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'''''Armillaria luteobubalina''''', comúnmente conocido como el '''hongo de la miel de Australia''', es una especie de hongo de la familia ''[[Physalacriaceae]]''. Ampliamente distribuido en el sur de Australia, es el hongo responsable de la enfermedad conocida como «pudrición de la raíz por armillaria», causa principal de muerte de los [[eucalipto]]s y la [[Extinción paulatina del bosque|acronecrosis]]. Es la más patógena y generalizada de las seis especies de ''[[Armillaria]]'' que se encuentran en Australia. El hongo también ha sido recogido en la Argentina y Chile. |
'''''Armillaria luteobubalina''''', comúnmente conocido como el '''hongo de la miel de Australia''', es una especie de hongo de la familia ''[[Physalacriaceae]]''. Ampliamente distribuido en el sur de Australia, es el hongo responsable de la enfermedad conocida como «pudrición de la raíz por armillaria», causa principal de muerte de los [[eucalipto]]s y la [[Extinción paulatina del bosque|acronecrosis]]. Es la más patógena y generalizada de las seis especies de ''[[Armillaria]]'' que se encuentran en [[Australia]]. El hongo también ha sido recogido en la [[Argentina]] y [[Chile]]. |
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==Descripción== |
==Descripción== |
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El cuerpo fructífero o [[basidiocarpo]] tiene color tostado crema, [[Píleo (micología)|copas]] que crecen hasta 10 cm de diámetro y [[Estipe (micología)|estipes]] que miden hasta 20 cm de largo por 15 [[mm]] de grosor. Los [[basidiocarpo]]s, que aparecen en la base de los árboles infectados y otras plantas leñosas en otoño (marzo-abril), son [[seta comestible|comestibles]] , pero requieren de cocción para eliminar el sabor amargo. El hongo se dispersa a través de las [[espora]]s producidas en las [[Láminas (micología)|láminas]] de la parte inferior de las copas, y también por [[propagación vegetativa|crecimiento vegetativo]] en las raíces de los árboles hospedantes. Un sistema de aireación que permite que se difunda de manera eficiente el oxígeno a través de [[micelio]]s rizomorfos (estructuras similares a raíces formadas por cordones de [[hifa]]s<ref>http://www.plantasyhongos.es/hongos/hongos_micelio.htm</ref>) facilita su capacidad de propagación vegetativa. |
El cuerpo fructífero o [[basidiocarpo]] tiene color tostado crema, [[Píleo (micología)|copas]] que crecen hasta 10 cm de diámetro y [[Estipe (micología)|estipes]] que miden hasta 20 cm de largo por 15 [[mm]] de grosor. Los [[basidiocarpo]]s, que aparecen en la base de los árboles infectados y otras plantas leñosas en otoño (marzo-abril), son [[seta comestible|comestibles]] , pero requieren de cocción para eliminar el sabor amargo. El hongo se dispersa a través de las [[espora]]s producidas en las [[Láminas (micología)|láminas]] de la parte inferior de las copas, y también por [[propagación vegetativa|crecimiento vegetativo]] en las raíces de los árboles hospedantes. Un sistema de aireación que permite que se difunda de manera eficiente el oxígeno a través de [[micelio]]s rizomorfos (estructuras similares a raíces formadas por cordones de [[hifa]]s<ref>http://www.plantasyhongos.es/hongos/hongos_micelio.htm</ref>) facilita su capacidad de propagación vegetativa. |
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''Armillaria luteobubalina'' fue descrita por primera vez en 1978, después de |
''Armillaria luteobubalina'' fue descrita por primera vez en 1978, después de ser descubierta años antes creciendo en una plantación de eucaliptos en el sureste de Australia. Se diferencia de otras especies australianas conocidas de ''Armillaria'' por su agresiva [[patogenicidad]]. Pueden pasar años sin que los árboles infectados empiecen a mostrar signos de enfermedad, lo que lleva a una subestimación del nivel de expansión de la enfermedad. Los estudios muestran que la propagación de la enfermedad en los bosques de eucaliptos está asociada con el abandono de tocones infectados después del desmonte (o eliminación de la vegetación nativa) de tierras en Australia para usos agrícolas y urbanos. Aunque se han propuesto varios métodos para controlar la propagación de la enfermedad, estos son en su mayoría económica o medioambientalmente inviables. Análisis [[Filogenia|filogenéticos]] han determinado que ''A. luteobubalina'' está estrechamente relacionado con ''[[Armillaria montagnei]]'' y que ambas especies, a su vez, están estrechamente relacionadas con la especie brasileña ''[[Armillaria paulensis]]''. La distribución de ''A. luteobubalina'' sugiere que es una especie antigua que se originó antes de la separación del supercontinente precursor [[Gondwana]]. |
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== Historia y filogenia == |
== Historia y filogenia == |
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La Armillaria luteobubalina fue descrita por primera vez en 1978 por los micólogos Roy Watling y Glen Kile , que estudiaron sus efectos en las plantaciones de [[Eucalyptus regnans]] cerca de [[Traralgon]], Victoria. La plantación, establecida en 1963, consistía en gran parte de los árboles con una altura media de unos 25 metros. Un racimo de árboles muertos y moribundos descubiertos en 1973 sugirió un ataque por parte de un virulento patógeno primario, es decir, capaz de infectar a un huésped antes de la invasión de otros patógenos, uno secundario. Esta conclusión era incompatible con el comportamiento de las especies patógenas de Armillaria conocidas en Australia en ese momento, [[Armillaria mellea]] y A. elegans. El estudio adicional durante los próximos años demostró que el hongo se extendió por el crecimiento subterráneo de micelios en sistemas de raíces, la expansión hacia el exterior desde el muñón infectado inicial en un promedio de 2,5 m por año. La mayoría de los registros australianos de infecciones Armillaria se refiere a A. mellea, basado en la presencia de rizomorfos negros.<ref name="Podger1978" /> |
