Sarcoscypha coccinea, commonly known as the scarlet elf cup, scarlet elf cap, or the scarlet cup, is a species of fungus in the family Sarcoscyphaceae of the order Pezizales. The fungus, widely distributed in the Northern Hemisphere, has been found in Africa, Asia, Europe, North and South America, and Australia. The type species of the genus Sarcoscypha, S. coccinea has been known by many names since its first appearance in the scientific literature in 1772. Phylogenetic analysis shows the species to be most closely related to other Sarcoscypha species that contain numerous small oil droplets in their spores, such as the North Atlantic island species S. macaronesica. Due to similar physical appearances and sometimes overlapping distributions, S. coccinea has often been confused with S. occidentalis, S. austriaca, and S. dudleyi.
The saprobic fungus grows on decaying sticks and branches in damp spots on forest floors, generally buried under leaf litter or in the soil. The cup-shaped fruit bodies are usually produced during the cooler months of winter and early spring. The brilliant red interior of the cups—from which both the common and scientific names are derived—contrasts with the lighter-colored exterior. The edibility of the fruit bodies is not clearly established, but its small size, tough texture and insubstantial fruitings would dissuade most people from collecting for the table. The fungus has been used medicinally by the Oneida Indians, and also as a colorful component of table decorations in England. Molliardiomyces eucoccinea is the name given to the imperfect form of the fungus that lacks a sexually reproductive stage in its life cycle.
Taxonomy, naming, and phylogeny
The species was originally named Helvella coccinea by the Italian naturalist Giovanni Antonio Scopoli in 1772. Other early names include Peziza coccinea (Nikolaus Joseph von Jacquin, 1774) and Peziza dichroa (Theodor Holmskjold, 1799). Although some authors in older literature have applied the generic name Plectania to the taxon following Karl Fuckel's 1870 name change (e.g. Seaver, 1928; Kanouse, 1948; Nannfeldt, 1949; Le Gal, 1953), that name is now used for a fungus with brownish-black fruit bodies. Sarcoscypha coccinea was given its current name by Jean Baptiste Émil Lambotte in 1889.
Obligate synonyms (different names for the same species based on one type) include Lachnea coccinea Gillet (1880), Macroscyphus coccineus Gray (1821), and Peziza dichroa Holmskjold (1799). Taxonomic synonyms (different names for the same species, based on different types) include Peziza aurantia Schumacher (1803), Peziza aurantiaca Persoon (1822), Peziza coccinea Jacquin (1774), Helvella coccinea Schaeffer (1774), Lachnea coccinea Phillips (1887), Geopyxis coccinea Massee (1895), Sarcoscypha coccinea Saccardo ex Durand (1900), Plectania coccinea (Fuckel ex Seaver), and Peziza cochleata Batsch (1783).
Sarcoscypha coccinea is the type species of the genus Sarcoscypha, having been first explicitly designated as such in 1931 by Frederick Clements and Cornelius Lott Shear. A 1990 publication revealed that the genus name Sarcoscypha had been used previously by Carl F. P. von Martius as the name of a tribe in the genus Peziza; according to the rules of Botanical Nomenclature, this meant that the generic name Peziza had priority over Sarcoscypha. To address the taxonomical dilemma, the genus name Sarcoscypha was conserved against Peziza, with S. coccinea as the type species, to "avoid the creation of a new generic name for the scarlet cups and also to avoid the disadvantageous loss of a generic name widely used in the popular and scientific literature". The specific epithet coccinea is derived from the Latin word meaning "deep red". The species is commonly known as the "scarlet elf cup", the "scarlet elf cap", or the "scarlet cup fungus".
