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1.
This study aimed to demonstrate the association of the ash dieback pathogen Hymenoscyphus fraxineus with leaf symptoms on Fraxinus excelsior and to test its pathogenicity towards leaves of three European ash species, F. excelsior, F. angustifolia and F. ornus, in wound inoculation experiments. On F. excelsior, H. fraxineus was isolated from 94% of leaf rachises with necrotic lesions and from 74% of necrotic leaflet midribs. Following wound inoculation of leaf rachises, in two separate experiments performed in 2010 and 2011, the ash dieback pathogen caused symptoms (necrotic rachis lesions, leaf wilting and premature leaf shedding) on all three ash species, while control leaves remained symptomless. Hymenoscyphus fraxineus was consistently reisolated from fungus‐inoculated rachises. All 10 isolates tested were pathogenic to the three ash species and varied in virulence. Koch's postulates for H. fraxineus as causal agent of leaf symptoms on F. excelsior were fulfilled in this study. Complemented with the isolation of the fungus from naturally infected, symptomatic leaf rachises of F. angustifolia and F. ornus in previous investigations, H. fraxineus was confirmed to be a leaf pathogen of these ash species as well. The leaf inoculation experiments showed that F. excelsior was highly susceptible to H. fraxineus, F. angustifolia was equally or slightly less susceptible, whereas F. ornus was the least affected species; however, F. ornus should also be regarded as a host tree for the ash dieback pathogen. This susceptibility ranking corresponds well with field observations and previous stem inoculation experiments.  相似文献   

2.
A large part of the area in Europe in which Fraxinus excelsior is native is currently affected by ash dieback, a threatening disease caused by the ascomycetous fungus Hymenoscyphus fraxineus. Fungi other than H. fraxineus also occur in large numbers on stems of the dying ash trees. To clarify their possible role in the dieback process, six fungal species common on dying stems and twigs of ash in Poland, i.e. Cytospora pruinosa, Diaporthe eres, Diplodia mutila, Fusarium avenaceum, F. lateritium and F. solani, were tested for pathogenicity using a test based on artificial wound inoculations of 6‐year‐old F. excelsior plants under field conditions, with H. fraxineus included for comparison. There were significant differences in index of pathogenicity among the fungi tested. Hymenoscyphus fraxineus (mean index 5.78) was the most pathogenic. Diplodia mutila (4.23) and C. pruinosa (4.02) were significantly less pathogenic than H. fraxineus, but significantly more than the other fungi. Diaporthe eres (2.43), F. avenaceum (1.92), F. solani (1.86) and F. lateritium (1.08) were the least pathogenic (< 0.0001). The extent of disease symptoms caused by F. solani and F. lateritium was statistically similar to the control (= 0.05). All tested fungi were successfully reisolated from inoculated stems. The contribution of the results to understanding the possible role of these fungi in the ash dieback process in F. excelsior, particularly in trees weakened after primary infection by H. fraxineus, is discussed.  相似文献   

3.
The future existence of common ash (Fraxinus excelsior), an important tree species throughout temperate Europe, is threatened. An invasive fungal disease (ash dieback) has spread through much of the distribution area of common ash. The causal agent of the disease is Hymenoscyphus fraxineus, a necrotrophic ascomycete, most probably introduced from Asia in the early 1990s. Hymenoscyphus fraxineus infects ash trees and saplings through their leaves, from which it grows into the stem. The fungus was studied intensively in recent years but there is still a need to address the topic from an evolutionary perspective. In this overview, some key evolutionary aspects of ash dieback are discussed, from the Red Queen dynamics of host–pathogen interactions to the probable consequences for virulence evolution of multiple infections. The progression of ash dieback in Europe does not show spatial differences, but studies show variation in susceptibility within host populations, a probable consequence of genetic differences, thus providing material for evolution of disease resistance or tolerance. Breeding programmes need to maintain the genetic diversity of Fraxinus, to enable it to withstand further threats such as climate change and the emerald ash borer. Because H. fraxineus reproduces exclusively sexually, the pathogen is likely to overcome a narrow genetic resistance. The introduction of further strains of H. fraxineus to Europe and the movement of infected plant material should be avoided. This case study shows that the integration of evolutionary ecology considerations would benefit plant disease management and biosecurity in general.  相似文献   