La Armillaria luteobubalina fue descrita por primera vez en 1978 por los micólogos Roy Watling y Glen Kile , que estudiaron sus efectos en las plantaciones de [[Eucalyptus regnans]] cerca de [[Traralgon]], Victoria. La plantación, establecida en 1963, consistía en gran parte de los árboles con una altura media de unos 25 metros. Un racimo de árboles muertos y moribundos descubiertos en 1973 sugirió un ataque por parte de un virulento patógeno primario, es decir, capaz de infectar a un huésped antes de la invasión de otros patógenos, uno secundario. Esta conclusión era incompatible con el comportamiento de las especies patógenas de Armillaria conocidas en Australia en ese momento, [[Armillaria mellea]] y A. elegans. El estudio adicional durante los próximos años demostró que el hongo se extendió por el crecimiento subterráneo de micelios en sistemas de raíces, la expansión hacia el exterior desde el muñón infectado inicial en un promedio de 2,5 m por año. La mayoría de los registros australianos de infecciones Armillaria se refiere a A. mellea, basado en la presencia de rizomorfos negros.<ref name="Podger1978" /> |
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== History and phylogeny == |
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''Armillaria luteobubalina'' was first described in 1978 by mycologists [[Roy Watling]] and [[Glen Kile]], who studied its effects on a fast-growing [[plantation]] of ''[[Eucalyptus regnans]]'' near [[Traralgon, Victoria]]. The plantation, established in 1963, consisted largely of trees with a mean height of about {{convert|25|m|ft|-1|abbr=on}}. A cluster of dead and dying trees discovered in 1973 suggested attack by a virulent primary [[plant pathology|pathogen]], that is, one capable of infecting a host before invasion by other, secondary pathogens. This finding was inconsistent with the pathogenic behaviour of the known ''Armillaria'' species in Australia at the time, ''[[Armillaria mellea|A. mellea]]'' and ''A. elegans''. Further study over the next few years showed that the fungus spread by the growth of underground [[mycelium|mycelia]] in root systems, expanding outward from the initial infected stump at an average of {{convert|2.5|m|ft|1|abbr=on}} per year. Most Australian records of ''Armillaria'' infections referred to ''A. mellea'', based on the presence of black [[rhizomorph]]s.<ref name="Podger1978" /> For over one hundred years, ''A. mellea'' was thought to be a [[pleomorphism (microbiology)|pleiomorphic]] (occurring in various distinct forms) species with a widespread distribution and host range, and variable pathogenicity.<ref name="Pegler2000" /> which led to great confusion among taxonomists and plant pathologists alike.<ref name="Watling1982"/> In 1973, [[Veikko Hintikka]] reported a technique to distinguish between ''Armillaria'' species by growing them together as single [[spore]] isolates on [[petri dish]]es and observing changes in the [[morphology (biology)|morphology]] of the cultures.<ref name="Hintikka1973" /> Using similar techniques, mycologists eventually determined that the ''Armillaria mellea'' [[species complex]] in Europe and North America in fact consisted of five and ten distinct "biological species", respectively.<ref name="Korhonen1978" /><ref name="Anderson1979" /> |
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Watling and Kile compared the macroscopic and microscopic characters of the pathogenic ''Armillaria'' with ''A. polymyces'' (now known as ''[[Armillaria obscura|A. obscura]]''), ''A. mellea'', ''[[Armillaria limonea|A. limonea]]'' and ''[[Armillaria novae-zelandiae|A. novae-zelandiae]]'' and found sufficient differences between them to warrant designating the species as new. Its [[botanical name|specific epithet]] is derived from the [[Latin]] ''lutea'' "yellow", and was chosen to highlight an important distinguishing characteristic: the strong yellow colour of the cap and lack of reddish or brown tones in the stem typical of other resident ''Armillaria''.<ref name="Podger1978" /> |
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A [[phylogenetic]] study of South American ''Armillaria'' species concluded that ''A. luteobubalina'' is in a lineage that includes ''[[Armillaria montagnei|A. montagnei]]'', and these are [[cladistics|sister]] to a lineage containing ''[[Armillaria paulensis|A. paulensis]]'',<ref name="Pildain2009" /> a species known from a single specimen collected in [[São Paulo]], Brazil.<ref name="Lima2008" /> Although they are very similar, specimens of ''A. luteobubalina'' have smaller spores than Argentinian specimens of ''A. montagnei'', and their distinctness is well-supported with phylogenetic analysis.<ref name="Pildain2010" /> Based on analysis of [[pectic]] enzymes, ''A. luteobubalina'' is closely related to ''[[Armillaria limonea|A. limonea]]'', a species found in New Zealand;<ref name="Coetzee2009" /> this result corroborates phylogenetic analyses reported in 2003<ref name="Coetzee2003" /> and 2006.<ref name="Maphosa2006" /> [[Molecular phylogenetics|Molecular]] analysis of 27 collections of ''A. luteobubalina'' from southwest Western Australia and one from Traralgon revealed four distinct polymorphic groups. The genetic variety suggests it is native to Australia.<ref name="Dunne2002" /><!-- cites previous two sentences --> |
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== Description == |
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[[File:Armillaria luteobubalina 67511.jpg|thumb|Mushroom caps are covered with tiny scales and have an umbo.]] |