S. coccinea var. jurana was described by Jean Boudier (1903) as a variety of the species having a brighter and more orange-colored fruit body, and with flattened or blunt-ended ascospores. Today it is known as the distinct species S. jurana. S. coccinea var. albida, named by George Edward Massee in 1903 (as Geopyxis coccinea var. albida), has a cream-colored rather than red interior surface, but is otherwise identical to the typical variety.
|Phylogeny and relationships of S. coccinea and related species based on ITS sequences and morphological characteristics.|
Within the large area that includes the temperate to alpine-boreal zone of the Northern Hemisphere (Europe and North America), only S. coccinea had been recognized until the 1980s. However, it had been known since the early 1900s that there existed several macroscopically indistinguishable taxa with various microscopic differences: the distribution and number of oil droplets in fresh spores; germination behavior; and spore shape. Detailed analysis and comparison of fresh specimens revealed that what had been collectively called "S. coccinea" actually consisted of four distinct species: S. austriaca, S. coccinea, S. dudleyi, and S. jurana.
The phylogenetic relationships in the genus Sarcoscypha were analyzed by Francis Harrington in the late 1990s. Her cladistic analysis combined comparisons of the sequences of the internal transcribed spacer in the non-functional RNA with fifteen traditional morphological characteristics, such as spore features, fruit body shape, and degree of curliness of the "hairs" that form the tomentum. Based on her analysis, S. coccinea is part of a clade that includes the species S. austriaca, S. macaronesica, S. knixoniana and S. humberiana. All of these Sarcoscypha species have numerous, small oil droplets in their spores. Its closest relative, S. macaronesica, is found on the Canary Islands and Madeira; Harrington hypothesized that the most recent common ancestor of the two species originated in Europe and was later dispersed to the Macaronesian islands.
Initially spherical, the fruit bodies are later shallowly saucer- or cup-shaped with rolled-in rims, and measure 2–5 cm (0.8–2.0 in) in diameter. The inner surface of the cup is deep red (fading to orange when dry) and smooth, while the outer surface is whitish and covered with a dense matted layer of tiny hairs (a tomentum). The stipe, when present, is stout and up to 4 cm (1.6 in) long (if deeply buried) by 0.3–0.7 cm (0.1–0.3 in) thick, and whitish, with a tomentum. Color variants of the fungus exist that have reduced or absent pigmentation; these forms may be orange, yellow, or even white (as in the variety albida). In the Netherlands, white fruit bodies have been found growing in the polders.
Sarcoscypha coccinea is one of several fungi whose fruit bodies have been noted to make a "puffing" sound—an audible manifestation of spore-discharge where thousands of asci simultaneously explode to release a cloud of spores.
Spores are 26–40 by 10–12 µm, elliptical, smooth, hyaline (translucent), and have small lipid droplets concentrated at either end. The droplets are refractive to light and visible with light microscopy. In older, dried specimens (such as herbarium material), the droplets may coalesce and hinder the identification of species. Depending on their geographical origin, the spores may have a delicate mucilaginous sheath or "envelope"; European specimens are devoid of an envelope while specimens from North America invariably have one.
The asci are long and cylindrical, and taper into a short stem-like base; they measure 300–375 by 14–16 µm. Although in most Pezizales all of the ascospores are formed simultaneously through delimitation by an inner and outer membrane, in S. coccinea the ascospores located in the basal parts of the ascus develop faster. The paraphyses (sterile filamentous hyphae present in the hymenium) are about 3 µm wide (and only slightly thickened at the apex), and contain red pigment granules.
Anamorphic or imperfect fungi are those that seem to lack a sexual stage in their life cycle, and typically reproduce by the process of mitosis in structures called conidia. In some cases, the sexual stage—or teleomorph stage—is later identified, and a teleomorph-anamorph relationship is established between the species. The International Code of Nomenclature for algae, fungi, and plants permits the recognition of two (or more) names for one and the same organism, one based on the teleomorph, the other(s) restricted to the anamorph. The name of the anamorphic state of S. coccinea is Molliardiomyces eucoccinea, first described by Marin Molliard in 1904. Molliard found the growth of the conidia to resemble those of the genera Coryne and Chlorosplenium rather than the Pezizaceae, and he considered that this suggested an affinity between Sarcoscypha and the family Helvellaceae. In 1972, John W. Paden again described the anamorph, but like Molliard, failed to give a complete description of the species. In 1984, Paden created a new genus he named Molliardiomyces to contain the anamorphic forms of several Sarcoscypha species, and set Molliardiomyces eucoccinea as the type species. This form produces colorless conidiophores (specialized stalks that bear conidia) that are usually irregularly branched, measuring 30–110 by 3.2–4.7 µm. The conidia are ellipsoidal to egg-shaped, smooth, translucent (hyaline), and 4.8–16.0 by 2.3–5.8 µm; they tend to accumulate in "mucilaginous masses".
