This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain. M A N U S C R I P T A C C E P T E D ACCEPTED MANUSCRIPT RESEARCH HIGHLIGHTS• P. cinnamomi survival propagules in annuals/herbaceous perennials in nature • Formation of stromata by Phytophthora cinnamomi, novel for the species • Dense oospore clusters of P. cinnamomi, forming viable colonies in new hosts • Thick walled chlamydospores inside naturally infected roots • Presence of haustoria suggest a biotrophic mode in asymptomatic host species M A N U S C R I P T A C C E P T E D ACCEPTED MANUSCRIPT ABSTRACTStudies were conducted to determine how Phytophthora cinnamomi survives during hot and dry Mediterranean summers in areas with limited surviving susceptible hosts.Two Western Australian herbaceous perennials Chamaescilla corymbosa and Stylidium diuroides and one Western Australian annual Trachymene pilosa were collected weekly from a naturally infested site from the Eucalyptus marginata (jarrah) forest from winter to spring and less frequently during summer 2011/12. Selfed oospores, thick walled chlamydospores and stromata of P. cinnamomi were observed in each species. Oospores and thick-walled chlamydospores germinated in planta confirming their viability. This is the first report of autogamy by P. cinnamomi in naturally infected plants. Stromata, reported for the first time for P. cinnamomi, were densely aggregated inside host cells, and germinated in planta with multiple germ tubes with hyphae capable of producing oospores and chlamydospores.Trachymene pilosa was completely asymptomatic, S. diuroides did not develop root lesions but some plants wilted, whilst C. corymbosa remained asymptomatic above ground but lesions developed on some tubers. The presence of haustoria suggests that P. cinnamomi grows biotrophically in some hosts. Asymptomatic, biotrophic growth of P. cinnamomi in some annual and herbaceous perennials and the production of a range of survival structures have implications for pathogen persistence over summer and its management. KEYWORDSPhytophthora cinnamomi, survival structures, life cycle, facultative homothallic, biotrophic growth, endophyte, haustoria M A N U S C R I P T A C C E P T E D ACCEPTED MANUSCRIPT 2 INTRODUCTIONPhytophthora cinnamomi is a soil-borne root pathogen with a broad host range and necrotrophic mode of infection (Cahill et al. 2008; Zentmyer 1980) and results in the death of many susceptible plant species and the degradation of ecosystems worldwide including 15 global biodiversity hotspots (Dunstan et al. 2010). Phylogenetically and taxonomically, this pathogen belongs to the class Oomycetes in which swimming zoos...

Resistant annual and herbaceous perennial plant species were identified as key hosts which allow Phytophthora cinnamomi to persist on severely impacted black gravel sites within the Eucalyptus marginata (jarrah) forest of southwest Western Australia. Of the annual and herbaceous perennial plant species present on black gravel sites, 15 out of 19 species were found to be hosts of P. cinnamomi, and 10 of these were symptomless hosts. In particular, the native annual Trachymene pilosa and the two native herbaceous perennials Stylidium diuroides and Chamaescilla corymbosa were commonly found to be hosts of the pathogen. Species from 12 new genera including three from new families (Crassulaceae, Droseraceae and Primulaceae) are reported for the first time to be hosts of P. cinnamomi. The species from which P. cinnamomi was recovered were the native species:

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Summary Phytophthora cinnamomi is a necrotrophic pathogen of woody perennials and devastates many biomes worldwide. A controlled perlite–hydroponic system with no other hyphae‐producing organisms as contaminants present allowed rapid assessment of ten annual and herbaceous perennial plant species most of which have a wide distribution within the jarrah (Eucalyptus marginata) forest in Western Australia where this pathogen has been introduced. As some annuals and herbaceous perennials have recently been reported as symptomatic and asymptomatic hosts, laboratory screening of some of the field‐tested annuals and herbaceous perennials and additional species was used to further evaluate their role in the pathogen's disease cycle. Nine of the species challenged with the pathogen were asymptomatic, with none developing root lesions; however, Trachymene pilosa died. The pathogen produced thick‐walled chlamydospores and stromata in the asymptomatic roots. Furthermore, haustoria were observed in the roots, indicating that the pathogen was growing as a biotroph in these hosts.