasive Pinus contorta and how this may affect the impact of a potential F. circinatum invasion. As there are no native pines or other Pinaceae in New Zealand, there are only a few pine insects, mainly accidental introductions. None of the insects recorded on pines in New Zealand is likely to be a vector, suggesting low disease risk. Of six potentially suitable biocontrol candidates, the European pine cone weevil Pissodes validirostris is the most promising regarding host specificity and impact on seed production, but there is uncertainty about its ability to act as a vector of F. circinatum. Our methodology to review and evaluate the vector potential of pine associates can be used as a generic framework to assess the potential impacts of F. circinatum invasion.     

Beyond the genomes of Fulvia fulva (syn. Cladosporium fulvum) and Dothistroma septosporum: New insights into how these fungal pathogens interact with their host plants

Fulvia fulva and Dothistroma septosporum are closely related apoplastic pathogens with similar lifestyles but different hosts: F. fulva is a pathogen of tomato, whilst D. septosporum is a pathogen of pine trees. In 2012, the first genome sequences of these pathogens were published, with F. fulva and D. septosporum having highly fragmented and near-complete assemblies, respectively. Since then, significant advances have been made in unravelling their genome architectures. For instance, the genome of F. fulva has now been assembled into 14 chromosomes, 13 of which have synteny with the 14 chromosomes of D. septosporum, suggesting these pathogens are even more closely related than originally thought. Considerable advances have also been made in the identification and functional characterization of virulence factors (e.g., effector proteins and secondary metabolites) from these pathogens, thereby providing new insights into how they promote host colonization or activate plant defence responses. For example, it has now been established that effector proteins from both F. fulva and D. septosporum interact with cell-surface immune receptors and co-receptors to activate the plant immune system. Progress has also been made in understanding how F. fulva and D. septosporum have evolved with their host plants, whilst intensive research into pandemics of Dothistroma needle blight in the Northern Hemisphere has shed light on the origins, migration, and genetic diversity of the global D. septosporum population. In this review, we specifically summarize advances made in our understanding of the F. fulva–tomato and D. septosporum–pine pathosystems over the last 10 years.     

Pinus radiata invasion in Australia: Identifying key knowledge gaps and research directions

Abstract Early recognition of plant invaders has been widely identified as the key to their successful management and yet too often species are only noticed and receive adequate attention once they have become widespread and control has become difficult and costly. Slow growing species are at particular risk of being overlooked, despite their ability to cause significant ecological damage. One such species, Pinus radiata (Monterey pine), has spread from large commercial plantations into native vegetation across the southern hemisphere. Here we review the status of P. radiata invasion in Australia, a country where the species has successfully naturalized but remains an invader of only low‐level concern. Patterns of spread in Australia mirror those in New Zealand and South Africa, two countries where invasive pines are considered major threats to biodiversity. While many areas adjacent to plantations remain free from invasion, dense infestations have occurred at several sites in areas of high conservation value demanding the implementation of adequate control measures. Expansion of Australia's plantation estate and increasing human disturbance of natural areas surrounding plantations will increase the likelihood and extent of invasion. Continued monitoring of wildling populations will determine the ability of pines to dominate native eucalypt forest and will provide insight into broader ideas of community invasibility.     

A look into the genetic diversity of Lecanosticta acicola in northern Europe

For northern Europe Lecanosticta acicola is an emerging pine needle pathogen. This study gives a first look into the population genetics of the pathogen in Estonia, the first population documented in that region. The main aim of this study was to investigate the genetic diversity and population structure of the pathogen in this new region for the fungus. For this purpose, 104 isolates from 2010 to 2017 were analysed with 11 microsatellite and mating type markers. The stand where the pathogen's jump from an exotic host to the native Scots pine was recorded was also involved in this analysis. The analysis revealed low genetic diversity and a high number of clones that indicated L. acicola is an invasive species in northern Europe. Results suggest that several separate introductions have taken place and anthropogenic activity has apparently affected the spread of the pathogen. Clonal reproduction is dominating and although sexual reproduction is possible, it probably takes place infrequently.     

