The cap of Amanita umbrinolutea is usually free of volval remnants, 45 - 90 mm wide, conico-paraboloid at first, then campanulate to convex and finally planar, umbonate, with a strongly striate margin (margins occupying 25 - 35% of the cap's radius. The cap has a distinctive pattern of color, often dark in the center, then pale, then dark over the inner edges of the lamellae and on the ridges between the marginal striations and at other times pallid in the center, but also strongly zonate; intensity of pigmentation is variable, with the center ranging from umber to grayish umber-brown to beige or pale grayish brown even within a single collection.
The gills are free, crowded, off-white to sordid pale cream in mass, and up to 6 mm broad; the short gills are truncate, of varying length, scattered and unevenly distributed.
The stem is 115 - 185 × 6 - 11 mm, pale cream to pale beige or isabella color or pale grayish brown, with faint appressed zigzag girdles of fibrils, and exannulate, with a fleshy membranous sack-like volva at the base. The 30 - 40 mm high volva is attached to the bottom 5 mm or so of the stipe.
The spores measure (9.5-) 10.5 - 13.4 (-17.2) × (8.5-) 9.5 - 12.5 (-14.8) µm and are subglobose (infrequently globose or broadly ellipsoid) and inamyloid. Clamps are absent from the bases of basidia.
Amanita umbrinolutea is widely distributed in Europe. The material I have examined has all been found in association with conifers (including pine and spruce); and its range extends eastward at least to northwestern Pakistan and northern India.—R. E. Tulloss
(Secr. ex Gillet) Bataille. 1910. Bull. Trimestriel Soc. Mycol. France 26: 139.
"Umber-zoned Ringless Amanita"
≡Amanita inaurata f. umbrinolutea Secr. ex Gillet. 1874. Champ. (Hyménomyc.) Croiss. France: 42. [Misapplication.]
≡Amanita vaginata var. umbrinolutea ("umbrino-lutea") (Secr. ex Gillet) E.-J. Gilbert. 1918. Gen. Amanita Pers.: 146.
≡Amanita vaginata subsp. badia var. umbrinolutea (Secr. ex Gillet) Konrad & Maubl. 1924. Icones Sel. Fung., Texte: 34.
≡Amanitopsis umbrinolutea (Secr. ex Gill.) E.-J. Gilbert. 1928. Bull. Trimestriel Soc. Mycol. France 44: 164.
≡Amanita vaginata f. umbrinolutea (Secr. ex Gillet) Veselý. 1933. Ann. Mycol. 31(4): 279.
≡Amanitopsis vaginata var. umbrinolutea (Secr. ex Gillet) E.-J. Gilbert. 1940. Iconogr. Mycol. (Milan) 27, suppl. (1): 75.
≡Amanitopsis umbrinolutea (Secr. ex Gillet) Courtec. "comb. ined." 1986. Clé Determin. Macroscop. Champ. Supér. Régions Nord France: 15/20b-24b. [The combination would have been superfluous if made.]
≡Amanitopsis vaginata var. umbrinolutea (Secr. ex Gillet) Wasser. 1988. Ukrayins'k. Bot. Zhurn. 45(6): 77. [Superfluous combination.]
The editors of this site owe a great debt to Dr. Cornelis Bas
whose famous cigar box files of Amanita nomenclatural information
gathered over three or more decades were made available to RET for computerization
and make up the lion's share of the nomenclatural information presented on this site.
This page reports original research by R. E. Tulloss
46 - 72 (-86) mm wide, zonate with three zones (with darkest zones over disc and marginal striations), over disc umber to grayish umber-brown (10YR 4/3-4) to beige or pale grayish brown (e.g., 10YR 7/3-4) even within single collection, in middle zone beige to grayish beige to somewhat yellowish gray-beige (10YR 6/3-4 or 10YR 7/3-4 or between 10YR 7/4 and 10YR 6-7/6), outer zone (over inner ends of striations and between striations) rather dark gray-brown (10YR 5/3-4 or 10YR 4/3) but sometimes paler, at first conico-paraboloid, soon broadly conical, then plano-conical, then plano-convex with umbo, finally subplanar with low obtuse subconical umbo; context white, membranous approximately in area of striations; margin sulcate-striate (0.25R - 0.35R), nonappendiculate; universal veil absent.
free and leaving narrow groove around stipe apex, crowded (approx. 10 - 12 in 10 mm arc at midradius of pileus), off-white to sordid pale cream, up to 6 mm broad, with thin rather straight gray-brown to yellowish gray-brown edge; lamellulae truncate, scattered, unevenly distributed (sometimes entirely missing from sector making up 25% or more of pileus), at most one between any pair of lamellae, of diverse lengths.
