Austropuccinia Psidii
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''Austropuccinia psidii'', commonly known as myrtle rust, guava rust, or ʻōhiʻa rust; is a
rust Rust is an iron oxide, a usually reddish-brown oxide formed by the reaction of iron and oxygen in the catalytic presence of water or air moisture. Rust consists of hydrous iron(III) oxides (Fe2O3·nH2O) and iron(III) oxide-hydroxide (FeO(OH ...
(a type of
plant pathogen Plant pathology (also phytopathology) is the scientific study of diseases in plants caused by pathogens (infectious organisms) and environmental conditions (physiological factors). Organisms that cause infectious disease include fungi, oomyc ...
) native to
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...
that affects plants in the family
Myrtaceae Myrtaceae, the myrtle family, is a family of dicotyledonous plants placed within the order Myrtales. Myrtle, pōhutukawa, bay rum tree, clove, guava, acca (feijoa), allspice, and eucalyptus are some notable members of this group. All speci ...
. It is a member of the fungal complex called the guava rust (''Puccinia psidii'') group. The
spore In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, f ...
s have a distinctive yellow to orange colour, occasionally encircled by a purple ring. They are found on lesions on new growth including shoots, leaves, buds and fruits. Leaves become twisted and may die. Infections in highly susceptible species may result in the death of the host plant. As of late 2013, it is infecting around 179 species in
New South Wales ) , nickname = , image_map = New South Wales in Australia.svg , map_caption = Location of New South Wales in AustraliaCoordinates: , subdivision_type = Country , subdivision_name = Australia , established_title = Before federation , es ...
and
Queensland ) , nickname = Sunshine State , image_map = Queensland in Australia.svg , map_caption = Location of Queensland in Australia , subdivision_type = Country , subdivision_name = Australia , established_title = Before federation , established_ ...
, from 41 genera (around 46% of genera in the Myrtaceae) in Australia.


Importance

''Austropuccinia psidii'', a myrtle rust or a rust fungus, has a large host range for infection making it difficult for disease control and prevention, but it has been seen to have biological uses for it. In Florida, the pathogen has been considered as a biological control agent for the invasive Australian plant, ''
Melaleuca quinquenervia ''Melaleuca quinquenervia'', commonly known as the broad-leaved paperbark, paper bark tea tree, punk tree or niaouli, is a small- to medium-sized tree of the myrtle family, Myrtaceae. It grows as a spreading tree up to tall, with its trunk cov ...
''. ''Melaleuca quinquenervia'' is a weed tree that have invaded south Florida and if left unchecked, it will cause of the most serious threats to the integrity native ecosystem, turning marshes in the Everglades into a swamp. Thus, with Autropuccinia psidii, the disease has to ability to disperse inoculum rapidly with the ability to attack healthy tissues, causing substantial levels of damage to their hosts. Thus, creating pustules and evenly death of the invasive species. ''Austropuccinia psidii'' can rapidly spread globally and can impact commercially and ecologically important species of Myrtaeceae such as '' Archirhodomyrtus beckleri'', ''
Decaspermum humile ''Decaspermum humile'', commonly known as the silky myrtle, is a tree from Australia and Asia. It can be used as bush food, as indicated by the alternate common name of currant myrtle. The tree features an attractive dark glossy crown. The new pi ...
'', '' Gossia hillii'' and '' Rhodamnia maideniana,'' that are extremely important in Australia affecting their native forest ecosystems and causing extinctions. ''Austropuccinia psidii'' has also been found to also be an invasive species that infects young growing tissues of plants. It infects developed leaves, floral bud, fruits and Coppice. This fungus is a serious problem because of its ability to spread rapidly from its production of a large number of small spores that can be easily dispersed over long distances by wind. These spores can also be dispersed by vectors that include animals such as birds, bats, possums, and insects that have been in contact with the rust spores. ''Austropuccinia psidii'' has a potential to cause damage to natural ecosystems. In Australia, many ecosystems are dominated by ''
Eucalyptus ''Eucalyptus'' () is a genus of over seven hundred species of flowering trees, shrubs or mallees in the myrtle family, Myrtaceae. Along with several other genera in the tribe Eucalypteae, including '' Corymbia'', they are commonly known as euca ...
'' which is also the dominant tree in most of Hawaii's forests. Infection from ''Autropuccinia psidii'' results in significant changes to the structure, composition, and the function of forests trees on a landscape level. Thus, it greatly impacts biodiversity of other flora and fauna in these ecosystems.


