9/18/2017 Morchella rufobrunnea - Wikipedia

Morchella rufobrunnea
From Wikipedia, the free encyclopedia

Morchella rufobrunnea, commonly known as the blushing morel, is a

species of ascomycete fungus in the family Morchellaceae. A choice Morchella rufobrunnea
edible species, the fungus was originally described as new to science in
1998 by mycologists Gastn Guzmn and Fidel Tapia from collections
made in Veracruz, Mexico. Its distribution was later revealed to be far
more widespread after several DNA studies suggested that it is common
in the West Coast of the United States, Israel, Australia, and Cyprus.

M. rufobrunnea grows in disturbed soil or in woodchips used in

landscaping, suggesting a saprophytic mode of nutrition. Reports from the
Mediterranean under olive trees (Olea europaea), however, suggest the
fungus may also be able to form facultative tree associations. Young fruit
bodies have conical, grayish caps covered with pale ridges and dark pits; Scientific classification
mature specimens are yellowish to ochraceous-buff. The surface of the Kingdom: Fungi
fruit body often bruises brownish orange to pinkish where it has been
touched, a characteristic for which the fungus is named, the New Latin Division: Ascomycota
rufobrunnea signifying "rufus brown". Mature fruit bodies grow to a
Class: Pezizomycetes
height of 9.015.5 cm (3.56.1 in). M. rufobrunnea differs from other
Morchella species by its urban or suburban habitat preferences, in the Order: Pezizales
color and form of the fruit body, the lack of a sinus at the attachment of
Family: Morchellaceae
the cap with the stipe, the length of the pits on the surface, and the
bruising reaction. A process to cultivate morels now known to be Genus: Morchella
M. rufobrunnea was described and patented in the 1980s.
Species: M. rufobrunnea
Binomial name
Contents Morchella rufobrunnea
Guzmn & F.Tapia (1998)
1 Taxonomy
2 Description
2.1 Similar species Morchella rufobrunnea
3 Habitat and distribution Mycological characteristics
4 Cultivation
5 References smooth hymenium
6 External links
cap is conical

Taxonomy or ovate
stipe is bare
The first scientifically described specimens of Morchella rufobrunnea were
collected in June 1996 from the Ecological Institute of Xalapa and other
spore print is cream
regions in the southern Mexican municipality of Xalapa, Veracruz, which
are characterized by a subtropical climate. The type locality is a to yellow
mesophytic forest containing oak, sweetgum, Clethra and alder at an
ecology is saprotrophic
altitude of 1,350 m (4,430 ft).[1] In a 2008 study, Michael Kuo determined
that the "winter fruiting yellow morel"erroneously referred to as edibility: choice
Morchella deliciosafound in landscaping sites in the western United

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States was the same species as M. rufobrunnea. According to Kuo,[2] David Arora depicts this species in his
popular 1986 work Mushrooms Demystified, describing it as a "coastal Californian form of Morchella deliciosa
growing in gardens and other suburban habitats".[3] Kuo suggests that M. rufobrunnea is the correct name for the
M. deliciosa used by western American authors.[4] Other North American morels formerly classified as deliciosa
have since been recategorized into two distinct species, Morchella diminutiva and M. sceptriformis
(=M. virginiana).[5]

Molecular analysis of nucleic acid sequences from the internal transcribed spacer and elongation factor EF-1
regions suggests that the genus Morchella can be divided into three lineages. M. rufobrunnea belongs to a lineage
that is basal to the esculenta clade ("yellow morels"), and the elata clade ("black morels").[6][7] This phylogenetic
placement implies that it has existed in its current form since the Cretaceous era (roughly 145 to 66 million years
ago), and all known morel species evolved from a similar ancestor.[8] M. rufobrunnea is genetically closer to the
yellow morels than the black morels.[7] M. anatolica, described from Turkey in 2012, is a closely related sister
species.[9]