115 - 155 (-185) × 6 - 11 mm, pale cream to pale beige or isabella color or pale grayish brownish, with faint appressed zigzag girdles of fibrils (at first very pale pinkish brownish, sometimes brownish yellow, later increasingly gray-brown, finally becoming quite distinct on pale background), narrowing upward, slightly or markedly flaring at apex; context hollow; exannulate; universal veil as saccate volva, copious, white to sordid white, with few or many orange-brown spots on exterior, with approximately upper third to half becoming grayish with age, 30 - 40 mm high, of which 25 - 35 mm free from stipe, at first fleshy membranous and up to 2 mm thick, circumcissile or 2 - 3 lobed, with limbus internus small, subfloccose, visible only in younger basidiomes, positioned at about midheight of free limb (volva bitangent), often evanescent.
Odorless. Taste indistinct.
105 - 160 (-195) µm thick, colorless at surface in layer ranging from minimal to 80 µm thick, subtended by brownish orange region (progressively paler toward pileus context) 60 - 120+ µm thick, gelatinized only just at surface, lacking pronounced division into supra- and subpellis; filamentous, undifferentiated hyphae 3.2 - 7.6 µm wide, branching, sometimes constricted at septa, with common slightly inflated intercalary segments up to 12.7 µm wide, occasionally with yellowish subrefractive walls, occasionally strongly pigmented (orange-brown) in most strongly pigmented region, dominantly subradially arranged; refractive or vascular hyphae 6.1 - 12.1 µm wide, sinuous, scattered throughout, locally common.
filamentous, undifferentiated hyphae 3.7 - 7.6 µm wide, branching, forming open lattice, in fascicles and singly, sometimes with yellowish subrefractive walls, often constricted at septa; acrophysalides narrowly clavate to clavate, up to 94 × 35 µm, with walls thin or up to 1.0 µm thick; inflated cells in subradially oriented chains just above interlamellar basidia giving impression of pseudoparenchymatous tissue, ellipsoid to subglobose (e.g., 32 × 27 µm) or broadly allantoid to broadly fusiform to elongate-ellipsoid to subcylindric (e.g., 35 × 19 µm); vascular hyphae 4.8 - 9.0 µm wide, scattered throughout to locally common.
bilateral; wcs = (40-) 50 - 80 µm (excellent rehydration, higher numbers from material preserved in FAA and glycerine); central stratum comprising filamentous, undifferentiated hyphae and plentiful inflated intercalary cells (ellipsoid to clavate to subfusiform, up to 83 × 26 µm); subhymenial base with angle of divergence predominantly 45° - 60°, with plentiful ellipsoid to elongate ellipsoid to subfusiform to obclavate to narrowly obclavate inflated cells (up to 75 × 32 µm); filamentous, undifferentiated hyphae 3.6 - 11.0 µm wide, branching, common in central stratum and subhymenial base; terminal, divergent inflated cells not observed; vascular hyphae 5.5 - 10.2 µm wide, sinuous, scattered.
wst-near = (75-) 85 - 105 (-120) µm (excellent rehydration, higher numbers from material preserved in FAA and glycerine); wst-far = (90-) 100 - 125 (-155) µm (excellent rehydration, higher values obtained from material preserved in FAA and glycerine); comprising inflated and partially inflated cells in branching structure (with uncommon uninflated hyphal segments here and there), with two to three layers of inflated cells between bases of longest basidia and subhymenial base; with most basidia arising from inflated cells.
33 - 77 (-85) × 11.6 - 20 µm, often thin-walled, also with walls up to 1.0 µm thick especially in broadest part, dominantly 4-, occasionally up to one-third 2-sterigmate, with sterigmata up to 7.6 × 3.1 µm; dextrinoid granules not observed; clamps not observed.