Taxonomy

The fungus was first described scientifically in 1884 as ''Puccinia psidii'' by G. Winter, then described again in 2006 as ''Uredo rangelii'' by mycologists J.A. Simpson, K. Thomas, and Cheryl Grgurinovic. Finally in 2017 the names were synonymised by Beenken in a new genus as ''Austropuccinia psidii''''.''


Development and symptoms

Myrtle rust is typically characterised by the appearance of urediniospores on the underside of the leaf, though urediniospores may also be found on the top of the leaf or on young stems. Initially, the disease appears as small purple or red brown flecks with a faint chlorotic halo on the leaf surface, which coalesce to form bright yellow pustules. As the rust develops, these pustules often fade to a grey brown colour. A high degree of pustule coalescence can result in distortion of the leaf. Myrtle rust also makes plants more susceptible to secondary infections, which may occur within days of the initial appearance of the pustules. Favourable conditions that increase the infection rate include: new tissue; high humidity; free water on plant surface for more than 6 hours; moderate temperatures, around 15–25 °C. Low light conditions (minimum of 8 hours) after spore contact can increase germination. The main ways in which myrtle rust can be spread are by: the movement of infected plant material, the movement of contaminated equipment, wind, water and gravity, animals, humans and/or vehicles. Myrtle rust may remain on a single host plant to complete its life cycle, which can be as short as 10–14 days.