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Up to {{convert|10|cm|abbr=on|0}} in diameter, the [[pileus (mycology)|cap]] is convex to flattened in shape with a central [[umbo (mycology)|umbo]] (a rounded elevation) and is various shades of cream, yellow and tan. The cap surface is covered with darker scales and feels rough to the touch. The cap edge, or margin, is rolled inward in young specimens. The crowded [[lamella (mycology)|gills]] are [[wikt:sinuate|sinuate]] and white to cream in colour initially, brownish-cream or pinkish brown in maturity, and sometimes with yellow or rust-coloured marks close to the margins.<ref name="fungimap" " /> The [[stipe (mycology)|stem]] is central (that is, it joins the cap in the centre) and is up to {{convert|20|cm|abbr=on|0}} long by {{convert|1.5|cm|abbr=on|0}} thick. It is slightly thicker at its base than its apex, sometimes almost bulb-like. The stem surface is streaked with fibrils that run up and down its length. It has a floppy yellow wool-like [[annulus (mycology)|ring]] which may develop irregular, jagged edges with time. The [[trama (mycology)|flesh]] is white, and in the stem has a woolly or stringy consistency.<ref name="Podger1978" /> Although it has a hot-bitter taste, ''Armillaria luteobubalina'' is [[edible mushroom|edible]], and cooking removes the bitterness.<ref name="Bougher1998" /> |
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=== Microscopic characteristics === |
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The [[spore print]] is white when fresh, but becomes more cream-coloured when dry. The smooth [[spore]]s are oval to ellipsoid, [[hyaline]] (translucent), non-[[amyloid]] (meaning they do not absorb [[iodine]] from [[Melzer's reagent]]), and typically measure 6.5–7.5 by 4.5–5.5 [[micrometre|μm]]. The [[basidia]] (spore-bearing cells) are thin-walled, hyaline, and lack [[clamp connection]]s at their bases. They are usually four-spored but occasionally two-spored, with [[sterigmata]] (projections that attach to the spores) up to 4 μm long. The cheilocystidia ([[cystidia]] that occur on the edge of a gill) are mostly club-shaped, thin-walled, hyaline, and measure 15–30 by 6–10 μm.<ref name="Podger1978" /> |
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=== Similar species === |
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Five other ''Armillaria'' species are found in Australia.<ref name="Robinson2003" /> Within the range of ''A. luteobubalina'', ''[[Armillaria hinnulea|A. hinnulea]]'' is restricted to [[gully]] habitats. ''[[Armillaria fumosa|A. fumosa]]'' is a rarer species found only in poorly drained or seasonally wet locations.<ref name="Kile&Watling1983" " /> ''A. luteobubalina'' and ''A. montagnei'' share cap features and a similar unpleasant flavour, but the latter species has an olive-tinged cap, larger spores (9.5–11 by 5.5–7 µm compared to 6.5–7.5 by 4.5–5.5 µm) and a more conspicuous annulus than those found in ''A. luteobubalina''.<ref name="Pildain2010" /> The morphology of the vegetative structures of ''[[Armillaria limonea|A. limonea]]'' is distinctly different than ''A. luteobubalina'', and can be used to distinguish the two species.<ref name="Shaw1981" /> ''A. novae-zelandiae'' has a sticky more flattened cap and stem below the ring and is found in wet forests, and ''A. pallidula'' is a species with cream gills maturing to pale pink found in tropical Australia arising from dead tree stumps or the roots of dead or living trees.<ref name="Fuhrer2005" /> ''A. luteobubalina'' is the only ''Armillaria'' species which occurs in Western Australia.<ref name="Bougher1998" /> Distinguishing Australian species is economically important, because ''A. luteobubalina'' is more pathogenic than the other members of the genus. A molecular diagnostic test, developed in 2002, can accurately identify each species using [[DNA]] extracted from its mycelia. Before this, species identification was limited to times when fruit bodies were in season. This technology also revealed a variation in the molecular material of ''A. luteobubalina'' that suggested sexual reproduction.<ref name="SmithWhite2002" /> |
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== Habitat and distribution == |
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[[File:Karri forest.jpg|thumb|left|''A. luteobubalina'' grows in [[Karri forest]]s, like this one in [[Pemberton, Western Australia]].]] |
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''Armillaria luteobubalina'' has been recorded in southeastern Australia, from the southeastern corner of [[Queensland]] through eastern [[New South Wales]] and across [[Victoria (Australia)|Victoria]] into southeastern [[South Australia]]. It also occurs in [[Tasmania]] and southwestern [[Western Australia]].<ref name="fungimap" " /> Those of the [[karri forest]]s (consisting largely of the species ''[[Eucalyptus diversicolor|E. diversicolor]]'') of the southwest have paler and yellower caps than those in the [[jarrah forest]]s (which contain predominantly ''[[Eucalyptus marginata]]'') further north.<ref name="Bougher1998" /> The fruit bodies arise on wood, especially on stumps or around the base of trees, and often in huge numbers. They usually appear between April and July, although most production occurs in the second half of May.<ref name="Kile1981b" /> Abundant in woodlands, it can invade gardens and orchards, where it can attack many woody plants.<ref name="Fuhrer2005" /><ref name="Smith1981" /> The honey fungus infected and killed many plants near tuart trees (''[[Eucalyptus gomphocephala]]'') which had been cut down near [[Kings Park, Western Australia|Kings Park]] in suburban Perth.<ref name="Bougher1998" /> ''Armillaria luteobubalina'' is commonly found in [[eucalyptus]] forests in Australia,<ref name="Podger1978" /><ref name="Kile1981b" /> and is thought to be the most [[pathogenic]] and most widespread ''Armillaria'' species in the major western Australian forest types.<ref name="Kile et al. 1983" " /><ref name="Pearce1989" /> The mushroom has also been reported from southern South America, in Argentina and Chile.<ref name="Pildain2009" /> A 2003 study of the molecular phylogenetics and pattern of its distribution in South America and Australia indicate that ''A. luteobubalina'' is an ancient species, originating before the separation of the precursor supercontinent [[Gondwana]]. Genetic differences between isolates in the South American and Australian populations indicate a long period of geographical separation, and the authors suggest that they "later might be regarded as independent taxa".<ref name="Coetzee2003" /> |