Similar species include S. dudleyi and S. austriaca, and in the literature, confusion amongst the three is common. Examination of microscopic features is often required to definitively differentiate between the species. Sarcoscypha occidentalis has smaller cups (0.5–2.0 cm wide), a more pronounced stalk that is 1–3 cm long, and a smooth exterior surface. Unlike S. coccinea, it is only found in the New World and in east and midwest North America, but not in the far west. It also occurs in Central America and the Caribbean. In North America, S. austriaca and S. dudleyi are found in eastern regions of the continent. S. dudleyi has elliptical spores with rounded ends that are 25–33 by 12–14 µm and completely sheathed when fresh. S. austriaca has elliptical spores that are 29–36 by 12–15 µm that are not completely sheathed when fresh, but have small polar caps on either end. The Macaronesian species S. macaronesica, frequently misidentified as S. coccinea, has smaller spores, typically measuring 20.5–28 by 7.3–11 µm and smaller fruit bodies—up to 2 cm (0.8 in) wide.
Ecology, habitat and distribution
A saprobic species, Sarcoscypha coccinea grows on decaying woody material from various plants: the rose family, beech, hazel, willow, elm, and, in the Mediterranean, oak. The fruit bodies of S. coccinea are often found growing singly or clustered in groups on buried or partly buried sticks in deciduous forests. A Hungarian study noted that the fungus was found mainly on twigs of European hornbeam (Carpinus betulus) that were typically less than 5 cm (2.0 in) long. Fruit bodies growing on sticks above the ground tend to be smaller than those on buried wood. Mushrooms that are sheltered from wind also grow larger than their more exposed counterparts. The fruit bodies are persistent and may last for several weeks if the weather is cool. The time required for the development of fruit bodies has been estimated to be about 24 weeks, although it was noted that "the maximum life span may well be more than 24 weeks because the decline of the colonies seemed to be associated more with sunny, windy weather rather than with old age." One field guide calls the fungus "a welcome sight after a long, desperate winter and ... the harbinger of a new year of mushrooming."
Common over much of the Northern Hemisphere, S. coccinea occurs in the Midwest, in the valleys between the Pacific coast, the Sierra Nevada, and the Cascade Range. Its North American distribution extends north to various locations in Canada and south to the Mexican state Jalisco. The fungus has also been collected from Chile in South America. It is also found in the Old World—Europe, Africa, Asia, Australia, and India. Specimens collected from the Macaronesian islands that once thought to be S. coccinea were later determined to be the distinct species S. macaronesica. A 1995 study of the occurrence of British Sarcoscypha (including S. coccinea and S. austriaca) concluded that S. coccinea was becoming very rare in Great Britain. All species of Sarcoscypha, including S. coccinea, are Red-Listed in Europe. In Turkey, it is considered critically endangered.
The red color of the fruit bodies is caused by five types of carotenoid pigments, including plectaniaxanthin and β-carotene. Carotenoids are lipid-soluble and are stored within granules in the paraphyses. British-Canadian mycologist Arthur Henry Reginald Buller suggested that pigments in fruit bodies exposed to the Sun absorb some of the Sun's rays, raising the temperature of the hymenium—hastening the development of the ascus and subsequent spore discharge.
Lectins are sugar-binding proteins that are used in blood typing, biochemical studies and medical research. A lectin has been purified and characterized from S. coccinea fruit bodies that can bind selectively to several specific carbohydrate molecules, including lactose.