4.
Dieback of European ash was first observed in Europe in the early 1990s. The disease is caused by the invasive ascomycete Hymenoscyphus fraxineus, proposed to originate from Far East Asia, where it has been considered a harmless saprotroph. This study investigates the occurrence of H. fraxineus in tissues of local ash species in the Russian Far East, and assesses its population‐specific genetic variation by ITS sequencing. Shoot dieback symptoms, characteristic of H. fraxineus infection on European ash, were common, but not abundant, on Fraxinus mandshurica and Fraxinus rhynchophylla trees in Far East Russia. High levels of pathogen DNA were associated with necrotic leaf tissues of these ash species, indicating that the local H. fraxineus population is pathogenic to their leaves. However, the low levels of H. fraxineus DNA detected in shoots with symptoms, the failure to isolate this fungus from such tissues, and the presence of other fungi with pathogenic potential in shoots with symptoms indicate that local H. fraxineus strains may not be responsible (or their role is negligible) for the observed ash shoot dieback symptoms in the region. Conspicuous differences in ITS rDNA sequences detected between H. fraxineus isolates from Russian Far East and European populations suggest that the current ash dieback epidemic in Europe might not directly originate from the Russian Far East. Revision of the herbarium material shows that the earliest specimen of H. fraxineus was collected in 1962 from the Russian Far East and the oldest H. fraxineus specimen of China was collected in 2004.  相似文献   

5.
Ash dieback is caused by Hymenoscyphus fraxineus, a cryptic species of the putatively harmless Hymenoscyphus albidus. Recently, H. fraxineus was found to be native to East Asia. However, the virulence of Asian H. fraxineus strains on Fraxinus excelsior and the virulence of European H. albidus on hosts other than F. excelsior and Fraxinus mandshurica have not yet been assessed. In a wound inoculation study, the virulence of four H. albidus and four European and Japanese H. fraxineus strains was assessed on F. excelsior and Fraxinus pennsylvanica in a climate chamber. Lesion lengths were measured after approximately three and a half months. No lesions were observed on the negative control or on trees inoculated with H. albidus. In contrast, inoculation with H. fraxineus induced typical symptoms of ash dieback on both tree species. Japanese H. fraxineus strains induced significantly longer lesions compared to European strains. Fraxinus excelsior was highly susceptible and developed lesions averaging lengths of 1·7 and 8·4 cm for European and Japanese strains, respectively. Fraxinus pennsylvanica was less susceptible and developed average lesion lengths of 1·6 and 4·8 cm for European and Japanese strains, respectively. Most strains were successfully reisolated from necrotic lesions or inocula, fulfilling Koch's postulates. The data show that additional introductions of H. fraxineus strains from the native range to Europe could pose a threat to the conservation of F. excelsior. In addition, introduction of H. fraxineus to North America could potentially have a negative effect on the indigenous F. pennsylvanica.  相似文献   

6.
The ascomycete Hymenoscyphus fraxineus causes devastating damage to the European common ash (Fraxinus excelsior). The fungus originates from Asia, where it coexists with native ash species and completes its life cycle by sporulating on degrading ash leaf litter on the forest floor. Given this life cycle of the fungus, genotypes of Hfraxineus with varying degrees of aggressiveness may coexist in infected European ash forests. To test this hypothesis, we cultured 19 single-spore isolates from apothecia collected in a trial of heavily infected ash in Denmark and carried out stem inoculations on young ash seedlings. Microsatellite markers revealed that the 19 isolates were all genetically unique and did not show any genetic structure. High variation was observed among the 19 isolates in pathogenicity on young seedlings. The variation (assessed as necrosis development) was highly significant, but not correlated with the vigour of the fungal isolates when grown on culture media. The results support the hypothesis that aggressiveness of Hfraxineus may not be crucial for its fitness. In this sense, ash dieback disease may differ from other recent emerging infectious diseases on trees. We discuss the variation in aggressiveness of Hfraxineus in relation to durability of resistance and future management of the ash dieback epidemic. The findings of this study call for more research into natural variation in endophytic versus pathogenic behaviour of Hfraxineus on European ash.  相似文献   