A molecular investigation of the genus Ecklonia (Phaeophyceae,Laminariales) with special focus on the Southern Hemisphere

Brown algae of the order Laminariales, commonly referred to as kelps, are the largest and most productive primary producers in the coastal inshore environment. The genus Ecklonia (Lessoniaceae, Phaeophyceae) consists of seven species with four species in the Northern Hemisphere and three in the Southern Hemisphere. It was recently transferred to the family Lessoniaceae based on phylogenetic analyses of nuclear and chloroplastic markers, though the type of the genus was not included and its relationship with allied genera Eckloniopsis and Eisenia remained unresolved. The present study is the first to produce a phylogeny focussed on the genus Ecklonia. It included sequences from nuclear, mitochondrial, and chloroplastic DNA, for most of the distribution range of the three current Southern Hemisphere species (Ecklonia radiata, Ecklonia maxima, and a sample of a putative Ecklonia brevipes specimen), sequences for East Asiatic species (Ecklonia cava, Ecklonia kurome, and Ecklonia stolonifera), as well as the closely related genera Eckloniopsis and Eisenia. Results confirmed E. radiata and E. maxima as two distinct species in South Africa, E. radiata as a single species throughout the Southern Hemisphere (in South Africa, Australia, and New Zealand) and East Asiatic species as a distinct lineage from the Southern Hemisphere clade. Results further pointed out a close sister relationship between Eckloniopsis radicosa and two Eisenia species (including the type species: Eisenia arborea) to the genus Ecklonia suggesting that the genera Eckloniopsis and Eisenia are superfluous.     

Genetic Conservation of Insular Populations of Monterey Pine (Pinus Radiata D. Don)

Monterey pine (Pinus radiata D. Don) has only five extant native populations: three disjunct populations along the coast of California, USA and two on Mexican islands. All populations have been influenced by human activity, but the island populations in particular have been affected by introduced biota. On Guadalupe Island, the pine population has suffered drastically from overgrazing by introduced goats. We visited both island populations and described their status, took measurements, and made seed collections. We counted approximately 200 mature pine trees and virtually no seedlings on Guadalupe Island: a reduction of approximately half the population in the last 50 years. The trees are all large (mean diameter of 144 cm) –considerably larger than trees from the other four populations – and arguably near the end of their natural lifespan. The population on Cedros Island is much more robust, with thousands of trees. None sampled were as large as those on Guadalupe Island (mean diameter of 20 cm) and many groves were young and even-aged – presumably the consequence of natural regeneration after a recent fire. Tissue samples from trees on both islands did not show evidence of infection from the pitch canker pathogen, Fusarium circinatum, that has caused significant mortality in the three mainland populations. Caution is recommended in any restoration activity for the Guadalupe Island pines. Inbreeding levels could indicate the need for some planting or seeding intervention but there are also risks associated with this. Natural regeneration – after goat removal – is preferred.     

Preliminary studies on a native Australian eucalypt forest invaded by exotic pines

Summary In some parts of Australia, the exotic tree species Pinus radiata is invading native eucalypt forests from adjacent plantations. At one site, measurements have been made in order to determine the dispersal gradient and rate of population increase of the self-sown pines. From these, it is evident that, barring further human interference, a new type of mixed forest community will develop.     

