Most members of the oomycete genus Phytophthora are primary plant pathogens. Both soil- and airborne Phytophthora species are able to survive adverse environmental conditions with enduring resting structures, mainly sexual oospores, vegetative chlamydospores and hyphal aggregations. Soilborne Phytophthora species infect fine roots and the bark of suberized roots and the collar region with motile biflagellate zoospores released from sporangia during wet soil conditions. Airborne Phytophthora species infect leaves, shoots, fruits and bark of branches and stems with caducous sporangia produced during humid conditions on infected plant tissues and dispersed by rain and wind splash. During the past six decades, the number of previously unknown Phytophthora declines and diebacks of natural and semi-natural forests and woodlands has increased exponentially, and the vast majority of them are driven by introduced invasive Phytophthora species. Nurseries in Europe, North America and Australia show high infestation rates with a wide range of mostly exotic Phytophthora species. Planting of infested nursery stock has proven to be the main pathway of Phytophthora species between and within continents. This review provides insights into the history, distribution, aetiology, symptomatology, dynamics and impact of the most important canker, decline and dieback diseases caused by soil- and airborne Phytophthora species in forests and natural ecosystems of Europe, Australia and the Americas.
During the course of the past three years, a new disease of Pinus radiata , referred to as 'Daño Foliar del Pino' (DFP) has appeared in the Arauco province of Chile and subsequently spread to other areas. The disease is typified by needle infections, exudation of resin at the bases of the needle brachyblasts and, in younger trees, necrotic lesions in the cambium, which eventually girdle the branches. The disease causes the death of young seedlings and mature trees can also succumb after a few years of successive infection, probably hastened by opportunistic fungi such as Diplodia pinea . Isolations on selective medium for Phytophthora spp. led to the consistent isolation of a Phytophthora sp. from needle tissue. DNA sequence comparisons for the ITS rDNA and cox II gene regions, and morphological observation showed that this oomycete represents a previously undescribed species for which the name Phytophthora pinifolia sp. nov. is provided. This new species is characterized by unbranched sporangiophores, and non-papillate, sub-globose to ovoid sporangia that are occasionally free from the sporangiophore with medium length pedicels. Despite using a number of oospore inducing techniques, oogonia/antheridia were not observed in isolates of P. pinifolia . Pathogenicity trials with P. pinifolia showed that it is pathogenic to P. radiata and causes rapid death of the succulent apical parts of young plants. Phytophthora pinifolia is the first Phytophthora known to be associated with needles and shoots of a Pinus sp. and its aerial habit is well matched with the occurrence and symptoms of DFP in Chile.
The Valdivian rainforest, one of the global hotspots of biodiversity, is a temperate rainforest originating as a Tertiary relic from the supercontinent Gondwana. In November 2014, a survey of Phytophthora diversity was performed in 13 natural forest stands and 20 forest streams and rivers in two protected areas near Valdivia and in a temperate montane forest in the Concepción area. One planted stand each of the introduced tree species Castanea sativa and Fagus sylvatica were also included. Using baiting assays, eight described species and four previously unknown taxa of Phytophthora were isolated from 86% of the 50 rhizosphere soil samples from seven of the eight tree species sampled in 12 forest stands, and from 20 streams: P. chlamydospora, P. cinnamomi, P. kernoviae, P. lacustris, P. plurivora, P. pseudosyringae, P. ×cambivora, P. ×stagnum, P. valdiviana nom. prov. from Clade 2b, P. madida nom. prov. from Clade 8a, and P. chilensis nom. prov. and P. pseudokernoviae nom. prov. The latter two species are the closest relatives of P. kernoviae from Clade 10. Phytophthora pseudokernoviae nom. prov. was also isolated from necrotic leaves of Drimys winteri. From the Valdivia river, a swarm of three Clade 6 hybrids was recovered. Each hybrid isolate resulted from multiple reticulation events with P. thermophila as maternal and both P. amnicola and P. chlamydospora as paternal parents. In addition, three previously unknown and recently described Nothophytophthora species, N. caduca, N. chlamydospora and N. valdiviana, were isolated from several forest streams. Phytophthora cinnamomi, the most common and widespread species in soils of native forests, was associated with severe dieback of Valdivian rainforest trees, in particular D. winteri, Luma apiculata, Nothofagus dombeyi and the endangered Saxegothaea conspicua. A first pathogenicity test demonstrated high aggressiveness of P. cinnamomi to several native tree species, including N. dombeyi, Blepharocalyx cruckshanksii and Gevuina avellana.