7.
The ash dieback fungus, Hymenoscyphus fraxineus, a destructive, alien pathogen of common ash (Fraxinus excelsior), has spread across Europe over the past 25 years and was first observed in the UK in 2012. To investigate the relationship of the pathogen's population structure to its mode of arrival, isolates were obtained from locations in England and Wales, either where established natural populations of ash had been infected by wind‐dispersed ascospores or where the fungus had been introduced on imported planting stock. Population structure was determined by tests for vegetative compatibility (VC), mating type and single‐nucleotide polymorphisms (SNPs). VC heterogeneity was high at all locations, with 96% of isolate pairings being incompatible. Frequencies of the MAT1‐1‐1 and MAT1‐2‐1 idiomorphs were approximately equal, consistent with H. fraxineus being an obligate outbreeder. Most SNP variation occurred within study location and there was little genetic differentiation between the two types of location in the UK, or between pathogen populations in the UK and continental Europe. There was modest differentiation between UK subpopulations, consistent with genetic variation between source populations in continental Europe. However, there was no evidence of strong founder effects, indicating that numerous individuals of H. fraxineus initiated infection at each location, regardless of the route of pathogen transmission. The ssRNA virus HfMV1 was present at moderate to high frequencies in all UK subpopulations. The results imply that management of an introduced plant pathogen requires action against its spread at the continental level involving coordinated efforts by European countries.  相似文献   

8.
Olive leprosy, caused by the fungus Phlyctema vagabunda, is a classic fruit rot disease widespread in the Mediterranean basin. From 2009 to 2013, new disease symptoms consisting of small circular necrotic leaf lesions, coin branch canker and shoot dieback were observed in Spanish and Portuguese olive orchards showing intense defoliation. Phlyctema‐like anamorphs were consistently isolated from leaves and shoots with symptoms. Representative isolates from affected leaves, shoots and fruits were characterized based on morphology of colonies and conidia, optimum growth temperature and comparison of DNA sequence data from four regions: ITS, tub2, MIT and rpb2. In addition, pathogenicity tests were performed on apple and olive fruits, and on branches and leaves of olive trees. Maximum mycelial growth rate ranged between 0.54 and 0.73 mm per day. Conidia produced on inoculated apple fruits showed slight differences in morphology among the representative fungal isolates evaluated. Phylogenetic analysis clustered all of the Phlyctema‐like isolates in the same clade, identifying them as Phlyctema vagabunda. On fruits, influence of wounding, ripening and cultivar resistance was studied, with cv. Blanqueta being the most susceptible cultivar. On branches, a mycelial‐plug inoculation method reproduced olive leprosy symptoms and caused shoot dieback. On leaves, Koch's postulates were fulfilled and the pathogen caused characteristic necrotic spots and plant defoliation. This is the first time that the pathogenicity of P. vagabunda in olive leaves has been demonstrated.  相似文献   

9.
The ontogeny and morphology of infection structures associated with the early stages of infection of Hymenoscyphus pseudoalbidus on common ash (Fraxinus excelsior) leaves and leaf petioles were investigated using light and scanning electron microscopy. Ascospores were produced in mature ascocarps and infections on ash leaves were first observed 2 weeks later. Ascospores developed germ tubes, followed by appressorium formation and penetration of epidermal cells on ash leaves and petioles. Chalara fraxinea spores, the anamorph of H. pseudoalbidus, appeared and were arranged in chains, surrounded by a membranous sheath, and varied considerably in size and shape. Host invasion and colonization of all cell types of leaves and petioles were observed using light microscopy. The role of leaves and petioles as sites of infection in the life cycle of H. pseudoalbidus and the disease cycle of ash dieback is discussed.  相似文献   

10.
The causative agent of dieback on European ash (Fraxinus excelsior) was first described as Chalara fraxinea based on cultural morphology because no sexual stage of the fungus was known. Later, based on culturing of ascospores of a candidate teleomorph, morphological comparison and nuclear ribosomal internal transcribed spacer sequencing, the sexual stage of C. fraxinea was assigned as Hymenoscyphus albidus, a native and widespread species in Europe. Recently, the morphological species concept of H. albidus was shown to cover two species that cannot be separated from each other based on teleomorph characters, but which can be distinguished by several DNA markers. As a result, the strains causing ash dieback were reassigned as H. pseudoalbidus. The closely related H. albidus is presumably a non‐pathogenic endophyte, but pathogenicity tests to confirm this hypothesis have not yet been performed. Genotyping of herbarium specimens has shown that H. pseudoalbidus was present in Switzerland for at least a decade prior to the epidemic outbreak in Europe. The origin of the ash dieback pathogen, and the general importance of correct pathogen identification to development of effective disease control, are discussed.  相似文献   