As an invasive species

''Puccinia psidii'' can have very serious consequences to various species of plants in the Myrtaceae. This family includes
guava Guava () is a common tropical fruit cultivated in many tropical and subtropical regions. The common guava ''Psidium guajava'' (lemon guava, apple guava) is a small tree in the myrtle family ( Myrtaceae), native to Mexico, Central America, the ...
(the original host of this rust in Brazil),
eucalyptus ''Eucalyptus'' () is a genus of over seven hundred species of flowering trees, shrubs or mallees in the myrtle family, Myrtaceae. Along with several other genera in the tribe Eucalypteae, including '' Corymbia'', they are commonly known as euca ...
,
melaleuca ''Melaleuca'' () is a genus of nearly 300 species of plants in the myrtle family, Myrtaceae, commonly known as paperbarks, honey-myrtles or tea-trees (although the last name is also applied to species of '' Leptospermum''). They range in size ...
, and a number of species native to
Hawaii Hawaii ( ; haw, Hawaii or ) is a state in the Western United States, located in the Pacific Ocean about from the U.S. mainland. It is the only U.S. state outside North America, the only state that is an archipelago, and the only stat ...
, including some endemic species (found nowhere else on Earth) and at least one important native forest tree. There are numerous strains of the ''Puccinia psidii'' rust—some known to be established in
Florida Florida is a state located in the Southeastern region of the United States. Florida is bordered to the west by the Gulf of Mexico, to the northwest by Alabama, to the north by Georgia, to the east by the Bahamas and Atlantic Ocean, and to ...
, and at least one reported from
California California is a U.S. state, state in the Western United States, located along the West Coast of the United States, Pacific Coast. With nearly 39.2million residents across a total area of approximately , it is the List of states and territori ...
and there is concern that strains may exist or develop that could be devastating to ʻōhiʻa ('' Metrosideros polymorpha''), one of Hawaii's dominant native trees, a foundation species for many remaining Hawaiian native ecosystems. However currently, the major threat of Puccinia in Hawaii is the massive damage it is doing to '' Eugenia koolauensis'', a federally listed endangered species. Myrtle rust was first recorded in
Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
in mid-2010 and currently poses a major threat to the continent's ecosystem given that almost 80 per cent of Australian native trees are Mytraceae, most indigenous species rely on healthy trees for their survival. Additionally it poses a major threat to Australia's primary industry sector. Its current range includes much of the eastern coastal fringe of the Australian mainland. Initial detection was in April 2010 in
Gosford Gosford is the city and administrative centre of the Central Coast Council local government area in the heart of the Central Coast region, about north of Sydney and about south of Newcastle. The city centre is situated at the northern extre ...
in the Central Coast region of
New South Wales ) , nickname = , image_map = New South Wales in Australia.svg , map_caption = Location of New South Wales in AustraliaCoordinates: , subdivision_type = Country , subdivision_name = Australia , established_title = Before federation , es ...
. It was initially quarantined and eradication thought viable. The New South Wales government spent $5 million attempting to eradicate the disease. However, efforts to contain it failed and it spread rapidly north and south along the eastern coast. In response to the increasing threat, a Myrtle Rust National Management Group was formed on 2 July 2010 with the aim of eradication however due to the extent of its spread at that point of time, the group conceded that it had become impossible to eradicate. By December 2010, it had significantly spread north along the coast and recorded in South East Queensland with isolated cases in
Far North Queensland Far North Queensland (FNQ) is the northernmost part of the Australian state of Queensland. Its largest city is Cairns and it is dominated geographically by Cape York Peninsula, which stretches north to the Torres Strait, and west to the Gulf C ...
cities of
Cairns Cairns (, ) is a city in Queensland, Australia, on the tropical north east coast of Far North Queensland. The population in June 2019 was 153,952, having grown on average 1.02% annually over the preceding five years. The city is the 5th-most-p ...
and Townsville. In January 2012, an isolated myrtle rust outbreak was reported in Victoria beginning in
Melbourne Melbourne ( ; Boonwurrung/Woiwurrung: ''Narrm'' or ''Naarm'') is the capital and most populous city of the Australian state of Victoria, and the second-most populous city in both Australia and Oceania. Its name generally refers to a met ...
's southern and eastern suburbs. Initial attempts to contain it were unsuccessful and by April, 2012 it had spread across much of the state via regional cities. By late 2015 myrtle rust was widespread in Queensland, NSW and Victoria. It has reached Tasmania, where it was detected in garden plants in the north-east in February 2015, and the Northern Territory, where it was detected on Melville Island in May 2015. The Tasmanian government is seeking to contain and eradicate myrtle rust from the state while the Northern Territory government has determined it is not possible to contain or eradicate the pathogen. In April 2017, New Zealand's Ministry for Primary Industries reported that myrtle rust had been detected on Raoul Island, off the New Zealand mainland; the following month, it was detected on the mainland, in
Kerikeri Kerikeri () is the largest town in Northland, New Zealand. It is a tourist destination north of Auckland and north of the northern region's largest city, Whangarei. It is sometimes called the Cradle of the Nation, as it was the site of th ...
. As of 2020 it is spreading in New Zealand, and infecting pōhutukawa ('' Metrosideros excelsa''), northern rātā ('' Metrosideros robusta''), southern rātā ('' Metrosideros umbellata''), ramarama (''
Lophomyrtus bullata ''Lophomyrtus bullata'', also known by its Māori name of ramarama, is a species of evergreen myrtle shrub in the genus ''Lophomyrtus'', family Myrtaceae. It is endemic to New Zealand. ''Lophomyrtus bullata'' grows to a height of 8 metres, pro ...
'') and rōhutu ''
Lophomyrtus obcordata ''Lophomyrtus obcordata'', also known as rōhutu or tutuhi, is a species of evergreen myrtle shrub in the genus ''Lophomyrtus'', family Myrtaceae. It is found throughout New Zealand New Zealand ( mi, Aotearoa ) is an island country in t ...
''.