Evaluation of the field impacts of simulated Bacillus thuringiensis‐transgenic Pinus radiata on nontarget native Lepidoptera and their natural enemies in a New Zealand plantation forest

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Protura in native and exotic forests in the North Island of New Zealand

Abstract

This study provides data on population and community ecology of Protura in native forests and Pinus radiata plantations in New Zealand. Abundance, age structure, sex ratios, biodiversity, and relationship with soil chemistry are discussed. Protura were significantly more abundant in pine plantations in comparison to native forests. Among native forests, Protura were most abundant under Southern beech (Nothofagus solandri). The abundance patterns may reflect the association between Protura and fungal communities in the soil. No correlation was found between soil organic matter, nitrogen, phosphorous, pH, cation exchange capacity, and the density of Protura. Protura assemblage composition was significantly related to forest type. New Zealand endemic species were associated with native forests; species with distribution outside New Zealand dominated in pine plantations. The distribution records within New Zealand were expanded for five species. The Protura fauna of New Zealand was increased to 18 species. Berberentulus capensis, Eosentomon australicum and Australentulus tillyardi are new records for New Zealand fauna.     

Pine plantations in south eastern Australia support highly impoverished ant assemblages (Hymenoptera: Formicidae)

Pitfall trap surveys of ants were undertaken in mature and intermediate-aged Pinus radiata plantations and parental eucalypt forest in central Victoria, Australia. Ant assemblages of pine plantations are an impoverished subset of the eucalypt forest assemblages. Very few ant morphospecies, most of them in very low abundance, were found in unthinned intermediate-aged pines, and the numbers were substantially higher in older, thinned pines in which understorey vegetation is considerably more developed. Implications for plantation management and conservation of epigaeic ant assemblages are discussed.     

Chloroplast DNA Transgresses Species Boundaries and Evolves at Variable Rates in the California Closed-Cone Pines (Pinus radiata, P. muricata, and P. attenuata)

We studied phylogenetic relationships among populations and species in the California closed-cone pines (Pinus radiata D. Don, P. attenuata Lemm., and P. muricata D. Don) via chloroplast DNA restriction site analysis. Data on genetic polymorphism within and among 19 populations in the three species were collected using9 to 20 restriction enzymes and 38 to 384 trees. Because only five clades and extremely low intraclade diversity were found, additional phylogenetic data were collected using a single representative per clade and two outgroup species, P. oocarpa Schiede and P. jeffreyi Loud. In total, 25 restriction enzymes were employed and approximately 2.7 kb surveyed (2.3% of genome). The five clades recognized were Monterey pine, knob-cone pine, and the southern, intermediate, and northern races of bishop pine. On the basis of bootstrapping, both Wagner and Dollo parsimony analyses strongly separated the northern and intermediate races of bishop pine from the southern race; knobcone pine from Monterey and bishop pines; and the closed-cone pines from the two outgroups. Approximate divergence times were estimated for the lineages leading to knob-cone pine and to the intermediate and northern populations of bishop pine. The position of Monterey pine relative to bishop pine within their monophyletic clade was unresolved. Surprisingly, Montery pine and the southern race of bishop pine were much more similar to one another than was the southern race of bishop pine to its conspecific intermediate and northern races. Both the Monterey and southern bishop pine lineages also evolved severalfold more slowly than did the knobcone pine and intermediate-northern bishop pine lineages. These results differ significantly from a recent allozyme study, corroborating previous observations that chloroplast genome phylogeny can depart substantially from that of nuclear genes.     

A FURTHER CONTRIBUTION TO CLOSED-CONE PINE (OOCARPAE) HISTORY

A new record of a fossil Monterey pine (Pinus radiata D. Don) is recorded from rocks of the Plio-Pleistocene age, southern California, and the affinities of other presumed fossil closed-cone pines are revised. This has led to the recognition of two new pine species, Pinus storeyana Axelrod of subsect. Oocarpae and Pinus verdiana Axelrod of subsect. Ponderosae. Geologic evidence indicates that forerunners of the present California and Mexican species of Oocarpae were in proximity in Mexico during the Miocene. They were isolated following opening of the Gulf of California, the northward movement of Baja and Alta California on the San Andreas fault system, and the spread of desert climate. New populations of Pinus radiata developed during the Pliocene (5–2 million years) and later as environmental diversity increased. This also enabled Pinus remorala Mason and P. muricata D. Don, which the fossil record suggests were distinct into the Late Pleistocene, to hybridize in the new topographic-climatic Holocene environments.     