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== Root rot == |
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[[File:Wielangta Unidentified Fungus 5164.jpg|thumb|Young fruit body found in [[Wielangta forest]] in southeastern [[Tasmania]], Australia]] |
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=== Appearance of infected trees === |
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Trees that are infected by ''A. luteobubalina'' show characteristic symptoms both above and below ground. Above the ground, the base of the tree develops inverted V-shaped lesions, and the infected wood undergoes [[white rot]], a fungal wood decay process where the [[cellulose]] and [[lignin]] of the [[Wood#Heartwood_and_sapwood|sapwood]] are both broken down, leaving the wood stringy. The [[bark]] of the stem dies and becomes discoloured up to {{convert|3|m|ft|0|abbr=on}} above the ground. Clusters of fruit bodies appear at the base of the tree in autumn. [[Crown (botany)|Crowns]] may show gradual deterioration, or tree death may occur suddenly. Below the ground, characteristic symptoms of infections include rotting the ends of tree roots, white-rotted sapwood, and the presence of fan-shaped areas of white mycelium below dead or infected bark.<ref name="Robinson2003" /> |
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=== Occurrence === |
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In selectively logged eucalypt forests in the central highlands of Victoria, it has been estimated that about 3–5% of the forest area is "moderately to severely affected" by ''Armillaria'' root rot caused by ''A. luteobubalina''.<ref name="Kile1983b" /> A review of eucalypt plantations planted in New South Wales from 1994 to 2005 found that infection by ''A. luteobubalina'' was rare, and only accounted for 1% of mortality in total. In this instance, the cases had been restricted to ''[[Eucalyptus nitens]]'' on the [[Dorrigo Plateau]].<ref name="Carnegie2007" /> Unlike other ''Armillaria'' species found in Australia's native forests, which require a host tree to become weakened by prior infection by a different species, ''A. luteobubalina'' is a primary pathogen, and can infect healthy trees.<ref name="Kile1983b" /><ref name="Kile1981" /> Tree roots may be infected for years before showing above-ground symptoms, making it difficult to accurately assess the true extent of disease in a forest stand.<ref name="Edgar1976" /> Surveys are usually conducted in autumn, to coincide with the appearance of fruit bodies; infection is assessed by the presence of basal scars on the trees, and the appearance of fruit bodies. Several factors, however—such as cost, variable on-site conditions, and non-symptomatic diseased trees—make it difficult for such surveys to reliably detect all infections. One study showed that above-ground examinations detected only 50% of the trees actually infected, leading to underestimation of the incidence of true infection by 20–40%. The study used more intensive surveying methods to determine that 25–30 year old karri regrowth forests in western Australia showed an average of 40–45% incidence of infection.<ref name="Robinson2003" /> |
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=== Disease spread === |
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[[File:Armillaria luteobubalina 151150.jpg|thumb|left|A dense cluster of fruit bodies growing on a rotting stump]] |
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Several studies have shown that the spread of ''Armillaria'' root rot in eucalypt forests is associated with infected [[tree stump|stumps]] that remain after an area has been [[logging|logged]].<ref name="Edgar1976" /><ref name="Pearce1986" /><ref name="Kellas1987" /> ''Armillaria luteobubalina'' can persist on these stumps, using them as a source of food for up to 25 or more years.<ref name="Kile1981" /> In one case reported in [[Ovens, Victoria]], the disease was spread to [[blueberry]] plants (''Vaccinium'' species) via buried fragments of infected ''Eucalyptus'' that remained following preparation of the previously forested site for planting.<ref name="Falk1995" /> In individual forest stands,<ref>A forest stand may be defined as: "A group of trees occupying a given area and sufficiently uniform in species composition, age, structure, site quality, and condition so as to be distinguishable from the forest on adjoining areas." From {{cite web |url=http://www.dnr.state.mn.us/forestry/subsection/glossary.html |title=Glossary – SFMRP – Division of Forestry: Minnesota DNR |publisher=Minnesota Department of Natural Resources |accessdate=2009-12-23}}</ref> fungal infection is usually found in discrete disease patches separated by stands of healthy trees—a discontinuous distribution.<ref name="Kile1981" /> Large-scale [[aerial photography]] can be used to identify regions of forest infected by the species.<ref name="Myers1983" /> The species also causes damage to trees and bushes in coastal [[dune]] woodlands, [[shrubland]], and [[heath (habitat)|heath]] communities.<ref name="Shearer1997" /> It can be found on a wide range of hosts, but is most commonly associated with (in order of decreasing frequency) jarrah (''[[Eucalyptus marginata]]''), bull banksia (''[[Banksia grandis]]''), marri (''[[Eucalyptus calophylla|E. calophylla]]''), ''[[Lasiopetalum floribundum]]'', and ''[[Acacia saligna]]''.<ref name="Shearer1988" /> It has also infected scattered populations of wandoo (''[[Eucalyptus wandoo|E. wandoo]]'').<ref name="Shearer1997b" /> The fungus has also been reported to infect ''[[Nothofagus]]'' species in Argentina, and ''[[Pinus radiata]]'' in Chile.<ref name="Pildain2009" /> |