On pileus: as scattered gelatinized microscopic fragments or absent. On stipe base, exterior surface: with scattered fascicles of subgelatinized to gelatinized filamentous, undifferentiated hyphae outermost and dominantly sublongitudinally oriented, with rather dense lattice of interwoven straight filamentous, undifferentiated hyphae as second layer (singly or in narrow fascicles, scattered hyphae exposed at surface partially gelatinized) with scattered inflated cells (collapsed, colorless to brown, ellipsoid to ovoid to subpyriform to subglobose to globose, up to 67 × 61 µm). On stipe base, interior: filamentous, undifferentiated hyphae 2.3 - 15.2 (-24) µm wide, branching, commonly curved or loosely coiled, dominating, rather densely interwoven, occasionally with yellowish subrefractive walls, singly or in fascicles; inflated cells terminal, singly or in chains of two, not tightly bound by surrounding hyphae, most frequently colorless, also slightly sordid to yellow-brown to brownish, globose to subglobose to subpyriform to broadly ovoid to ovoid (up to 105 × 80 µm, but rarely larger than 90 × 75 µm) and broadly clavate to clavate to subfusiform to subcylindric (up to 75 × 38 µm), with walls thin or up to 0.6 µm thick, unevenly distributed, plentiful locally; vascular hyphae 3.2 - 9.8 µm wide, scattered, sinuous. On stipe base, inner surface: with much area comprising exposed interior tissue (near top of limb especially), with scattered fascicles of sublongitudinally oriented gelatinized hyphae, especially lower on limb with such fascicles densely packed in broad patches or strips suggesting pileipellis.
longitudinally acrophysalidic; filamentous, undifferentiated hyphae 3.8 - 8.7 µm wide, branching, common, singly or in fascicles, plentiful near surfaces; acrophysalides up to 275 × 46 µm, dominating away from surfaces, with walls thin or up to 0.8 µm thick; refractive hyphae 8.9 - 19.0 µm wide, sinuous, rarely branching.
1. Elements of universal veil.
2. Elements of hymenium, subhymenial tree, and central stratum.
[257/10/6] (95-) 10.5 - 13.4 (-17.2) × (8.5-) 9.5 - 12.5 (-14.8) µm, (L = 11.0 - 12.3 (-13.2) µm; L' = 11.9 µm; W = 10.1 - 11.4 (-12.1) µm; W' = 11.0 µm; Q = (1.0-) 1.04 - 1.15 (-1.34); Q = 1.07 - 1.11; Q' = 1.09), hyaline, colorless, thin-walled, smooth, inamyloid, subglobose to (occasionally) broadly ellipsoid; apiculus sublateral, truncate-conic to cylindric; contents monoguttulate; occasionally with some spores containing central flocculent body appearing to be supported by radiating filaments attached to spore walls; color in deposit unrecorded for collections examined.
Subgregarious. Estonia: In Picea forest with plentiful Corylus on rather acid, but probably mineral-rich, loamy soil. India: At 1800 m elev. In mixed forest of Quercus incana Roxb. and Pinus roxburghii Sarg. or with Cedrus deodara (Roxb.) Loud (mycorrhizal association traced). Pakistan: At 2500 m elev. With P. roxburghii and P. wallichiana A. B. Jackson or with P. excelsa Lam.
ESTONIA: ca. 16 km S of Villandi, 26.viii. 1989 C. Bas 9203 (in herb. David T. Jenkins; L; RET 024-4).
INDIA: UTTARAKHAND—ca. Shimla along path to Glen, 3.ix.1987 S. L. Stephenson 87-5 (BPI; RET 327-5); Kullu, Jalori Pass, 11.viii.1984 A. Kumar s.n. (BPI 71983; HPU 1398 as "ceciliae"), s.n. (HPUB 3714).
NORWAY: Gjövik Co. - tract around Honne in Biri [NN 8858 (1816 I)], 24.viii.1985 T. E. Brandrud & J. Stordal 24317 (O).
PAKISTAN: NW FRONTIER PROV.—Hazara Distr. - Ayoubia-Khanspur, 10.viii.1995 A. N. Khalid 10895 (LAH; RET 291-9), 10.viii.1997 S. H. Iqbal & A. N. Khalid s.n. (LAH; RET 270-8).
Because of the incomplete state of knowledge of the taxonomy of A. umbrinolutea (see below), material from Pakistan was compared to a collection from Estonia and a second, clearly conspecific collection from Norway. It is particularly important that the Estonian collection was described in considerable detail in its fresh state by the collector, Dr. C. Bas (Rijksherbarium, Leiden) and therefore is known to strongly conform to Secretan's description of the present species.
The European and Pakistani collections were compared:
· Habit and pigmentation of the basidiomes are extremely similar in the European and Pakistani collections.
· A few inflated cells in the volva of the Pakistani material were slightly larger than any seen in the European material. Otherwise, the structure of the universal veil is identical in the two sets of material.