Host genera

Species within the following plant genera have been recorded with the infection: *'' Acmena'' *'' Agonis'' *''
Angophora ''Angophora'' is a genus of nine species of trees and shrubs in the myrtle family, Myrtaceae. Endemic to eastern Australia, they differ from other eucalypts in having juvenile and adult leaves arranged in opposite pairs, sepals reduced to projec ...
'' *'' Asteromyrtus'' *''
Austromyrtus ''Austromyrtus'' is a genus of shrubs in the myrtle family Myrtaceae. Three species are found along the east coast of Australia; in Queensland and in New South Wales and A. lotoides being endemic to New Caledonia. The fruits of ''A. dulcis'' hav ...
'' *'' Backhousia'' *''
Callistemon ''Callistemon'' is a genus of shrubs in the family Myrtaceae, first described as a genus in 1814. The entire genus is endemic to Australia but widely cultivated in many other regions and naturalised in scattered locations. Their status as a se ...
'' *''
Chamelaucium ''Chamelaucium'', also known as waxflower, is a genus of shrubs endemic to south western Western Australia. They belong to the myrtle family Myrtaceae and have flowers similar to those of the tea-trees (''Leptospermum''). The most well-known spe ...
'' *''
Choricarpia ''Backhousia'' is a genus of thirteen currently known species of flowering plants in the family Myrtaceae. All the currently known species are endemic to Australia in the rainforests and seasonally dry forests of Queensland, New South Wales and ...
'' *'' Decaspermum'' *''
Eucalyptus ''Eucalyptus'' () is a genus of over seven hundred species of flowering trees, shrubs or mallees in the myrtle family, Myrtaceae. Along with several other genera in the tribe Eucalypteae, including '' Corymbia'', they are commonly known as euca ...
'' *'' Eugenia'' *'' Gossia'' *''
Lenwebbia ''Lenwebbia'' is a genus of shrubs or small trees in the myrtle family Myrtaceae.Snow et al. 2003 The genus is named to honour the Australian plant ecologist Dr. Leonard Webb. The genus occurs in mesic forests along or near the east coast of Au ...
'' *''
Leptospermum ''Leptospermum'' is a genus of shrubs and small trees in the myrtle family Myrtaceae commonly known as tea trees, although this name is sometimes also used for some species of ''Melaleuca''. Most species are endemic to Australia, with the greate ...
'' *''
Lophomyrtus ''Lophomyrtus'' is a genus of the myrtle family described as a genus in 1941. The entire genus is endemic to New Zealand. It consists of evergreen shrubs or trees, noted for their colorful leaves, which are purple, chocolate, red or bronze-gree ...
'' *''
Melaleuca ''Melaleuca'' () is a genus of nearly 300 species of plants in the myrtle family, Myrtaceae, commonly known as paperbarks, honey-myrtles or tea-trees (although the last name is also applied to species of '' Leptospermum''). They range in size ...
'' *''
Metrosideros ''Metrosideros'' is a genus of approximately 60 trees, shrubs, and vines mostly found in the Pacific region in the family Myrtaceae. Most of the tree forms are small, but some are exceptionally large, the New Zealand species in particular. The n ...
'' *''
Myrtus ''Myrtus'' (commonly called myrtle) is a genus of flowering plants in the family Myrtaceae. It was first described by Swedish botanist Linnaeus in 1753. Over 600 names have been proposed in the genus, but nearly all have either been moved ...
'' *'' Pilidiostigma'' *''
Rhodamnia ''Rhodamnia'' is a group of rainforest trees and shrubs in the myrtle family described as a genus in 1822.Govaerts, R., Sobral, N., Ashton, P., Barrie, F., Holst, B.K., Landrum, L.L., Matsumoto, K., Fernanda Mazine, F., Nic Lughadha, E., Proenç ...
'' *''
Rhodomyrtus ''Rhodomyrtus'' is a group of shrubs and trees in the family Myrtaceae described as a genus in 1841. The genus is native to southern China, the Indian Subcontinent, Southeast Asia, Melanesia, and Australia. Its greatest levels of diversity are o ...
'' *''
Ristantia ''Ristantia'' is a group of plants in the family Myrtaceae described as a genus in 1982. The entire genus is endemic to the State of Queensland in Australia. ;Species # ''Ristantia gouldii'' Peter G. Wilson & B. Hyland - a tree species listed a ...
'' *''
Stockwellia ''Stockwellia'' is a monotypic genus in the flowering plant family Myrtaceae. The sole species in the genus, ''Stockwellia quadrifida'' (commonly known as Vic Stockwell's puzzle), is endemic to Queensland. Description ''Stockwellia quadrifida' ...
'' *''
Syncarpia ''Syncarpia'' is a small group of trees in the myrtle family (Myrtaceae) described as a genus in 1839. They are native to Queensland and New South Wales in Australia.Govaerts, R., Sobral, N., Ashton, P., Barrie, F., Holst, B.K., Landrum, L.L., ...
'' *'' Syzygium'' *'' Tristania'' *''
Ugni ''Ugni'' is a genus of plants in the myrtle family Myrtaceae, described as a genus in 1848. It is native to western Latin America from the Valdivian temperate rain forests of southern Chile (including the Juan Fernández Islands) and adjacent ...
'' *''
Uromyrtus ''Uromyrtus'' is a genus of plants in the myrtle family Myrtaceae described as a genus in 1941. The greatest diversity of species are found in New Caledonia and the remainder are found in Australia, New Guinea and Borneo.Govaerts, R., Sobral, N., ...
'' *''
Waterhousea ''Syzygium'' () is a genus of flowering plants that belongs to the myrtle family, Myrtaceae. The genus comprises about 1200 species, and has a native range that extends from Africa and Madagascar through southern Asia east through the Pacific. I ...
'' *'' Xanthostemon''