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''Armillaria luteobubalina'' uses "an elaborate, sophisticated aeration system" that enables it to efficiently deliver oxygen into the [[rhizomorph]]s, helping it thrive in low-oxygen environments.<ref name="Kile1983" /><ref name="Pareek2006" /> When grown in [[microbiological culture|culture]], the [[mycelium]] develops into a continuous region of tissue with a perforated crust. This tissue is [[hydrophobic]] and resistant to becoming waterlogged. Rhizomorphs develop beneath clusters of so-called "air-pores" near the perforations. These gas spaces connect the atmosphere with the central canal of the rhizomorph, to facilitate [[diffusion]] of oxygen and satiate the organism's high oxygen requirement during growth. This aeration system is thought to be an important factor in the organism's pathogenicity, allowing it to grow on wet or waterlogged root surfaces and send hyphae or rhizomorphs into live roots or cut stumps, where conditions may be [[hypoxia (environmental)|hypoxic]].<ref name="Pareek2006" /> The rhizomorphs have a dichotomous branching pattern, so that they split or bifurcate at various intervals. Experiments and field observations have shown that this allows the fungus to be a more aggressive and virulent pathogen than ''Armillaria'' species whose rhizomorphs branch [[monopodial]]ly (where lateral branches grow from a main stem).<ref name="Morrison2004" /> Although the structure of ''A. luteobubalina'' rhizomorphs is specialised for spread in potentially anaerobic conditions, the soil mycelium is adaptive and can amplify the absorptive surface of peripheral hyphae in response to the presence of nutrient-rich soil.<ref name="Pareek2001" /> |
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=== Control === |
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Methods for controlling the spread of ''Armillaria'' root rot include physical removal of infected trees, stumps and large roots;<ref name="Robinson2001b" /> fumigation of soil around infected hosts; and injection of fumigants directly into infected hosts.<ref name="Robinson2001" /> These methods are often not practical due to high cost, introduction of toxic chemicals that affect other organisms, or health and safety issues for the operator.<ref name="Robinson2001" /><ref name="Schütt1985" " /> [[Biological pest control|Biological control]] is another method that has been investigated to control root rot caused by ''A. luteobubalina''. In one study, thinning stumps of ''Eucalyptus diversicolor'' were simultaneously inoculated with ''A. luteobubalina'' and one of the [[saprobic]] [[wood-decay fungus|wood decay fungi]] ''[[Coriolus versicolor]]'', ''[[Stereum hirsutum]]'' and ''[[Xylaria hypoxylon]]''; all three fungi significantly reduced infection by ''A. luteobubalinea''.<ref name="Pearce1990" /> These results were echoed in another study of stumps in karri regrowth forests, where it was shown that the presence of other wood decay fungi suppressed the growth of ''A. luteobubalina'' on the stump base.<ref name="Pearce1989" /> |
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== See also == |
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{{Portal|Fungi}} |
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* [[List of Armillaria species|List of ''Armillaria'' species]] |
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{{Clear}} |
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== References == |
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{{Reflist|colwidth=30em|refs= |
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<ref name="Anderson1979">{{cite journal |author=Anderson JB, Ullrich RC. |year=1979 |title=Biological species of ''Armillaria'' in North America |journal=Mycologia |volume=71 |issue=2 |pages=401–14 |jstor=3759160 |doi=10.2307/3759160}}</ref> |
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<ref name="Bougher1998">{{cite book |title=Fungi of Southern Australia|author=Bougher NL, Syme K. |year=1998|pages=192–93|publisher=[[University of Western Australia]] Press |location=Nedlands, WA |isbn=1-875560-80-7}}</ref> |
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<ref name="Carnegie2007">{{cite journal |author=Carnegie AJ.|year=2007|title=Forest health condition in New South Wales, Australia, 1996–2005. II. Fungal damage recorded in eucalypt plantations during forest health surveys and their management |journal=Australasian Plant Pathology |volume=36 |issue=3 |pages=225–39 |doi=10.1071/AP07021}}</ref> |
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<ref name="Coetzee2003">{{cite journal |author=Coetzee MPA, Wingfield BD, Bloomer P, Ridley GS, Wingfield MJ. |year=2003 |title=Molecular identification and phylogeny of ''Armillaria'' isolates from South America and Indo-Malaysia |journal=Mycologia |volume=95 |issue=2 |pages=285–93|doi=10.2307/3762039 |pmid=21156614}}</ref> |
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<ref name="Coetzee2009">{{cite journal |author=Coetzee MPA, Maphosa L, Mwenje E, Wingfield MJ, Wingfield BD. |year=2009 |title=Characterisation of ''Armillaria'' species based on pectic isozyme analyses |journal=Fungal Diversity |volume=36 |pages=9–16 |url=http://www.fungaldiversity.org/fdp/sfdp/FD36-2.pdf |format=PDF}}</ref> |
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<ref name="Dunne2002">{{cite journal |author=Dunne CP, Glen M, Tommerup IC, Shearer L, Jardy GE St J. |year=2002 |title=Sequence variation in the rDNA ITS of Australian ''Armillaria'' species and intra-specific variation in ''A. luteobubalina'' |journal=Australasian Plant Pathology |volume=31 |issue=3 |pages=241–51 |doi=10.1071/AP02015}}</ref> |
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<ref name="Edgar1976">{{cite journal |author=Edgar JG, Kile GA, Almond CA. |year=1976 |title=Tree decline and mortality in selectively logged eucalypt forests in central Victoria |journal=Australian Forestry |volume=39 |issue=4 |pages=288–303 |doi=10.1080/00049158.1976.10675656}}</ref> |