Terpenes as Useful Markers in Differentiation of Natural Populations of Relict Pines Pinus heldreichii,P. nigra,and P. peuce

Comparative analysis of terpene diversity and differentiation of relict pines Pinus heldreichii, P. nigra, and P. peuce from the central Balkans was performed at the population level. Multivariate statistical analyses showed that the composition of needle terpenes reflects clear divergence among the pine species from different subgenera: P. peuce (subgenus Strobus) vs. P. nigra and P. heldreichii (subgenus Pinus). In addition, despite the described morphological similarities and the fact that P. nigra and P. heldreichii may spontaneously hybridize, our results indicated differentiation of their populations naturally growing in the same area. In accordance with recently proposed concept of ‘flavonic evolution’ in the genus Pinus, we assumed that the terpene profile of soft pine P. peuce, defined by high amounts of six monoterpenes, is more basal than those of hard pines P. nigra and P. heldreichii, which were characterized by high content levels of mainly sesquiterpenes. In order to establish precise positions of P. heldreichii, P. nigra and P. peuce within the taxonomic and phylogenetic tree, as well as develop suitable conservation strategies and future breeding efforts, it is necessary to perform additional morphological, biochemical, and genetic studies.     

Beneath the bark: associations among Sirex noctilio development,bluestain fungi,and pine host species in North America

1. Invasive species with global distributions encounter unique environmental and biotic variables that can greatly affect the magnitude of their impact. The European woodwasp, Sirex noctilio Fabricius, is a prime example that has invaded climatically and ecologically distinct ecosystems across the Southern Hemisphere and, more recently, North America. 2. Northeastern North America presents a unique set of conditions, including pine host species, native parasitoids, a diverse assemblage of native co‐colonising insects, and fungal associates of these co‐colonisers. In North America, S. noctilio attacks both a naturalised ancestral host (Pinus sylvestris L.) from Europe and a naïve native host (P. resinosa). A large assemblage of insects and their associated bluestaining ophiostomatoid fungi colonise these pines. Competition between S. noctilio and this group is a hypothesised mechanism of biotic resistance in the invaded region of North America, possibly via superior resource capture abilities or alteration of host tissue by bluestain fungi. 3. Investigating these ecological interactions is challenging because they manifest deep in the xylem tissue. To overcome this, 30 experimentally stressed trees were systematically dissected with an electric log splitter to investigate the effects of bluestain fungi and tree factors on S. noctilio development and parasitism by native hymenopterans. 4. Body size and colonisation density were affected by pine species, with S. noctilio being 25% larger and densities three‐fold greater in P. sylvestris than P. resinosa. Survivorship was slightly negatively correlated with the proportional volume of bluestain infection. Interestingly, rhyssine (Rhyssa and Megarhyssa spp.) parasitism responded positively to greater S. noctilio density, but there was no density relationship with Ibalia leucospoides ensiger parasitism. Pine host species appears to play a strong role in S. noctilio development, which is important considering uninvaded regions of North America have a diversity of pine species that likely vary in their susceptibility to this woodborer.     

Transgenic Pinus radiata from Agrobacterium tumefaciens–mediated transformation of cotyledons