Environmental impacts

Since first being detected in 2010, myrtle rust has spread rapidly with entire plant species now under threat. In Australia, the family Myrtaceae - which includes eucalypts, melaleuca and lilly pilly - is diverse, widespread and important to many native ecosystems. The impact of myrtle rust has now been seen in a range of forest ecosystems including coastal heath, coastal and river wetlands, sand island ecosystems and subtropical and tropical rainforests. A number of plant species are now at risk of becoming extinct with about 40 plant species considered highly susceptible, such as the endangered ''Rhodamnia angustifolia''. Native animals are also likely to suffer significant impacts. Myrtle rust grows in shoots, fruits and flowers, destroying the food relied on by some species of flying foxes, lorikeets and honey eaters. There is the strong possibility that some of these species will become regionally extinct, and their loss could have serious flow-on effects.


Life cycle and disease cycle

Rust fungi typically have complex life cycles that include stages of sexual and asexual reproduction that occasionally occur on phylogenetically distinct host plants. ''Austropuccinia psidii''’s life cycle is quite controversial with one study showing that the rust is
autoecious A heteroecious parasite is one that requires at least two hosts. The ''primary host'' is the host in which the parasite spends its adult life; the other is the ''secondary host''. Both hosts are required for the parasite to complete its life cycle ...
while another study finding it to be
heteroecious A heteroecious parasite is one that requires at least two hosts. The ''primary host'' is the host in which the parasite spends its adult life; the other is the ''secondary host''. Both hosts are required for the parasite to complete its life cycle ...
but with an alternate host that has yet to be found. Looking at the fungus as a polycyclic pathogen, Stage I consists of aeciospores inoculating young leaf/shoot/fruit/flower bud. Once infected, aeciospores germinate and penetrate the host by creating a haustoria. Colonies and urediniosori forms after penetration. In Stage II, the production of urediniospores go back to inoculate young leaf/shoot/fruit/flower bud, important for secondary infection and contributing to the polycyclic nature of the pathogen. Stage II is always is also where urediniospores germinate, penetration of the host occurs, resulting in the development of an uridiniosori. In Stage III, urediniospores germinate and the host is penetrated with the development of a haustoria. After penetration, colonies are formed and teliosori are developed. Teliosori produce teliospores which can further germinate. Once they're germinated, basidiospores are developed. In Stage IV, basidiospores are spread, inoculating young leaf/shoot/fruit/flower bud. As a result, basidiospores germinate penetrating the host with the development of a haustorium. Aeciosori is formed with the production of aeciospores and the cycle repeats.