11.
Aleppo pine is a thermophilic species that has become a popular tree species in plantings even beyond its natural habitat. In 1999 the pathogenic fungus Gremmeniella abietina was isolated for the first time in Aleppo pine plantations in Spain. The main objective of this study was to analyse the variation in susceptibility to G. abietina (European race, biotype A) infections among trees from five Aleppo pine provenances. Artificial inoculations were performed at two different times in January 2012. The susceptibility of the trees was evaluated by recording disease severity and the length of internal stem necrosis produced by the pathogen. In addition, the concentrations of two flavanone compounds were quantified as putative indicators of resistance. Provenances and G. abietina isolates both significantly affected levels of necrosis although the results were dependent on the time of inoculation. There was a relationship between provenance and resistance, with trees of higher elevation provenances being more susceptible to disease than trees of lower elevation provenances. The amount of naringenin flavanone from trees was significantly different between provenances, suggesting that it is a possible indicator value for resistance. Conversely, no differences were found in the concentration of eriodictyol in trees between provenances. The results show that the provenance‐dependent variation in the susceptibility of Aleppo pine to G. abietina should be considered in selection of seed sources for restoration.  相似文献   

12.
Cytospora species are ubiquitous pathogens of numerous woody plants, causing dieback and wood cankers in agronomic crops, timber trees and wildland trees (e.g. Prunus, Eucalyptus and Salix, respectively). Cytospora chrysosperma, C. cincta and C. leucostoma have been reported from grapevines in Iran showing symptoms of one or more recognized trunk diseases (esca, botryosphaeria‐, eutypa‐ and phomopsis diebacks); however, only C. chrysosperma was shown to be pathogenic to grapevine. To understand the potential role of Cytospora species in the grapevine trunk‐disease complex, 21 Cytospora isolates were examined that were recovered from dieback and wood cankers of Vitis vinifera and Vitis interspecific hybrids in seven northeastern U.S. states and two Canadian provinces. Phylogenetic analyses of ITS and translation elongation factor 1‐α identified two novel species: Cytospora vinacea sp. nov. and Cytospora viticola sp. nov. Differences in culture morphology and conidial dimensions also distinguished the species. When inoculated to the woody stems of potted V. vinifera ‘Thompson Seedless’ in the greenhouse, both species were pathogenic, based on development of wood lesions and fulfilment of Koch's postulates. Cytospora viticola was the most virulent based on lesion length at 12 months post‐inoculation. As cytospora canker shares some of the same general dieback‐type symptoms as botryosphaeria‐, eutypa‐ and phomopsis diebacks, it may be considered part of the grapevine trunk‐disease complex in eastern North America.  相似文献   

13.
Hymenoscyphus fraxineus is an invasive fungus in Europe and causes a severe decline affecting ash, which began in the late 1990s. One of the symptoms associated with the disease is lesions in the outer bark of the collar area. However, the aetiology of these basal lesions, and in particular the relative roles of H. fraxineus and Armillaria species, is still controversial; moreover, little is known about the influence of environmental factors on the disease epidemiology. This study therefore surveyed 42 plots located in northeastern France, in an area affected by ash decline since 2008, in order to determine which environmental factors condition the severity of lesions associated with H. fraxineus on ash collar. The spatial pattern that is a consequence of the invasive spread of the disease was taken into account in the analysis, using a spatial hierarchical Bayesian model fitted by integrated nested laplace approximation (INLA). Results show that while basal lesions are tightly associated with H. fraxineus, their severity is influenced by the Armillaria species present in the plot. Sites with vegetation indicating moist conditions, or more humid topographical positions, were associated with more developed basal lesions.  相似文献   