Sarcoscypha coccinea was used as a medicinal fungus by the Oneida Indians, and possibly by other tribes of the Iroquois Six Nations. The fungus, after being dried and ground up into a powder, was applied as a styptic, particularly to the navels of newborn children that were not healing properly after the umbilical cord had been severed. Pulverized fruit bodies were also kept under bandages made of soft-tanned deerskin. In Scarborough, England, the fruit bodies used to be arranged with moss and leaves and sold as a table decoration.
The species is said to be edible, inedible, or "not recommended", depending on the author. Although its insubstantial fruit body and low numbers do not make it particularly suitable for the table, one source claims "children in the Jura are said to eat it raw on bread and butter; and one French author suggests adding the cups, with a little Kirsch, to a fresh fruit salad." The fruit bodies have been noted to be a source of food for rodents in the winter, and slugs in the summer.
- "Sarcoscypha coccinea (Scop.) Lambotte 1889". MycoBank. International Mycological Association. Retrieved 2010-08-21.
- Scopoli JA. (1772). Flora carniolica (in Latin) 2 (2 ed.). Vienna, Austria: Sumptibus J.T. Trattner. p. 479.
- Jacquin NJ. (1774). Flora Austriaca 2. Vienna, Austria. p. 40.
- Holmskjold T. (1799). Beata ruris otia fungis danicis (in Latin) 2. p. 20.
- Fuckel L. (1870). "Symbolae mycologicae. Beiträge zur Kenntnis der rheinischen Pilze" [Contributions to the knowledge of mushrooms of the Rhein]. Jahrbücher des Nassauischen Vereins für Naturkunde (in German). 23–24: 324.
- Seaver FJ. (1928). The North American Cup-Fungi (Operculates). New York, New York: Self published. pp. 191–2.
- Kanouse BC. (1948). "The genus Plectania and its segregates in North America". Mycologia 40 (4): 482–97. doi:10.2307/3755155. JSTOR 3755155.
- Nannfeldt JA. (1949). "Contributions to the microflora of Sweden. 7. A new winter Discomycete, Urnula hiemalis Nannf. n. sp., and a short account of the Swedish species of Sarcoscyphaceae". Svensk Botanisk Tidskrift 43: 468–84.
- Le Gal M. (1953). "Les Discomycètes de Madagascar" [The Discomycetes of Madagascar]. Prodrome à Flore Mycologique de Madagascar et Dépendanes (in French) (Paris, France: Muséum National d’Histoire Naturelle) 4: 1–465.
- Korf RP; Harrington FA. (1990). "Proposal to conserve a type for Sarcoscypha (Fries) Boudier, S. coccinea (Jacq.) Lambotte (Fungi)". Taxon 39 (2): 342–3. doi:10.2307/1223069. JSTOR 1223069.
- "Lachnea coccinea (Jacq.) Gillet 1880". MycoBank. International Mycological Association. Retrieved 2010-08-21.
- "Macroscyphus coccineus (Scop.) Gray 1821". MycoBank. International Mycological Association. Retrieved 2010-08-21.
- "Peziza dichroa Holmsk. 1799". MycoBank. International Mycological Association. Retrieved 2010-08-21.
- Korf and Harrington (1990), citing Clements FE; Shear CL. (1931). The Genera of Fungi. New York, NY.
- von Martius CFP. (1817). Flora Cryptogamica Erlangensis (in Latin). Nuremberg, Germany: J.L. Schrag. p. 469.
- McKnight VB; McKnight KH. (1987). A Field Guide to Mushrooms, North America. Boston, Massachusetts: Houghton Mifflin. p. 33. ISBN 978-0-395-91090-0.
- Burrows I. (2005). Food from the Wild. London, UK: New Holland Publishers. p. 91. ISBN 978-1-84330-891-1.
- Arora D. (1986). Mushrooms Demystified: A Comprehensive Guide to the Fleshy Fungi. Berkeley, California: Ten Speed Press. p. 836. ISBN 978-0-89815-169-5.