The specific epithet rufobrunnea derives from the Latin roots ruf- (rufuous, reddish) and brunne- (brown).[3]
Vernacular names used for the fungus include "western white morel",[10] "blushing morel",[4] andaccounting for
the existence of subtropical species in the "blushing clade""red-brown blushing morel".[11]

Description
Fruit bodies of M. rufobrunnea can reach 6.021.0 cm (2.48.3 in) tall,
although most are typically found in a narrower range, 9.015.5 cm (3.5
6.1 in). The conical to roughly cylindrical hymenophore (cap) is typically 6.0
8.5 cm (2.43.3 in) high by 3.04.5 cm (1.21.8 in) wide. Its surface is
covered with longitudinal anastomosed ridges and crosswise veins that form
broad, angular, elongated pits. Young fruit bodies are typically dark grey with
sharply contrasting beige or buff ridges, while mature specimens fade to
ochraceous-buff. The cylindrical stipe is often strongly wrinkled, enlarged at
Tiny dark granules are on the apex the base and measures 3070 cm (1228 in) by 12.5 cm (0.41.0 in) thick. It
of the stipe. is typically covered with a dark brown to greyish pruinescence, often fading at
maturity, a useful character to discriminate it from similar species, such as M.
tridentina or M. sceptriformis. The stipe and hymenophore often exhibit
ochraceous, orange or reddish stains, although this feature is neither constant
nor exclusive to M. rufobrunnea and can be seen in a number of Morchella
species, such as Morchella tridentina (=Morchella frustrata), M. esculenta, M.
guatemalensis, the recently described M. fluvialis (Clowez et al. 2014), and
most likely M. anatolica.[12]
The translucent, egg-shaped spores
are up to 24 m long. In deposit, the spores are pale orange to yellowish orange. Ascospores are
egg-shaped, measuring 2024 by 1416 m when mature, but smaller (14.5
19 by 910 m) in immature fruit bodies. They are thin-walled, hyaline
(translucent), and inamyloid. The cylindrical asci (spore-bearing cells) are 300360 by 1620 m with walls up to
1.5 m thick. Paraphyses measure 90184 by 1018.5 m (69 m thick if immature); they are hyaline, have a
septum at the base, and comprise either one or two cells. The flesh is made of thin-walled, hyaline hyphae
measuring 39 m wide.[1]

Morchella rubobrunnea is an edible fungus;[13] it has been described variously as "one of the tastiest members of
the morel family",[10] and alternately as "bland in comparison to other morel species".[14] Individual specimens
over 1 pound (0.45 kg) have been reported.[10]

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Similar species

Morchella tridentina (=Morchella frustrata) is also rufescent and very similar to M. rufobrunnea. It is found in
mountainous forests and maquis and forms a marked sinus at the attachment of the cap with the stem, which is pure
white. At maturity, it develops more or less parallel, ladderlike interconnecting ridges. Microscopically, it often has
moniliform paraphyses with septa extending in the upper half and has more regularly cylindrical or clavate 'hairs'
on the stem, up to 100 m long.[12] M. guatemalensis, found in Central America, has a color ranging from yellow
to yellowish-orange, but never grey, and it has a more distinct reddish to wine red bruising reaction.
Microscopically, it has smaller paraphyses, measuring 56103 by 6.513 m. The New Guinean species
M. rigidoides has smaller fruit bodies that are pale ochre to yellow, without any grey. Its pits are less elongated
than those of M. rufobrunnea, and it has wider paraphyses, up to 30 m.[1]

Morchella americana (=M. esculentoides), is widely distributed in North America, north of Mexico and has similar
colours to mature fruit bodies of M. rufobrunnea, but lacks the bruising reaction. M. diminutiva, found in
hardwood forests of eastern North America, has a smaller fruit body than M. rufobrunnea, up to 9.4 cm (3.7 in) tall
and up to 2.7 cm (1.1 in) wide at its widest point. Morchella sceptriformis (=Morchella virginiana) is found in
riparian and upland ecosystems from Virginia to northern Mississippi, usually in association with the American
tulip tree (Liriodendron tulipifera).[5]