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<ref name="Falk1995">{{cite journal |author=Falk SP, Parbery DG. |year=1995 |title=''Armillaria luteobubalina'' population-structure in horticultural plantings in Victoria, Australia |journal=Mycological Research |volume=99 |issue=2 |pages=216–20 |doi=10.1016/S0953-7562(09)80888-5}}</ref> |
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<ref name="fungimap">{{cite book |title=Fungi Down Under: The Fungimap Guide to Australian Fungi |author=Grey P. |year=2005 |publisher=Royal Botanic Gardens |location=Melbourne |isbn=0-646-44674-6 |page=23}}</ref> |
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<ref name="Hintikka1973">{{cite journal |author=Hintikka V. |year=1973 |title=A note on the polarity of ''Armillaria mellea'' |journal=Karstenia |volume=13 |pages=32–39}}</ref> |
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<ref name="Kellas1987">{{cite journal |author=Kellas JD, Kile GA, Jerrett RG, Morgan BJT. |year=1987 |title=The occurrence and effects of ''Armillaria luteobubalina'' following partial cutting in mixed eucalypt stands in the Wombat Forest, Victoria |journal=Australian Forest Research |volume=17 |issue=3 |pages=263–76}}</ref> |
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<ref name="Kile1981">{{cite journal |author=Kile GA. |year=1981 |title=''Armillaria luteobubalina'' – a primary cause of decline and death of trees in mixed species eucalypt forests in central Victoria |journal=Australian Forest Research |volume=11 |issue=1 |pages=63–77 |issn=0004-914X }}</ref> |
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<ref name="Kile1981b">{{cite journal |author=Kile GA, Watling R. |year=1981 |title=An expanded concept of ''Armillaria luteobubalina'' |journal=Transactions of the British Mycological Society |volume=77 |issue=1 |pages= 75–83 |doi=10.1016/S0007-1536(81)80181-7}}</ref> |
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<ref name="Kile1983">{{cite journal |author=Kile GA. |year=1983 |title=Identification of genotypes and the clonal development of ''Armillaria luteobubalina'' Watling and Kile in eucalypt forests |journal=Australian Journal of Botany |volume=31 |issue=6 |pages=657–71 |doi=10.1071/BT9830657}}</ref> |
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<ref name="Kile1983b">{{cite journal |author=Kile GA. |year=1983 |title= Armillaria root rot in eucalypt forests aggravated endemic disease |journal=Pacific Science |volume=37 |issue=4 |pages=459–64|issn=0030-8870}}</ref> |
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<ref name="Kile et al. 1983">{{cite journal |author=Kile GA, Watling R, Malajczuk N, Shearer BL. |year=1983 |title=Occurrence of ''Armillaria luteobubalina'' Watling and Kile in Western Australia |journal=Australasian Plant Pathology |volume=12 |issue=2 |pages=18–20 |doi=10.1071/APP9830018}}</ref> |
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<ref name="Kile&Watling1983">{{cite journal |author=Kile GA, Watling R. |year=1983 |title=''Armillaria'' species from south-eastern Australia |journal=Transactions of the British Mycological Society |volume=81 |issue=1 |pages=129–40 |issn=0007-1536 |doi=10.1016/S0007-1536(83)80212-5}}</ref> |
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<ref name="Korhonen1978">{{cite journal |author=Korhonen K. |year=1978 |title=Interfertility and clonal size in the ''Armillaria mellea'' complex |journal=Karstenia |volume=18 |pages=31–42}}</ref> |
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<ref name="Myers1983">{{cite journal |author=Myers BJ, Kile GA, Runnalls R. |year=1983 |title=Evaluation of large scale aerial photography for identifying areas of Armillaria root rot in mixed species eucalypt forest |journal=Australian Forestry |volume=46 |issue=1 |pages=39–42 |issn=0004-9158 |doi=10.1080/00049158.1983.10674375}}</ref> |
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<ref name="Lima2008">{{cite journal |author=Lima MLA, Asai T, Capelari M.|year=2008 |title=''Armillaria paulensis'': a new South American species |journal=Mycological Research |volume=112 |issue=9 |pages=1122–28 |pmid=18692378 |doi=10.1016/j.mycres.2008.03.006}}</ref> |
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<ref name="Maphosa2006">{{cite journal |author=Maphosa L, Wingfield BD, Coetzee MPA, Mwenje E, Wingfield MJ. |year=2006 |title= Phylogenetic relationships among ''Armillaria'' species inferred from partial elongation factor 1-alpha DNA sequence data |journal=Australasian Plant Pathology |volume=35 |issue=5 |pages=513–20 |doi=10.1071/AP06056}}</ref> |
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<ref name="Morrison2004">{{cite journal |author=Morrison DJ. |year=2004 |title=Rhizomorph growth habit, saprophytic ability and virulence of 15 ''Armillaria'' species |journal=Forest Pathology |volume=34 |issue=1 |pages=15–26 |doi=10.1046/j.1439-0329.2003.00345.x}}</ref> |
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<ref name="Pareek2001">{{cite journal |author=Pareek M, Cole L, Ashford AE. |year=2001 |title=Variations in structure of aerial and submerged rhizomorphs of ''Armillaria luteobubalina'' indicate that they may be organs of absorption |journal=Mycological Research |volume=105 |issue=11 |pages=1377–87 |doi=10.1017/S0953756201004622}}</ref> |
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<ref name="Pareek2006">{{cite journal |author=Pareek M, Allaway WG, Ashford AE. |year= 2006|title= ''Armillaria luteobubalina'' mycelium develops air pores that conduct oxygen to rhizomorph clusters |journal= Mycological Research|volume=110 |issue=1 |pages=38–51 |doi=10.1016/j.mycres.2005.09.006 |pmid=16376531}}</ref> |
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<ref name="Pearce1986">{{cite journal |author=Pearce MH, Malajczuk N, Kile GA. |year=1986 |title=The occurrence and effects of ''Armillaria luteobubalina'' in the karri (''Eucalyptus diversicolor'' F. Muell.) forests of Western Australia |journal=Australian Forest Research |volume=16 |issue=3 |pages=243–59}}</ref> |