- Boudier E. (1903). "Note sur quelques ascomycètes du Jura" [Notes on some ascomycetes of Jura]. Bulletin de la Société Zoologique de France (in French) 19: 193–9.
- "Species synonymy: Sarcoscypha jurana (Boud.) Baral". Index Fungorum. CAB International. Retrieved 2010-08-22.
- Massee G. (1892). British Fungus-Flora. A Classified Text-Book of Mycology 4. London, UK: George Bell & Sons. p. 378.
- Harrington FA. (1998). "Relationships among Sarcoscypha species: evidence from molecular and morphological characters". Mycologia 90 (2): 235–43. doi:10.2307/3761299. JSTOR 3761299.
- Baral HO. (1984). "Taxonomische und ökologische Studien über Sarcoscypha coccinea agg., Zinnoberrote Kelchbecherlinge. (Kurzfassung)" [Taxonomic and economic study on Sarcoscypha coccinea]. Zeitschrift für Mykologie (in German) 50 (1): 117–46.
- Harrington FA; Potter D. (1997). "Phylogenetic relationships within Sarcoscypha based upon nucleotide sequences of the internal transcribed spacer of nuclear ribosomal DNA". Mycologia 89 (2): 258–67. doi:10.2307/3761080. JSTOR 3761080.
- Van Duuren Y; Van Duuren G. (2005). "Witte Rode kelkzwammen en op excursie met Hans-Otto Baral" [White Sarcoscypha coccinea fruitbodies and a foray with Hans-Otto Baral] (PDF). Coolia 48 (3): 169–70.
- Buller, 1958, pp. 329–31. Retrieved 2010-08-22.
- Orr DB; Orr RT. (1980). Mushrooms of Western North America. California Natural History Guides. Berkeley, California: University of California Press. pp. 24–5. ISBN 978-0-520-03660-4.
- Harrington FA. (1990). "Sarcoscypha in North America (Pezizales, Sarcoscyphaceae)". Mycotaxon 38: 417–58.
- Merkus E. (1976). "Ultrastructure of the ascospore wall in Pezizales (Ascomycetes)—IV. Morchellaceae, Helvellaceae, Rhizinaceae, Thelebolaceae, and Sarcoscyphaceae. General discussion". Persoonia 9: 1–38.
- Hanlin RT. (1990). Illustrated Genera of Ascomycetes. St. Paul, Minnesota: American Phytopathological Society. pp. 54–5. ISBN 978-0-89054-107-4.
- Molliard M. (1904). "Forme conidienne de Sarcoscypha coccinea (Jacq.) Cooke. Bull" [Condidial form of Sarcoscypha coccinea (Jacq.) Cooke. Bull]. Bulletin trimestriel de la Société mycologique de France (in French) 20: 138–41.
- Paden JW. (1972). "Imperfect states and the taxonomy of the Pezizales". Persoonia 6: 405–14.
- Paden JW. (1984). "A new genus of Hyphomycetes with teleomorphs in the Sarcoscyphaceae (Pezizales, Sarcoscyphineae)". Canadian Journal of Botany 62 (2): 211–8. doi:10.1139/b84-035.
- Miller HR; Miller OK. (2006). North American Mushrooms: A Field Guide to Edible and Inedible Fungi. Guilford, Connecticut: Falcon Guide. p. 536. ISBN 978-0-7627-3109-1.
- Denison WC. (1972). "Central American Pezizales. IV. The genera Sarcoscypha, Pithya, and Nanoscypha". Mycologia 64 (3): 609–23. doi:10.2307/3757876. JSTOR 3757876.
- Kuo M. (2012). "Sarcoscypha dudleyi & Sarcoscypha austriaca". MushroomExpert.Com. Retrieved 2016-03-22.
- Korf RP; Zhuang W-Y. (1991). "A preliminary Discomycete flora of Macaronesia: part 11, Sarcoscyphineae". Mycotaxon 40: 1–11.
- Kuo M. (2006). "Sarcoscypha coccinea". MushroomExpert.com. Retrieved 2016-03-22.