Habitat and distribution

A predominantly saprophytic species, Morchella rufobrunnea fruit bodies grow singly or in clusters in disturbed
soil or woodchips used in landscaping. Large numbers can appear the year after wood mulch has been spread on
the ground.[13] Typical disturbed habitats include fire pits, near compost piles, logging roads, and dirt
basements.[15] Fruiting usually occurs in the spring, although fruit bodies can be found in these habitats most of the
year. Other preferred habitats include steep slopes and plateaus, and old-growth conifer forests.[10] In Cyprus, the
fungus is frequently reported from coastal, urban and suburban areas under olive trees (Olea europaea).[12][16]

Morchella rufobrunnea ranges from Mexico through California and Oregon in the United States.[2] It has also been
introduced to central Michigan from California.[5] It is one of seven Morchella species that have been recorded in
Mexico.[1] In 2009, Israeli researchers used molecular genetics to confirm the identity of the species in northern
Israel, where it was found growing in gravelly disturbed soil near a newly paved path at the edge of a grove. This
was the first documented appearance of the fungus outside the American continent. Unlike North American
populations that typically fruit for only a few weeks in spring, the Israeli populations have a long-season ecotype,
fruiting from early November to late May (winter and spring). This period corresponds to the rainy season in Israel
(October to May), with low to moderate temperatures ranging from 1528 C (5982 F) during the day and 5
15 C (4159 F) at night.[17]

Cultivation
Morchella rufobrunnea is the morel that is cultivated commercially per US patents 4594809[18] and 4757640.[5][19]
This process was developed in 1982 by Ronald Ower with what he thought was Morchella esculenta;[18]
M. rufobrunnea had not yet been described. The cultivation protocol consists of preparing a spawn culture that is
mixed with nutrient-poor soil. This mixture is laid on nutrient-rich soil and kept sufficiently moist until fruiting. In
the nutrient-poor substrate, the fungus forms sclerotiahardened masses of mycelia that serve as food reserves.
Under appropriate environmental conditions, these sclerotia grow into morels.[20]

The fruit bodies of Morchella rufobrunnea have been cultivated under controlled conditions in laboratory-scale
experiments. Primordia, which are tiny nodules from which fruit bodies develop, appeared two to four weeks after
the first watering of pre-grown sclerotia incubated at a temperature of 16 to 22 C (61 to 72 F) and 90% humidity.
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Mature fruit bodies grew to 7 to 15 cm (3 to 6 in) long.[21]

The early stages of fruit body development can be divided into four
discrete stages. In the first, disk-shaped knots measuring 0.51.5 mm
(0.020.06 in) appear on the surface of the substrate. As the knot expands
in size, a primordial stipe emerges from its center. The stipe lengthens,
orients upward, and two types of hyphal elements develop: long, straight
and smooth basal hairy hyphae and short stipe hyphae, some of which are
inflated and project out of a cohesive layer of tightly packed hyphal
elements. In the final stage, which occurs when the stipe is 23 mm (0.08
A collection of mature specimens from
0.12 in) long, immature caps appear that have ridges and pits with distinct
San Francisco, US
filament-like paraphyses. Extracellular mucilage that covers the ridge layer
imparts shape and rigidity to the tissue and probably protects it against
dehydration.[22]