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<ref name="Pearce1989">{{cite journal |author=Pearce MH, Malanjczuk N. |year=1989 |title=Stump colonization by ''Armillaria luteobubalina'' and other wood decay fungi in an age series of cut-over stumps in Karri (''Eucalyptus diversicolor'') regrowth forests in south-western Australia |journal=New Phytologist |volume=115 |issue=1 |pages=129–38 |jstor=2557060 |doi=10.1111/j.1469-8137.1990.tb00930.x}}</ref> |
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<ref name="Pearce1990">{{cite journal |author=Pearce MH, Malajczuk N. |year=1990 |title= Inoculation of ''Eucalyptus diversicolor'' thinning stumps with wood decay fungi for control of ''Armillaria luteobubalina''|journal=Mycological Research |volume=94 |issue=1 |pages=32–37 |issn=0953-7562 |doi=10.1016/S0953-7562(09)81261-6}}</ref> |
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<ref name="Pegler2000">{{cite book|chapter=Taxonomy, nomenclature and description of ''Armillaria'' |title=''Armillaria'' Root Rot: Biology and Control of Honey Fungus |author=Pegler DN. |editor-last=Fox RTV |year=2000|publisher=Intercept |location=Andover |isbn=1-898298-64-5 |pages=81–93}}</ref> |
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<ref name="Pildain2009">{{cite journal |author=Pildain MB, Coetzee MPA, Rajchenberg RH, Petersen RH, Wingfield MJ, Wingfield BD. |year=2009 |title=Molecular phylogeny of ''Armillaria'' from the Patagonian Andes |journal=Mycological Progress |volume=8 |issue=3 |pages=181–94 |doi=10.1007/s11557-009-0590-8}}</ref> |
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<ref name="Pildain2010">{{cite journal |author=Pildain MB, Coetzee MPA, Wingfield BD, Wingfield MJ. |year=2010 |title=Taxonomy of ''Armillaria'' in the Patagonian forests of Argentina |journal=Mycologia |volume=102 |issue=2 |pages=392–403 |url= |doi=10.3852/09-105 |pmid=20361506}}</ref> |
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<ref name="Podger1978">{{cite journal |author=Podger FD, Kile GA, Watling R, Fryer J. |year=1978 |title=Spread and effects of ''Armillaria luteobubalina'' sp. nov. in an Australian ''Eucalyptus regnans'' plantation |journal=Transactions of the British Mycological Society |volume=71 |issue=1 |pages=77–87 |doi=10.1016/S0007-1536(78)80009-6}}</ref> |
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<ref name="Robinson2001">{{cite journal |author=Robinson RM, Smith RH. |year=2001 |title=Fumigation of regrowth karri stumps with metham-sodium to control ''Armillaria luteobubalina'' |journal=Australian Forestry |volume=64 |issue=4 |pages=209–15 |url=http://www.forestry.org.au/pdf/pdf-members/afj/AFJ%202001%20v64/AFJ%20Dec%202001%2064(4)/Robinson.PDF |format=PDF |doi=10.1080/00049158.2001.10676190}}</ref> |
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<ref name="Robinson2001b">{{cite journal |author=Robinson RM. |year=2003 |title=Short-term impact of thinning and fertilizer application on ''Armillaria'' root disease in regrowth karri (''Eucalyptus diversicolor'' F. Muell.) in Western Australia |journal=Forest Ecology and Management |volume=176 |issue=1–3 |pages=417–26 |doi=10.1016/S0378-1127(02)00231-1}}</ref> |
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<ref name="Robinson2003">{{cite journal |author=Robinson RM, Williams MR, Smith RH. |year=2003 |title=Incidence of Armillaria root disease in Karri regrowth forest is underestimated by surveys of aboveground symptoms |journal=Australian Forestry |volume=66 |pages=273–78 |url=http://svc246.wic013v.server-web.com/pdf/pdf-members/afj/AFJ%202003%20v66/AFJ%20Dec%202003%2066-4/04%20dec%20printproof%20robinson.pdf |format=PDF |doi=10.1080/00049158.2003.10674921 |issue=4}}</ref> |
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<ref name="Schütt1985">{{cite journal |author= Schütt P.|year=1985 |title=Control of root and butt rots: limits and prospects |journal=European Journal of Forest Pathology |volume=15 |issue=5–6 |pages=357–63 |doi=10.1111/j.1439-0329.1985.tb01112.x}}</ref> |
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<ref name="Shaw1981">{{cite journal |author=Shaw CG, MacKenzie M, Toes EHA, Hood IA. |year=1981 |title=Cultural characteristics and pathogenicity to ''Pinus radiata'' of ''Armillaria novae-zelandiae'' and ''A. limonea'' |journal=New Zealand Journal of Forestry Science |volume=11 |pages=65–70}}</ref> |
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<ref name="Shearer1988">{{cite journal |author=Shearer BL, Tippett JT. |year=1988 |title=Distribution and impact of ''Armillaria luteobubalina'' in the ''Eucalyptus marginata'' forest of south-western Australia |journal=Australian Journal of Botany |volume=36 |issue=4 |pages=433–45 |doi=10.1071/BT9880433}}</ref> |
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<ref name="Shearer1997">{{cite journal |author=Shearer BL, Crane CE, Fairman RG, Grant MJ. |year=1997 |title=Occurrence of ''Armillaria luteobubalina'' and pathogen-mediated changes in coastal dune vegetation of south-western Australia |journal=Australian Journal of Botany |volume=45 |issue=5 |pages=905–17 |doi=10.1071/BT96084}}</ref> |
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<ref name="Shearer1997b">{{cite journal |author=Shearer BL, Byrne A, Dillon M, Buehrig R. |year=1997 |title=Occurrence of ''Armillaria luteobubalina'' and its impact on community diversity and structure in ''Eucalyptus wandoo'' woodland of southern Western Australia |journal=Australian Journal of Botany |volume=45 |issue=1 |pages=151–65 |doi=10.1071/BT95083 }}</ref> |
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<ref name="Smith1981">{{cite journal |author=Smith L, Kile GA. |year=1981 |title=Distribution and hosts of Armillaria root rot in Melbourne suburban gardens |journal=Australasian Plant Pathology |volume=10 |issue=3 |pages=41–42|issn=0815-3191 |doi=10.1071/APP9810041}}</ref> |