· A very slight difference in the numerical parameters of the lamellae was noted, but this was attributed to comparison of rehydrated tissue in the European collections with material preserved in liquid in the Pakistani collections.
· Structure of the lamella trama in the two sets of material is nearly identical.
· Allowing for the presence of giant spores (seven of 20 measured) in one of the Pakistani collections (Khalid 10895), A. umbrinolutea spores from Europe and Pakistan were very similar: [100/5/2] (10.0-) 10.5 - 12.8 (-17.2) × (9.0-) 9.5 - 11.5 (-13.7) µm, (L = 11.0 - 12.2 µm; L' = 11.5 µm; W = 10.1 - 11.0 µm; W' = 10.5 µm; Q = (1.04-) 1.05 - 1.15 (-1.26); Q = 1.09 - 1.11; Q' = 1.10) in the European material and [40/2/2] (10.5-) 11.4 - 13.9 (-16.4) × (9.5-) 10.5 - 12.6 (-14.3) µm, (L = 12.1 - 13.2 µm; L' = 12.7; W = 11.1 - 12.1 µm; W' = 11.6 µm; Q = 1.04 - 1.13 (-1.34); Q = 1.09; Q' = 1.09) in the Pakistani material. [For the best preserved Pakistani collection (date 10.viii.1997) the match to the European data is very good: [20/1/1] (10.5-) 11.0 - 12.8 (-15.1) × (9.5-) 9.9 - 11.6 (-13.0) µm, (L = 12.1 µm; W = 11.1 µm; Q = 1.04 - 1.11 (-1.34); Q = 1.09).]
We conclude that the Pakistani and European collections are conspecific.
The volva of A. umbrinolutea is rather unusual since it remains coherent throughout the life of the basidiome, but also includes inflated cells that may eventually discolor to brown as are found in taxa of Amanita section Vaginatae with less coherent universal veil such as A. reidiana Tulloss, A. ceciliae (Berk. & Broome) Bas, A. olivaceogrisea Kalamees, A. sinicoflava Tulloss, and A. submembranacea (Bon) Gröger. Recently, two taxa that are somewhat similar macroscopically to A. umbrinolutea have been described in detail from southwestern China—A. atrofusca Zhu L. Yangand A. lignitincta Zhu L. Yang nom. prov.; however, neither has a subhymenium largely composed of inflated cells as in the present species. This can be seen in the superb and detailed illustrations of Yang (1997: figs. 56 & 67). In Tulloss' current classification of the species of section Vaginatae (unpub. data), the species that is most similar phenetically to A. umbrinolutea is A. pekeoides G. S. Ridl. (1991), described from New Zealand. Amanita pekeoides can be distinguished (Tulloss, unpub. data) from the present taxon by its having
· a pileus (at least sometimes) with vinaceous tints
· a pileipellis 70 - 100 µm thick
· inflated cells in its universal veil that are apparently all unpigmented and are smaller [up to 53 × 27 µm (most under 30 × 20 µm)], less common, and thin-walled
· lamellae lacking a colored edge
· smaller inflated cells in the lamella trama (none seen larger than 35 × 20 µm).
The great variation in spore dimensions given by European mycologists for the present species is somewhat puzzling and has suggested to several authors that there may be more than one species involved. The same impression can be obtained from the variety of colors (e.g., with some strong reddish tints to the dark zones) depicted for the pileus of the present species in various illustrations. It is also possible that some investigators have reported finding giant spores, and some have not. The situation of A. umbrinolutea taxonomy prior to the current work is discussed in brief by Fraiture (1993: 70-73). Fraiture says that most current authors describe the spores as globose and 10 - 12 (-13) µm. Allowing for the usual overemphasis of roundness common to most descriptions of subglobose spores in Amanita (Tulloss, 1994), this size is in approximate agreement with spore data from the collections we examined.
The unusual and unexplained phenomenon of spores on occasional basidiomes having central flocculent bodies supported by fine filaments attached to the interior of the spore wall was noted both in Bas 9203 and in an English collection [Epsom Common, Prince's Covert, 1.xi.1984 C. Whaley s.n. (K)] discussed by Tulloss and Halling (1997: 287).—revisions by R. E. Tulloss
—R. E. Tulloss
Information to support the viewer in reading the content of "technical" tabs
can be found here.
(Secr. ex Gillet) Bataille
"Umber-zoned Ringless Amanita"
1. Amanita umbrinolutea, Switzerland.
2. Amanita umbrinolutea, Germany.
1. Elements of universal veil.
2. Elements of hymenium, subhymenial tree, and central stratum.