14.
Dieback of European ash (Fraxinus excelsior), caused by the ascomycete Hymenoscyphus pseudoalbidus (anamorph Chalara fraxinea), started around 1992 in Poland and has since then spread over large geographical areas. By November 2010, the disease had been recorded in 22 European countries. The gradual expansion and high intensity of the ash dieback epidemic in Europe may suggest that H. pseudoalbidus is an invasive alien organism. In Norway, ash dieback was first reported in spring 2008, and a survey in early summer of the same year revealed that the disease had spread over large parts of the southern and eastern regions of the country. The distance from the southernmost to the northernmost infected stands was, at that time, about 400 km. Some old necrotic lesions were also observed, indicating that the ash dieback pathogen is likely to have been present in Norway since at least 2006. In 2009, a spore sampler was installed in a diseased ash stand at Ås, South‐Eastern Norway. Sampling started in late July and continued until late September. Large numbers of ascospores resembling those of H. pseudoalbidus were observed, with the maximum number of spores occurring from the end of July to mid‐August. The deposition of ascospores occurred mainly between 6 and 8 a.m. Ascospores are most likely to be the primary source initiating host infections and responsible for the rapid recent spread of H. pseudoalbidus in Europe.  相似文献   

15.
Puccinia psidii has long been considered a significant threat to Australian plant industries and ecosystems. In April 2010, P. psidii was detected for the first time in Australia on the central coast of New South Wales (NSW). The fungus spread rapidly along the east coast and in December 2010 was found in Queensland (Qld) followed by Victoria a year later. Puccinia psidii was initially restricted to the southeastern part of Qld but spread as far north as Mossman. In Qld, 48 species of Myrtaceae are considered highly or extremely susceptible to the disease. The impact of P. psidii on individual trees and shrubs has ranged from minor leaf spots, foliage, stem and branch dieback to reduced fecundity. Tree death, as a result of repeated infection, has been recorded for Rhodomyrtus psidioides. Rust infection has also been recorded on flower buds, flowers and fruits of 28 host species. Morphological and molecular characteristics were used to confirm the identification of P. psidii from a range of Myrtaceae in Qld and compared with isolates from NSW and overseas. A reconstructed phylogeny based on the LSU and SSU regions of rDNA did not resolve the familial placement of P. psidii, but indicated that it does not belong to the Pucciniaceae. Uredo rangelii was found to be con‐specific with all isolates of P. psidii in morphology, ITS and LSU sequence data, and host range.  相似文献   

16.
Genetic, phenotypic and host range diversity among Pseudomonas savastanoi isolates from Myrtus communis were investigated. Thirty‐one isolates from six Sardinian commercial myrtle orchards and three isolates from plants growing spontaneously on the island of Rhodes (Greece) were compared with reference strains of Psavastanoi from olive, oleander, ash and myrtle. Multilocus sequence analysis (MLSA) indicated the presence of a monomorphic population with a very low level of variability. Conversely, Biolog phenotypic fingerprinting and phytohormone production analyses showed a considerable metabolic diversity, as bacteria obtained from single infected tissue differed more than bacteria obtained from different orchards. When pathogenicity tests were carried out on myrtle plants, different types of symptoms were induced: knots, canker lesions with or without tissue proliferations and, occasionally, wilting of the inoculated twig, a symptom never reported before for Psavastanoi. Comparable symptoms were also observed in the natural environment both on spontaneous and cultivated plants. Moreover, the host range of the myrtle population was heterogeneous and not well defined. Some isolates showed a wide host range whilst others were pathogenic only to their natural host. Overall these findings suggest that the diversity of the Psavastanoi population from myrtle does not depend so much on the locality or the natural host and does not allow the Sardinian and Greek isolates, together with previously characterized myrtle strains, to be ascribed to a known pathovar of Psavastanoi, nor to propose their belonging, as a whole, to a new pathovar.  相似文献   

17.
Conifer species, which have formed the foundation of commercial forestry industry in many countries, are known to be invasive in natural ecosystems, especially in the Southern Hemisphere. Controlling isolated invasive conifers before they reach reproductive maturity is an essential element of any strategy that aims to reduce spread rate of these species. Using a novel helicopter‐mounted spot‐application gun, which delivers a precise dosage to the crown of each tree, the objective of this research was to test the efficacy of three triclopyr‐based treatments against the four most vigorous wilding conifer species (Pinus contorta, Pinus nigra, Pinus sylvestris and Pseudotsuga menziesii) under New Zealand field conditions. Herbicides tested were triclopyr at two different rates and a combination of triclopyr and picloram. Treated trees covered a wide range of heights (c. 0.5–16 m), and measurements of mortality taken two years post‐herbicide application were used to examine variation in efficacy of the herbicides. Successful treatment was defined by a mortality rate of 85% or higher. A logistic regression model was fitted to the mortality data and used to derive threshold tree heights at which 85% mortality occurred, H85. For all four species, the most effective treatment was application of 1000 ml of herbicide mixture per tree that contained 120 and 20 g, respectively, of the active ingredients triclopyr and picloram. There was a significant decline in efficacy of this treatment with increases in tree size for all four species. Values of H85 for this treatment were 7.4 m for P. nigra, 8.3 m for P. menziesii, 9.7 m for P. contorta and >10 m for P. sylvestris. The methods developed here could be used to effectively manage emerging conifer infestations before they become problematic.  相似文献   