A method for Agrobacterium tumefaciens-mediated transformation of Pinus radiata cotyledon explants was developed using commercially available open-pollinated seed. Pinus radiata is the most widely planted commercial conifer species in the Southern Hemisphere. Reports on transformation of this species have relied on particle bombardment of embryogenic callus derived from immature embryos. The main drawback to the method is the small number of genotypes that are amenable to transformation and regeneration. Since more than 80% of genotypes of radiata pine can be regenerated using cotyledons from mature seed, cotyledon explants were cocultivated with A. tumefaciens strain AGL1 containing a plasmid coding for the neomycin phosphotransferase II (nptII) gene and the -glucuronidase (GUS) gene (uidA). Transformed shoots were selected using either geneticin or kanamycin. Critical factors for successful transformation were survival of the cotyledons after cocultivation and selection parameters. Of the 105 putative transformants that were recovered from selection media, 70% were positive for integration of the nptII gene when analysed by PCR. GUS histochemical assay for uidA expression was unreliable because of reaction inhibition by unidentified compounds in the pine needles. Further, only 4 of the 26 independent transformants characterised by PCR and Southern analysis contained an intact copy of both genes. The remaining 22 transformants appeared to have a truncated or rearranged copy of the T-DNA. It is possible that the truncation/rearrangements are due to the Cauliflower mosaic virus (CaMV) 35S promoter. Analysis of the T-DNA junction sites and sequencing of the introduced DNA will help elucidate the nature of T-DNA insertion so that genetic modification of radiata pine can be targeted effectively.Communicated by P. Debergh     

Exotic Pine Plantations in Victoria, Australia: A Threat to Epigaeic Beetle (Coleoptera) Assemblages?

Ground-dwelling invertebrates were sampled by pitfall traps over a 14 week period in parent eucalypt forest and three stages of exotic Pinus radiata plantations established after forest clearance in central Victoria. The four treatments each yielded numerous beetle morphospecies, and the assemblages corresponded only partially with the understorey vegetation. More than 200 beetle morphospecies were captured during this short survey, with 30 found in all four treatments; few morphospecies were abundant (only five with >100 individuals in a total of 3382 beetles). Each treatment had unique morphospecies, but all were rich, with the lowest diversity being 91 morphospecies (young pines). These data confirm that beetle diversity can remain substantial in exotic softwood plantations, but considerable care is needed to interpret this apparent diversity in relation to forest management and the effect of replacement of native forests by exotic taxa.     

High resolution analysis of mating systems: inbreeding in natural populations of Pinus radiata

Pinus radiata has a history of population bottlenecks and is currently restricted to five relatively small populations, three in mainland California, and two on islands off the coast of Baja California. Using highly polymorphic microsatellite markers and a newly developed statistical approach, we were able to estimate individual inbreeding coefficients and can thus analyse the mating system with high resolution. We find a bimodal distribution of inbreeding coefficients: most individuals result from selfing whereas few (in the mainland populations) to a modest number (in the island populations) are likely selfed. In most other pine species and presumably in the ancestral P. radiata population, occurrence of mature selfed individuals would be impossible because of the high genetic load. We therefore conclude that inbreeding depression has been purged in P. radiata and that the mating system has changed as a consequence.     

Deciphering the worldwide invasion of the Asian long‐horned beetle: A recurrent invasion process from the native area together with a bridgehead effect

Retracing introduction routes is crucial for understanding the evolutionary processes involved in an invasion, as well as for highlighting the invasion history of a species at the global scale. The Asian long‐horned beetle (ALB) Anoplophora glabripennis is a xylophagous pest native to Asia and invasive in North America and Europe. It is responsible for severe losses of urban trees, in both its native and invaded ranges. Based on historical and genetic data, several hypotheses have been formulated concerning its invasion history, including the possibility of multiple introductions from the native zone and secondary dispersal within the invaded areas, but none have been formally tested. In this study, we characterized the genetic structure of ALB in both its native and invaded ranges using microsatellites. In order to test different invasion scenarios, we used an approximate Bayesian “random forest” algorithm together with traditional population genetics approaches. The strong population differentiation observed in the native area was not geographically structured, suggesting complex migration events that were probably human‐mediated. Both native and invasive populations had low genetic diversity, but this characteristic did not prevent the success of the ALB invasions. Our results highlight the complexity of invasion pathways for insect pests. Specifically, our findings indicate that invasive species might be repeatedly introduced from their native range, and they emphasize the importance of multiple, human‐mediated introductions in successful invasions. Finally, our results demonstrate that invasive species can spread across continents following a bridgehead path, in which an invasive population may have acted as a source for another invasion.