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Pine-pine gall rust, also known as western gall rust, is a
fungal disease A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from th ...
of
pine A pine is any conifer tree or shrub in the genus ''Pinus'' () of the family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. The World Flora Online created by the Royal Botanic Gardens, Kew and Missouri Botanical Garden accep ...
trees. It is caused by ''Endocronartium harknessii'' (asexual name is ''Peridermium harknessii''), an
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 ...
, endocyclic,
rust fungus Rusts are plant diseases caused by pathogenic fungi of the order Pucciniales (previously known as Uredinales). An estimated 168 rust genera and approximately 7,000 species, more than half of which belong to the genus ''Puccinia'', are currently a ...
that grows in the
vascular cambium The vascular cambium is the main growth tissue in the stems and roots of many plants, specifically in dicots such as buttercups and oak trees, gymnosperms such as pine trees, as well as in certain other vascular plants. It produces secondary xyle ...
of the host. The disease is found on pine trees (''
Pinus A pine is any conifer tree or shrub in the genus ''Pinus'' () of the family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. The World Flora Online created by the Royal Botanic Gardens, Kew and Missouri Botanical Garden accep ...
'' spp.) with two or three needles, such as
ponderosa pine ''Pinus ponderosa'', commonly known as the ponderosa pine, bull pine, blackjack pine, western yellow-pine, or filipinus pine is a very large pine tree species of variable habitat native to mountainous regions of western North America. It is the ...
,
jack pine Jack pine (''Pinus banksiana'') is an eastern North American pine. Its native range in Canada is east of the Rocky Mountains from the Mackenzie River in the Northwest Territories to Cape Breton Island in Nova Scotia, and the north-central and ...
and
scots pine ''Pinus sylvestris'', the Scots pine (UK), Scotch pine (US) or Baltic pine, is a species of tree in the pine family Pinaceae that is native to Eurasia. It can readily be identified by its combination of fairly short, blue-green leaves and orang ...
.Peterson, Roger S. "Western Gall Rust on Hard Pines." U.S. Department of Agriculture Forest Service (1960): n. pag. Web. It is very similar to pine-oak gall rust, but its second host is another ''Pinus'' species. The fungal infection results in
gall Galls (from the Latin , 'oak-apple') or ''cecidia'' (from the Greek , anything gushing out) are a kind of swelling growth on the external tissues of plants, fungi, or animals. Plant galls are abnormal outgrowths of plant tissues, similar to be ...
formation on branches or trunks of infected hosts. Gall formation is typically not detrimental to old trees, but has been known to kill younger, less stable saplings. Galls can vary from small growths on branch extremities to grapefruit-sized galls on trunks.


Hosts and symptoms

The hosts of the
aecia An aecium (plural aecia) is a specialised reproductive structure found in some plant pathogenic rust fungi that produce aeciospores. Aecia may also be referred to as "cluster cups". The term aecidium (plural aecidia) is used interchangeably but i ...
l stage of the fungus includes two and three needled ''Pinus'' spp. The most important aecial hosts include jack pine ('' P. banksiana''), lodgepole pine ('' P. contorta''), western yellow pine ('' P. ponderosa''), and the European Scots pine ('' P. sylvestris''). A variety of other pines, such as ''
Pinus nigra ''Pinus nigra'', the Austrian pine or black pine, is a moderately variable species of pine, occurring across Southern Europe from the Iberian Peninsula to the eastern Mediterranean, on the Anatolian peninsula of Turkey, Corsica and Cyprus, as wel ...
'', '' P. mugo'', '' P. palustris'', '' P. banksiana'', '' P. muricata'', and '' P. radiata'' have also been reported as hosts to pine-pine gall rust (''E. harknessii''), but these pine species are considered less valuable. Because ''E. harknessii'' is an autoecious short-cycle rust lacking telial a host, there is no alternative hosts. The pathogen can infect actively growing shoots of any age very quickly without infecting an alternative host, making the disease cycle more destructive that typical rust species that switch between hosts. This also prevents control of the fungus by management of the alternative host species. Symptoms of pine-pine gall rust can be quite conspicuous and are useful for diagnostic identification. Pine-Pine Gall Rust: ''Endocronartium harknessii''
Cornell University Cornell University is a private statutory land-grant research university based in Ithaca, New York. It is a member of the Ivy League. Founded in 1865 by Ezra Cornell and Andrew Dickson White, Cornell was founded with the intention to teach an ...
Department of Plant Pathology and Plant Microbe Biology
The most prominent symptoms are hip
canker A plant canker is a small area of dead tissue, which grows slowly, often over years. Some cankers are of only minor consequence, but others are ultimately lethal and therefore can have major economic implications for agriculture and horticultur ...
s and swollen, spherical to oblong growths (galls) on the branches, stems, or main trunk of the host. The galls on small twigs of 1–2 years of age are often
pear Pears are fruits produced and consumed around the world, growing on a tree and harvested in the Northern Hemisphere in late summer into October. The pear tree and shrub are a species of genus ''Pyrus'' , in the family Rosaceae, bearing the p ...
-shaped. Gall formation results from the overproduction of
xylem Xylem is one of the two types of transport tissue in vascular plants, the other being phloem. The basic function of xylem is to transport water from roots to stems and leaves, but it also transports nutrients. The word ''xylem'' is derived from ...
tissue stimulated by the fungus. Witches brooming occasionally occurs along with galling. The bark on large galls will slough off over time, exposing the smooth wood beneath. Signs of pine-pine gall rust include the pale yellow aecia (1-8mm in diameter) formed on galls in spring as well as the yellow-orange
aeciospore Aeciospores are one of several different types of spores formed by Rusts. They each have two nuclei and are typically seen in chain-like formations in the aecium. References Fungal morphology and anatomy {{mycology-stub ...
s contained within. Immature galls are spindle-shaped. Infections usually occur on more succulent (thick) branches. When mature, galls can grow as large as a softball (30.5 cm circumference), but most are the size of a golf ball (4.2 cm). Before the production of spores, the needles on highly infected branches become
chlorotic In botany, chlorosis is a condition in which leaves produce insufficient chlorophyll. As chlorophyll is responsible for the green color of leaves, chlorotic leaves are pale, yellow, or yellow-white. The affected plant has little or no ability to ...
or red, eventually turning brown when the branch dies. Western gall rust can cause dwarfing to occur if leader stem of younger pines are infected. Early identification of the disease is important for treatment, but is often hard to diagnose due to the inconspicuous initial symptoms and a lack of galls until the second year. A red pigmentation may develop on the epidermal galls of ''P. contorta'' seedlings 14–28 days after infection, but this is not always a reliable indicator.