18.
A survey of bleeding canker disease, caused by Pseudomonas syringae pv. aesculi, was undertaken across Ireland. Incidence has become severe and can be considered epidemic, as 61% of the 1587 horse chestnut trees surveyed showed symptoms of the disease. Bacteria were isolated from a sample of trees and characterized using gyrBDNA sequencing. DNA was also extracted directly from wound tissue. The Irish P. syringae pv. aesculi genotype was identical to genotypes previously sequenced with gyrB from the UK and some other locations in Europe. Real‐time PCR, using existing primers and a newly designed, more pathovar‐specific primer set, was assessed for use in disease screening. With molecular screening, a total of 11 trees from a sample of 55 tested positive for P. syringae pv. aesculi in Ireland. It was more efficient to extract DNA directly from wound tissue, especially fresh bark, for disease detection than to undertake bacterial isolation with subsequent molecular analysis. A further set of sequencing primers was developed for the amplification of the gyrB gene from P. syringae pv. aesculi and their specificity was shown using a diverse sample of bacterial isolate DNAs. The study also isolated and identified other bacterial species from diseased material; some of these are known pathogens (Brenneria nigrifluens, P. marginalis and P. syringae) or have previously been identified as potentially beneficial endophytes of host trees (Erwinia billingiae, E. tolentana, P. fluorescens, P. putida and Raoultella).  相似文献   

19.
Citrus postbloom fruit drop (PFD) is caused by Colletotrichum acutatum and C. gloeosporioides. These pathogens attack the flowers and cause premature fruit drop and the retention of fruit calyces. This study was designed to characterize the spatial and temporal dynamics of PFD in commercial citrus‐growing areas to better understand the disease spread. Experiments were carried out in three young orchards (500 trees each) in two municipalities in Sao Paulo State, Brazil. Symptoms of PFD on the flowers and presence of persistent calyces were assessed in each of three orchards for three years. Logistic, Gompertz and monomolecular models were fitted to the incidence data over time from the trees with symptoms. The spatial pattern of diseased trees was characterized by a dispersion index and by Taylor′s power law. An autologistic model was used for the spatiotemporal analysis. The logistic model provided the best fit to the disease incidence data, which had a fast progress rate of 0·53 per day. During the early epidemic of PFD, the spatial pattern of diseased trees was random, which suggested that inoculum spread was due to mechanisms other than rain splash. As the disease incidence increased (up to 12·6%), the spatial pattern of diseased trees became aggregated. The rapid rate of disease progress and the distribution of PFD suggest that dispersal of the pathogen is possibly related to a mechanism other than splash dispersal, which is more typical of other fruit diseases caused by Colletotrichum spp.  相似文献   

20.
The purpose of this study was to test the hypothesis that Chrysoporthe cubensis on native trees in South America could be the source of the pathogen that causes severe stem cankers and often mortality in commercially propagated Eucalyptus trees. This was done by investigating populations originating from two adjacent Eucalyptus (Myrtaceae) plantations in Colombia, and wild Miconia rubiginosa trees (Melastomataceae) growing alongside these stands. Polymorphic microsatellite markers were used to quantify allele sizes in 20 and 39 isolates from the two Eucalyptus stands and 32 isolates from adjacent M. rubiginosa trees. Gene and genotypic diversities were calculated from these data, and population differentiation and assignment tests were performed to ascertain whether the populations were genetically different. Results showed that there were no differences between any of the populations using these techniques, and that they can be treated as a single population. Therefore, the results support the hypothesis that host switching has occurred in C. cubensis in Colombia.  相似文献   

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