Wednesday, November 11, 2015

Armillaria mella

Nomenclature:

Armilliaria mellea is a complex group of roughly ten very closely related species that used to be named Armillaria melleus [6,13,14]. The taxonomy of this group has been difficult, mycologists have been making corrections to this genus for quite some time. At first they thought to put the honey colored mushrooms in the Armillaria genus, but with further research they discovered that over 200 species of them did not actually belong to that genera [10].  


Classification of A. mellea has been difficult due to the species not being easily identifiable in the field [7]. Molecular methods such as DNA sequencing are required in addition to field characteristics to help identify the fungus [7].

General Description:

The word melle means honey, and A. mellea are commonly called honey mushrooms [6,14]. The mushroom did not get this nickname from its taste, but from the honey colored caps that are produced [2].The caps of these mushrooms are convex in shape with brown to black hairs up near the center [14]. The entire cap is close to a yellowish brown color, but with age the caps will darken [14]. Most of the mushrooms spotted in the field will have a ring around the upper part of the stalk called a veil, this is known to disappear on some of them [3,14]. The veil is thick, flaring outward from the stalk, and is white in color [14].

A. mellea is a soil-borne fungus that can be found year round [12]. They are most common in late winter and after rain in the fall [2,10].

                                                   Photo of A. mella mushrooms
Photo © Pam Kaminski [6] 

Parasitic:

A. mellea are plant pathogens meaning that they infect and kill plants [1,2]. It causes root disease, rot, and wood decay in a broad range of host plants [2,9,15]. The most susceptible hosts are ones that have been previously weakened or injured, but A. mellea is known to infect healthy trees as well [4,15]. The fungus attacks the tissue located between the xylem and phloem of the tree which causes death in some of the main roots located close to the trunk [1]. A. mellea are considered very serious pathogens because of how well they can spread and infect. The fungus can infect neighboring trees by moving through the soil [2].

Mycorrhizal:

Interestingly, A. mellea can also form mycorrhizal relationships with certain plants. The fungus forms endotrophic mycorrhiza with orchids located in the tropics [9,14]. The orchid genus Gastrodia actually requires an infection from A. mellea to complete its life cycle and grow the structure called protocorm [14]. The mycelium of A. mellea is now within the protocom of the orchid and connects to nearby trees forming a parasitic relationship with the tree [14]. This relationship between A. mellea and the trees allows for the appropriate nutrients to benefit the orchids’ growth [14].

How to spot an infected tree:

An infection caused by A. mellea can be difficult to see unless it is in the sexual form where mushrooms have developed [15]. If there are not mushrooms present and the tree has multiple dead branches, yellow colored leaves, or reduced growth, then one can pull bark off of the tree [4,6,15]. By pulling the bark off the tree, the mycelium or dark rhizomorphs of the fungi will be visible. The mycelium is often white and describes as looking fan shaped [4,15]. If a tree is infected it may die rather quickly, or slowly over a period of time [4,6].

Photo of the dark rhizomorphs growing on a tree.
Photo ©  Dave Kelly                                 
http://www.first-nature.com/fungi/armillaria-mellea.php


Photo is of the fan shaped mycelial growth. 















                             




















Distribution:

A. mellea is commonly found throughout temperate and tropical regions [5,14,15]. They occur in the Eastern part of North America ranging from the Great Lakes to the east and gulf coast [10].  These fungi are native to California where people refer to them as “oak root fungus” or “shoestring rot” due to the darkly pigmented rhizomorphs. Rhizomorphs resemble thick string like structures colored black [1,2].

According to a study completed by Robert Raabe, the following U.S states have reported A. mellea findings: California, Washington, New Mexico, Idaho, Colorado, Arizona, Utah, Montana, New Jersey, Pennsylvania, New York, Maine, Maryland, as well as some North central states. This study also displays other countries that have found A. mellea including but not subject to: Germany, Italy, India, Poland, Canada, Switzerland, Denmark, Australia, Tanganyika, Russia, etc. [13] This particular study gives insight as to how far spread this particular fungus is.
Interestingly A. mellea was found and identified in Cape Town South Africa [5]. The fungus was DNA sequenced which linked them to the species that originated in Europe. A. mellea is believed to have been introduced to South Africa from Europe in the form of potted citrus or grapes tracing back to 300+ years ago [5].

Habitat:

A. mellea are commonly found on stumps and logs of living or dead trees [2,14]. They can also be found scattered on the ground but if this is the case then they are growing from roots or wood buried underneath the ground [2,14]. They typically need a root or piece of stem to survive, and they will survive for many years this way [4]. This fungus has an extremely large host range, but they are most commonly found on herbaceous plants, deciduous trees, conifers, and orchards [2,4,8,13,14]. A. mellea has also been recorded on hardwoods, gardens, forest shrubs, grape and citrus plants [4,5]. In South Africa the fungus has been found on Pinus and Eucalyptus species in addition to oak trees [5].