References
1. Guzmn G, Tapia F (1998). "The known morels in Mexico, a description of a new blushing species,
Morchella rufobrunnea, and new data on M. guatemalensis" (http://www.cybertruffle.org.uk/cyberliber/5935
0/0090/004/0705.htm). Mycologia. 90 (4): 70514. JSTOR 3761230 (https://www.jstor.org/stable/3761230).
doi:10.2307/3761230 (https://doi.org/10.2307%2F3761230).
2. Kuo M. (2008). "Morchella tomentosa, a new species from western North America, and notes on M.
rufobrunnea" (http://www.mushroomexpert.com/Kuo_M_2008_Morchella_tomentosa.pdf) (PDF).
Mycotaxon. 105: 4416.
3. Arora D. (1986). Mushrooms Demystified: A Comprehensive Guide to the Fleshy Fungi. Berkeley,
California: Ten Speed Press. ISBN 978-0-89815-169-5.
4. Kuo M. (2005). Morels. Ann Arbor, Michigan: University of Michigan Press. p. 173. ISBN 978-0-472-
03036-1.
5. Kuo M, Dewsbury DR, O'Donnell K, Carter MC, Rehner SA, Moore JD, Moncalvo JM, Canfield SA,
Stephenson SL, Methven AS, Volk TJ (2012). "Taxonomic revision of true morels (Morchella) in Canada
and the United States". Mycologia. 104 (5): 115977. PMID 22495449 (https://www.ncbi.nlm.nih.gov/pubm
ed/22495449). doi:10.3852/11-375 (https://doi.org/10.3852%2F11-375).
6. Kanwal HK, Acharya K, Ramesh G, Reddy MS (2011). "Molecular characterization of Morchella species
from the Western Himalayan region of India". Current Microbiology. 62 (4): 124552. PMID 21188589 (http
s://www.ncbi.nlm.nih.gov/pubmed/21188589). doi:10.1007/s00284-010-9849-1 (https://doi.org/10.1007%2F
s00284-010-9849-1).
7. Barseghyan GS, Kosakyan A, Isikhuemhen OS, Didukh M, Wasser SP. "Phylogenetic analysis within genera
Morchella (Ascomycota, Pezizales) and Macrolepiota (Basidiomycota, Agaricales) inferred from nrDNA
ITS and EF-1 sequences". In Misra JK, Tewari JP, Desmukh SK. Systematics and Evolution of Fungi.
Progress in Mycological Research. 2. Boca Raton, Florida: Science Publishers. pp. 159205. ISBN 978-1-
57808-723-5.
8. O'Donnell K, Rooney AP, Mills GL, Kuo M, Weber NS, Rehner SA (2011). "Phylogeny and historical
biogeography of true morels (Morchella) reveals an early Cretaceous origin and high continental endemism
and provincialism in the Holarctic" (http://www.cbs.knaw.nl/morchella/images/FGB%202011%20Morchella.
pdf) (PDF). Fungal Genetics and Biology. 48 (3): 25265. PMID 20888422 (https://www.ncbi.nlm.nih.gov/p
ubmed/20888422). doi:10.1016/j.fgb.2010.09.006 (https://doi.org/10.1016%2Fj.fgb.2010.09.006).
9. Takn H, Bykalaca S, Hansen K, O'Donnell K (2012). "Multilocus phylogenetic analysis of true morels
(Morchella) reveals high levels of endemics in Turkey relative to other regions of Europe". Mycologia. 104
(2): 44661. PMID 22123659 (https://www.ncbi.nlm.nih.gov/pubmed/22123659). doi:10.3852/11-180 (http
s://doi.org/10.3852%2F11-180).
10. Jones B. (2013). The Deerholme Mushroom Book: From Foraging to Feasting (https://books.google.com/bo
oks?id=X5BVD0F7J6MC&pg=PA19). Victoria, British Columbia: TouchWood Editions. p. 19. ISBN 978-1-
77151-003-5.
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11. Pilz D, McLain R, Alexander S, Villarreal-Ruiz L, Berch S, Wurtz TL, Parks CG, McFarlane E, Baker B,
Molina R, Smith JE (2007). Ecology and Management of Morels Harvested From the Forests of Western