During various surveys of Phytophthora diversity in Europe, Chile and Vietnam slow growing oomycete isolates were obtained from rhizosphere soil samples and small streams in natural and planted forest stands. Phylogenetic analyses of sequences from the nuclear ITS, LSU, β-tubulin and HSP90 loci and the mitochondrial cox1 and NADH1 genes revealed they belong to six new species of a new genus, officially described here as Nothophytophthora gen. nov., which clustered as sister group to Phytophthora. Nothophytophthora species share numerous morphological characters with Phytophthora: persistent (all Nothophytophthora spp.) and caducous (N. caduca, N. chlamydospora, N. valdiviana, N. vietnamensis) sporangia with variable shapes, internal differentiation of zoospores and internal, nested and extended (N. caduca, N. chlamydospora) and external (all Nothophytophthora spp.) sporangial proliferation; smooth-walled oogonia with amphigynous (N. amphigynosa) and paragynous (N. amphigynosa, N. intricata, N. vietnamensis) attachment of the antheridia; chlamydospores (N. chlamydospora) and hyphal swellings. Main differing features of the new genus are the presence of a conspicuous, opaque plug inside the sporangiophore close to the base of most mature sporangia in all known Nothophytophthora species and intraspecific co-occurrence of caducity and non-papillate sporangia with internal nested and extended proliferation in several Nothophytophthora species. Comparisons of morphological structures of both genera allow hypotheses about the morphology and ecology of their common ancestor which are discussed. Production of caducous sporangia by N. caduca, N. chlamydospora and N. valdiviana from Valdivian rainforests and N. vietnamensis from a mountain forest in Vietnam suggests a partially aerial lifestyle as adaptation to these humid habitats. Presence of tree dieback in all forests from which Nothophytophthora spp. were recovered and partial sporangial caducity of several Nothophytophthora species indicate a pathogenic rather than a saprophytic lifestyle. Isolation tests from symptomatic plant tissues in these forests and pathogenicity tests are urgently required to clarify the lifestyle of the six Nothophytophthora species.
Phytophthora pinifolia is the causal agent of the recently discovered needle disease of Pinus radiata in Chile, referred to as "Daño Foliar del Pino" (DFP). The genetic structure of the pathogen population is unknown, which hinders our understanding of its appearance and spread in Chile since 2004. In this study, a population of 88 cultures of P. pinifolia isolated from P. radiata at several localities in Chile was evaluated for genotypic diversity using amplified fragment length polymorphisms (AFLPs). Results of the AFLP analyses showed that the P. pinifolia population in Chile consists of two near identical genotypes but with no genetic differentiation based on geography, year of isolation or the part of the tree from which the isolates were obtained. Mating experiments did not lead to the production of gametangia suggesting that the organism is sterile. The fact that a single clonal genotype dominates the population of P. pinifolia in Chile supports the hypothesis that P. pinifolia was recently introduced into this country and that its impact is due to a new and susceptible host encounter.
In 2014, a new and serious leaf and shoot disease of unknown aetiology appeared in Eucalyptus plantations of North Sumatra, Indonesia. The disease is characterized by black necrotic spots that initially appear on young leaves and petioles, which become scab‐like as the lesions age. Infected trees respond to infection by producing shoots with small leaves that commonly appear feathered. Fruiting structures typical of most foliar pathogens are not seen, but using scanning electron microscopy (SEM), fungal spores are evident and associated with the scab‐like structures. Using culture‐dependent methods, cultures resembling a species of Elsinoe were isolated from the lesions. DNA sequence comparisons for four gene regions, as well as morphological observations, showed that the fungus is an undescribed species in Elsinoe, for which the name Elsinoe necatrix sp. nov. is provided. Pathogenicity trials on a Eucalyptus clone with the Elsinoe species resulted in scab‐like structures similar to those observed under field conditions and the fungus was easily reisolated from the resulting lesions. This study includes a description of the pathogen and characterization of the disease, for which the name Eucalyptus scab and shoot malformation is suggested.
Phytophthora pinifolia causes a needle and shoot disease in Pinus radiata, referred to as 'Daño Foliar del Pino'. This newly discovered disease requires intensive research efforts that necessitate the processing of large numbers of samples for which accurate identification, often by people not experienced in Phytophthora taxonomy, is required. The aim of this study was, therefore, to develop species-specific primers for P. pinifolia that amplify the internal transcribed spacer region of the ribosomal operon and the nuclear Ypt1 gene, respectively. The primers were tested over several Phytophthora spp., as well as fungi isolated from P. radiata. In all cases, only P. pinifolia was amplified. In addition to the species-specific primers, a PCR-restriction fragment length polymorphism protocol using available Phytophthora genus-specific primers was also used to generate a species-specific profile for P. pinifolia. This provided a characteristic profile that allows the identification of P. pinifolia, and it could also discriminate between 27 different species of Phytophthora. Both techniques reported in this study make it possible to identify large numbers of P. pinifolia cultures accurately and efficiently, which will be important for both quarantine work and biological research on this important new pathogen.
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