Look alikes:

It is always best to be cautious when identifying mushrooms because many will have one or more look alikes. A. mellea has two look alike fungi that are commonly misidentified [2,8]. Armillaria bulbosa is one of these look alikes [2]. This one is especially important to beware of because it is a poisonous mushroom [2]. A. bulbosa has a ring on the stalk the same way A. mellea does, but the cap is a smoother texture [2]. However the main look alike is Armillaria tabescens. This fungus clusters on wood and has the same spore print color as A. mellea, though it has a darker color cap and does not have a veil [2]. This is an edible fungus [8].

Nutrition:
A. mellea obtains all of its required nutrition through the roots and lower trunk of their host tree [2]. They do this using extracellular enzymes to obtain food from the organic matter they are decaying [6].

How is A. mellea used?

As mentioned earlier, A. mellea will form a symbiotic relationship with the orchid Gastrodia elata, and this is currently being used in Chinese medicine [3]. Also, people in subarctic regions have been known to use this fungus as a way to mark their trails at night [2]. The fungi will luminesce at night allowing people to mark where they have previously been on a trail [2]. 

Toxicity:

A. mellea is an edible fungus when cooked correctly [2,14]. When preparing these mushrooms to eat they must be fully cooked and the water they were cooked in has to be drained [2]. Only the firm caps should be eaten while the stalks are thrown out [2]. This species has been recorded to substitute shiitake mushrooms, but has also caused some gastrointestinal discomfort [2,14].

Damage to crops:

Unfortunately A. mellea is the leading killer of gardens, plantations, and orchards [4]. When an infection occurs in an orchard the fungus will prevent any new development in those areas [4,8].

Bioluminescence:

The genus Armillaria  have the ability to luminesce during the night [7]. The species mellea is commonly called “foxfire” due to the glow produced at night [2,6]. The fungi will glow the brightest when they are still growing in decaying wood [7]. The best time to observe the glow is at night during the fall months. They can be exposed by kicked up the leaf litter on the ground [7]. The bioluminescence of fungi has been researched, but mycologists do not yet have a clear understanding of why they do so [2].

                     Photo of A. mellea during the night time, commonly referred to as "foxfire".
                                         Picture by Smita Bernadet Kujur
                                                       http://strangemushrooms.blogspot.com/2010/10/glowing-armillaria-mellea.html

References:
1.      Agriculture and Natural Resources. 2015. Armillaria root rot- Armillaria mellea. University of California.
http://www.ipm.ucdavis.edu/PMG/r302100811.html

2.      Arora, D. 1986. Mushrooms Demystified. Berkeley, CA: Published by Ten Speed Press. 196-197 p.

3.      Barron, G. 1999. Mushrooms of Northeast North America Midwest to New England. Auburn, WA. Published by Lone Pine Publishing. 246 p.


4.      Basnayake V. 2001. Armillaria mellea. North Caroline State University.
http://www.cals.ncsu.edu/course/pp728/Armillaria/Armillaria.htm

5.      Coetzee M, Wingfield BD, Harrington TC, Steimel J, Coutinho TA, Wingfield MJ. 2001. The root rot fungus Armillaria mellea introduced into South Africa by early Dutch settlers. Molecular Ecology.
http://www.public.iastate.edu/~tcharrin/Dutch%20Armillaria.pdf

6.      Emberger, G. 2008. Armillaria mellea. Messiah College.

7.      Huffman DM, Tiffany LH, Knaphus G, Healy RA. 2008. Mushrooms and Other Fungi of the Midcontinental United States. Iowa City, IA: Published by University of Iowa Press. 128-129 p.

8.      Kerik CD, Harold S, Beckman TG. Armillaria Root and Crown Rot. Department of Plant Pathology. University of Georgia.
               http://www.ent.uga.edu/peach/peachhbk/fungal/armillariacr.pdf
9.      Kirk PM, Cannon PF, Minter DW, Stalpers JA. 2008. Dictionary of the Fungi. Wallingford, UK. Published by Cab International. 49 p.


10.   Kuo, M. (2004, October). Armillaria mellea. Retrieved from the MushroomExpert.Com Web site: http://www.mushroomexpert.com/armillaria_mellea.html

11.   Muszynska B, Sulkowska-Ziaja K, Wolkowska M, Ekiert H. Chemical, pharmacological, and biological characterization of the culinary-medicinal honey mushroom, Armillaria mellea (Vahl) P. Kumm. (Agaricomycetideae). Europe PubMed Central.
http://www.ncbi.nlm.nih.gov/pubmed/22135893

12.   Oak Woodland Management. 2015. Armillaria mellea: Native Soil Fungus Causing Root Rot.
 University of California.

13.   Raabe R. 1962. Host list of the root rot fungus,Armillaria mellea. Hilgardia 33(2):25-88.

14.   Webster J, Weber RWS. 2007. Introduction to Fungi. Cambridge, NY: Published by Cambridge University Press. 1 p.


15.   Williams RE, Shaw CG, Wargo PM, Sites WH. 1989. Armillaria Root Disease. U.S Department of Agriculture Forest Service.
               http://www.na.fs.fed.us/spfo/pubs/fidls/armillaria/armillaria.htm