North America. General Technical Report PNW-GTR-710 (http://www.fs.fed.us/pnw/publications/gtr710/pn
w_gtr710a.pdf) (PDF) (Report). Portland, Oregon: United States Department of Agriculture, Forest Service,
Pacific Northwest Research Station. p. 4.
12. Loizides M, Alvarado P, Clowez P, Moreau PA, de la Osa LR, Palazn A (2015). "Morchella tridentina, M.
rufobrunnea, and M. kakiicolor: a study of three poorly known Mediterranean morels, with nomenclatural
updates in section Distantes". Mycological Progress. 14 (3): 1030. doi:10.1007/s11557-015-1030-6 (https://d
oi.org/10.1007%2Fs11557-015-1030-6).
13. Davis RM, Sommer R, Menge JA (2012). Field Guide to Mushrooms of Western North America (https://boo
ks.google.com/books?id=obp7jddvjt4C&pg=PA392). Berkeley, California: University of California Press.
p. 392. ISBN 978-0-520-95360-4.
14. Bone E. (2011). Mycophilia: Revelations from the Weird World of Mushrooms. New York, New York:
Rodale. p. 137. ISBN 978-1-60961-987-9.
15. Wood M, Stevens F. "Morchella rufobrunnea" (http://www.mykoweb.com/CAF/species/Morchella_rufobrun
nea.html). California Fungi. MykoWeb. Retrieved 2014-03-29.
16. Loizides M, Bellanger J-M, Lowez P, Richard F, Moreau P-A (2016). "Combined phylogenetic and
morphological studies of true morels (Pezizales, Ascomycota) in Cyprus reveal significant diversity,
including Morchella arbutiphila and M. disparilis spp. nov.". Mycological Progress. 15: 39.
doi:10.1007/s11557-016-1180-1 (https://doi.org/10.1007%2Fs11557-016-1180-1).
17. Masaphy S, Zabari L, Goldberg D (2009). "New long-season ecotype of Morchella rufobrunnea from
northern Israel" (http://www.redalyc.org/pdf/685/68511349005.pdf) (PDF). Micologia Aplicada
International. 21 (2): 4555. ISSN 1534-2581 (https://www.worldcat.org/issn/1534-2581).
18. 4594809 (https://worldwide.espacenet.com/textdoc?DB=EPODOC&IDX=4594809), Ower R, Mills GL,
Malachowski JA., "Cultivation of Morchella", published 17 June 1986, assigned to Neogen Corporation
19. States4757640 United States 4757640 (https://worldwide.espacenet.com/textdoc?DB=EPODOC&IDX=Unit
ed), Malachowski JA, Mills GL, Ower RD., "Cultivation of Morchella", published 19 July 1988, assigned to
Neogen Corporation
20. Stamets P. (2000). Growing Gourmet and Medicinal Mushrooms (3rd ed.). Berkeley, California: Ten Speed
Press. p. 421. ISBN 978-1-58008-175-7.
21. Masaphy S. (2010). "Biotechnology of morel mushrooms: Successful fruiting body formation and
development in a soilless system". Biotechnology Letters. 32 (10): 15237. PMID 20563623 (https://www.nc
bi.nlm.nih.gov/pubmed/20563623). doi:10.1007/s10529-010-0328-3 (https://doi.org/10.1007%2Fs10529-01
0-0328-3).
22. Masaphy S. (2005). "External ultrastructure of fruit body initiation in Morchella". Mycological Research.
109 (4): 50812. PMID 15912939 (https://www.ncbi.nlm.nih.gov/pubmed/15912939).
doi:10.1017/S0953756204002126 (https://doi.org/10.1017%2FS0953756204002126).

External links
Morchella rufobrunnea (http://www.indexfungorum.org/Names/NamesRecord.asp?RecordID=445064) in
Index Fungorum
Morchella rufobrunnea (http://www.mycobank.org/MycoTaxo.aspx?Link=T&Rec=445064) in MycoBank.

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