Disease cycle

Pine-pine gall rust has characteristically brown to yellow-orange sori visible on large globular galls on pines. Gall formation on trunks occurs over 2–4 years and is stimulated by the pathogen, which causes cells to grow and divide quickly at the site of initial infection. When mature, the gall splits open and the yellow-orange aeciospores are dispersed and carried to new hosts by wind. Because ''P. harknessii'' does not require an alternate host, the aeciospores can infect another pine directly. This typically occurs during moist conditions and rarely infects older shoots. Infection continues on the host shoots and needles until they have reached 90% of their elongation. Gall mortality is associated with squirrel feeding or invasion and inactivation by secondary organisms. The life cycle of ''E. Harknessii'' is different from other pine stem rust in that it is autoecious, making large stands of pines ideal hosts for survival and reproduction of the fungus. Wind carries
teliospore Teliospore (sometimes called teleutospore) is the thick-walled resting spore of some fungi ( rusts and smuts), from which the basidium arises. Development They develop in '' telia'' (sing. ''telium'' or ''teliosorus''). The telial host is the p ...
s to pines shoots that then germinate under cool humid conditions, producing germ tubes with up to three side branches that act as
basidia A basidium () is a microscopic sporangium (a spore-producing structure) found on the hymenophore of fruiting bodies of basidiomycete fungi which are also called tertiary mycelium, developed from secondary mycelium. Tertiary mycelium is highly-c ...
. The basidium directly penetrate the
cuticle A cuticle (), or cuticula, is any of a variety of tough but flexible, non-mineral outer coverings of an organism, or parts of an organism, that provide protection. Various types of "cuticle" are non- homologous, differing in their origin, structu ...
and
epidermis The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and hypodermis. The epidermis layer provides a barrier to infection from environmental pathogens and regulates the amount of water rele ...
. Other pines rusts like ''
Cronartium ribicola ''Cronartium ribicola'' is a species of rust fungus in the family Cronartiaceae that causes the disease white pine blister rust. Other names include: (French), (German), (Spanish). ''Cronartium ribicola'' is native to China, and was subsequ ...
'' and '' C. comandrae'' infect through the
stoma In botany, a stoma (from Greek ''στόμα'', "mouth", plural "stomata"), also called a stomate (plural "stomates"), is a pore found in the epidermis of leaves, stems, and other organs, that controls the rate of gas exchange. The pore is bor ...
ta. After penetration and establishment of a intracellular infection structure, primary
hypha A hypha (; ) is a long, branching, filamentous structure of a fungus, oomycete, or actinobacterium. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium. Structure A hypha consists of one or ...
e are produced, infecting the epidermis and cortex intercellularly.
Haustoria In botany and mycology, a haustorium (plural haustoria) is a rootlike structure that grows into or around another structure to absorb water or nutrients. For example, in mistletoe or members of the broomrape family, the structure penetrates t ...
extend through neighboring cells and the cortex to reach the vascular cambium before the host becomes dormant follow the first infection season. The cambium is invade inwardly through the
phloem Phloem (, ) is the living biological tissue, tissue in vascular plants that transports the soluble organic compounds made during photosynthesis and known as ''photosynthates'', in particular the sugar sucrose, to the rest of the plant. This tran ...
and cortex, as opposed to a vertical or peripheral hyphael growth. Initiation of gall formation is through exogenous stimulation of the cambium and pith rays, causing an increased production of ray
parenchyma Parenchyma () is the bulk of functional substance in an animal organ or structure such as a tumour. In zoology it is the name for the tissue that fills the interior of flatworms. Etymology The term ''parenchyma'' is New Latin from the word π ...
. The host reacts by
hyperplasia Hyperplasia (from ancient Greek ὑπέρ ''huper'' 'over' + πλάσις ''plasis'' 'formation'), or hypergenesis, is an enlargement of an organ or tissue caused by an increase in the amount of organic tissue that results from cell proliferati ...
(increased division) providing the resources needed for further hyphae proliferation in the cortex, phloem, and cambium until the galls death. The gall will enlarge for 2 years and sporulate on the third.
Spermogonia {{Short pages monitor