Infection pathway

Beginning with 1) extracellular pre-penetration processes (adhesion, germination, and
appressorium An appressorium is a specialized cell typical of many fungal plant pathogens that is used to infect host plants. It is a flattened, hyphal "pressing" organ, from which a minute infection peg grows and enters the host, using turgor pressure capable ...
formation 2) intercellular post-penetration processes (nutrient acquisition, ETS, ETI) and 3) dispersal of newly formed uredospores (reproduction and sporulation). To infect the plant, uredospores must adhere to the leaf surface which can be enhanced by the secretion of extracellular compounds. Both physical and chemical cues by neutralized spore germination inhibitors induces germination and break
dormancy Dormancy is a period in an organism's life cycle when growth, development, and (in animals) physical activity are temporarily stopped. This minimizes metabolic activity and therefore helps an organism to conserve energy. Dormancy tends to be clo ...
. After the penetration of urediniospores by a combination of extracellular secretions and physical force, the germ tube is formed and elongated in a favorable area fueled by spore protein reserves from an appressorium. Once the hyphae is within the cell apoplast, the hyphae penetrates a host
mesophyll cell A leaf (plural, : leaves) is any of the principal appendages of a vascular plant plant stem, stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", wh ...
to form a haustoria to acquire nutrients and suppress plant defenses. The hyphae and haustoria continually secrete effectors prevent plant resistance. Chemical signaling between plant and pathogen are continuously occurring to determine infection or any defense responses to the pathogen. Nutrient acquisition fuels pathogen growth by the accumulation of carbon from the host plant. This results in sporulation where the hyphae move toward plant surfaces to make urediniospores and to create a sori that ruptures through the dermal surface for penetration and infection.


Management

The original plan to eradicate myrtle rust from Australia was declared to be infeasible by the Myrtle Rust National Management Group in December 2010. The Myrtle Rust Response Plan was cancelled and focus was placed on minimising the spread and the impacts on myrtle rust. The Australian Government, through the Department of Agriculture Fisheries and Forestry, established the Myrtle Rust Coordination Group to manage the investment of $1.5 million of research funding. In 2016, The National Environmental Science Programme (http://www.environment.gov.au/science/nesp) hosted a national workshop on myrtle rust to discuss research findings and future management options. Participants included the Cooperative Research Centre for Plant Biosecurity, state and federal agencies, and botanical and plant conservation experts. Discussions centred around the impact on native species in Australia. A key outcome of the workshop included agreement that there is a need for a nationally coordinated approach through a long-term National Action Plan which aims to ensure that no species or ecosystems are lost to its impact. Practical measures to minimise the risk of increasing the distribution of myrtle rust include: not moving plant matter from one site to another; minimising pathogen spread by arriving and leaving each site clean of the pathogen, and avoiding areas that may contain myrtle rust-infected plant matter.


References


External links

*
'Beyond Myrtle Rust' New Zealand Research programme
* 2010. Loope, Lloyd L
A Summary of Information on the Rust ''Puccinia psidii'' Winter (Guava Rust) with Emphasis on Means to Prevent Introduction of Additional Strains to Hawaii
U.S. Geological Survey Open-File Report 2010–1082. * '
''Puccinia psidii'' species information
'' from th
Hawaiian Ecosystems at Risk project (HEAR)
* 2007. Eloise M. Killgore & Ronald A. He
Ohia Rust
{{Taxonbar, from1=Q7900369, from2=Q7258196 Fungal plant pathogens and diseases Pucciniales Fungi described in 2006 Fungi described in 2017 psidii Fruit tree diseases Fungi of South America Fungi of New Zealand