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<ref name="SmithWhite2002">{{cite journal |author=Smith-White JL, Summerell BA, Gunn LV, Rinzin C, Porter C, Burgess LW. |year=2002 |title=Molecular detection and differentiation of Australian ''Armillaria'' species |journal=Australasian Plant Pathology |volume=31 |issue=1 |pages=75–79 |doi=10.1071/AP01061}}</ref> |
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<ref name="Watling1982">{{cite journal|author=Watling R, Kile GA, Gregory NM. |title=The genus ''Armillaria'' – nomenclature, typification, the identity of ''Armillaria mellea'' and species differentiation |journal=Transactions of the British Mycological Society |year=1982 |volume=78 |issue=2 |pages=271–85 |doi=10.1016/S0007-1536(82)80011-9}}</ref> |
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}} |
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== External links == |
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* {{IndexFungorum|308928}} |
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{{featured article}} |
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{{DEFAULTSORT:Armillaria luteobubalina}} |
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[[Category:Armillaria|luteobubalina]] |
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[[Category:Edible fungi]] |
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[[Category:Fungal tree pathogens and diseases]] |
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[[Category:Fungi described in 1978]] |
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[[Category:Fungi of Australia]] |
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[[Category:Fungi of South America]] |
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== Referencias == |
== Referencias == |
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Revisión del 10:57 2 dic 2014
| Armillaria luteobubalina | ||
|---|---|---|
 | ||
| Taxonomía | ||
| Reino: | Fungi | |
| División: | Basidiomycota | |
| Clase: | Agaricomycetes | |
| Subclase: | Agaricomycetidae | |
| Orden: | Agaricales | |
| Familia: | Physalacriaceae | |
| Género: | Armillaria | |
| Especie: |
A. luteobubalina Watling & Kile (1978) | |
Armillaria luteobubalina, comúnmente conocido como el hongo de la miel de Australia, es una especie de hongo de la familia Physalacriaceae. Ampliamente distribuido en el sur de Australia, es el hongo responsable de la enfermedad conocida como «pudrición de la raíz por armillaria», causa principal de muerte de los eucaliptos y la acronecrosis. Es la más patógena y generalizada de las seis especies de Armillaria que se encuentran en Australia. El hongo también ha sido recogido en la Argentina y Chile.
Descripción
El cuerpo fructífero o basidiocarpo tiene color tostado crema, copas que crecen hasta 10 cm de diámetro y estipes que miden hasta 20 cm de largo por 15 mm de grosor. Los basidiocarpos, que aparecen en la base de los árboles infectados y otras plantas leñosas en otoño (marzo-abril), son comestibles , pero requieren de cocción para eliminar el sabor amargo. El hongo se dispersa a través de las esporas producidas en las láminas de la parte inferior de las copas, y también por crecimiento vegetativo en las raíces de los árboles hospedantes. Un sistema de aireación que permite que se difunda de manera eficiente el oxígeno a través de micelios rizomorfos (estructuras similares a raíces formadas por cordones de hifas[1]) facilita su capacidad de propagación vegetativa.
Armillaria luteobubalina fue descrita por primera vez en 1978, después de ser descubierta años antes creciendo en una plantación de eucaliptos en el sureste de Australia. Se diferencia de otras especies australianas conocidas de Armillaria por su agresiva patogenicidad. Pueden pasar años sin que los árboles infectados empiecen a mostrar signos de enfermedad, lo que lleva a una subestimación del nivel de expansión de la enfermedad. Los estudios muestran que la propagación de la enfermedad en los bosques de eucaliptos está asociada con el abandono de tocones infectados después del desmonte (o eliminación de la vegetación nativa) de tierras en Australia para usos agrícolas y urbanos. Aunque se han propuesto varios métodos para controlar la propagación de la enfermedad, estos son en su mayoría económica o medioambientalmente inviables. Análisis filogenéticos han determinado que A. luteobubalina está estrechamente relacionado con Armillaria montagnei y que ambas especies, a su vez, están estrechamente relacionadas con la especie brasileña Armillaria paulensis. La distribución de A. luteobubalina sugiere que es una especie antigua que se originó antes de la separación del supercontinente precursor Gondwana.
Historia y filogenia
La Armillaria luteobubalina fue descrita por primera vez en 1978 por los micólogos Roy Watling y Glen Kile , que estudiaron sus efectos en las plantaciones de Eucalyptus regnans cerca de Traralgon, Victoria. La plantación, establecida en 1963, consistía en gran parte de los árboles con una altura media de unos 25 metros. Un racimo de árboles muertos y moribundos descubiertos en 1973 sugirió un ataque por parte de un virulento patógeno primario, es decir, capaz de infectar a un huésped antes de la invasión de otros patógenos, uno secundario. Esta conclusión era incompatible con el comportamiento de las especies patógenas de Armillaria conocidas en Australia en ese momento, Armillaria mellea y A. elegans. El estudio adicional durante los próximos años demostró que el hongo se extendió por el crecimiento subterráneo de micelios en sistemas de raíces, la expansión hacia el exterior desde el muñón infectado inicial en un promedio de 2,5 m por año. La mayoría de los registros australianos de infecciones Armillaria se refiere a A. mellea, basado en la presencia de rizomorfos negros.[2]
Referencias
- ↑ http://www.plantasyhongos.es/hongos/hongos_micelio.htm
- ↑ Podger FD, Kile GA, Watling R, Fryer J. (1978). «Spread and effects of Armillaria luteobubalina sp. nov. in an Australian Eucalyptus regnans plantation». Transactions of the British Mycological Society 71 (1): 77-87. doi:10.1016/S0007-1536(78)80009-6.