- Baral HO. (2004). "Host specificity, plant communities". The European and North-American species of Sarcoscypha. Retrieved 2010-08-22.
- Bratek Z; Balazs T; Halasz K; Zld-Balogh A. (2003). "Adatok a Sarcoscypha es Microstoma Nemzetsegek fajainak karpat-medencei elterjedese ismeretehez" [Data on the genera Sarcoscypha and Microstoma in the Carpathian basin]. Mikologiai Kozlemenyek (in Hungarian) 42 (3): 3–16. ISSN 0133-9095.
- Brown RP. (1980). "Observations on Sarcoscypha coccinea and Disciotis venosa in North Wales during 1978–1979". Bulletin of the British Mycological Society 14 (2): 130–5. doi:10.1016/S0007-1528(80)80008-3.
- Smith AH; Weber NS. (1980). The Mushroom Hunter's Field Guide. Ann Arbor, Michigan: University of Michigan Press. p. 28. ISBN 978-0-472-85610-7.
- Abel D; Horn B; Kay R (1993). A Guide to Kansas Mushrooms. Lawrence, Kansas: University Press of Kansas. p. 238. ISBN 978-0-7006-0571-2.
- Téllez-Bañuelos C; Guzmán-Dávalos L; Guzmán G. (1988). "Contribucion al conocimiento de los hongos de le reserva de la biosfera de la Sierra de Manantlan, Jalisco" [Contribution to the knowledge of the fungi from the biosphere reserve of Sierra de Manantlan state of Jalisco Mexico]. Revista Mexicana de Micologia (in Spanish) 4: 123–30. ISSN 0187-3180.
- Tortella GR; Rubilar O; Gianfreda L; Valenzuela E; Diez MC. (2008). "Enzymatic characterization of Chilean native wood-rotting fungi for potential use in the bioremediation of polluted environments with chlorophenols". World Journal of Microbiology and Biotechnology 24 (12): 2805–18. doi:10.1007/s11274-008-9810-7.
- Zhishu B; Zheng G; Taihui L. (1993). The Macrofungus Flora of China's Guangdong Province (Chinese University Press). New York, NY: Columbia University Press. p. 37. ISBN 962-201-556-5.
- Butterfill GB; Spooner BM. (1995). "Sarcoscypha (Pezizales) in Britain". Mycologist 9 (1): 20–6. doi:10.1016/S0269-915X(09)80243-7.
- Baral HO. (2004). "Geographical Distribution". The European and North-American species of Sarcoscypha. Retrieved 2016-03-22.
- Çinar H; Sermenli HB; Işiloğlu M. (2014). "Some critically endangered species From Turkey" (PDF). Fungal Conservation (4): 26–28.
- Arpin N. (1968). "Les caroténoïdes des Discomycètes: essai chimiotaxinomique" [Carotenoids of the Discomycetes: chemotaxonomic analysis]. Bulletin Mensuel de la Société Linnéenne de Lyon (in French) 28 (Suppl): 1–169.
- Buller, 1958, p. 301. Retrieved 2010-08-22.
- Antoniuk VO. (2005). "[Purification and study of carbohydrate specificity of lectin from Sarcoscypha coccinea (Fr.) Lambette]" (PDF). Ukrainskii Biokhimicheskii Zhurnal (in Ukrainian) 77 (3): 96–103. PMID 16566135.
- Dickinson C; Lucas J. (1982). VNR Color Dictionary of Mushrooms. Van Nostrand Reinhold. pp. 20–1. ISBN 978-0-442-21998-7.
- Baker N. (2006). Nick Baker's British Wildlife. New Holland Publishers. p. 155. ISBN 978-1-84537-271-2.
- Davidson JL; Davidson A; Saberi H; Jaine T. (2006). The Oxford Companion to Food. Oxford [Oxfordshire]: Oxford University Press. p. 234. ISBN 978-0-19-280681-9.
- Buller AHR. (1958). Researches on Fungi 6. New York, New York: Hafner Publishing.
|Wikimedia Commons has media related to Sarcoscypha coccinea.|