The following material that is not otherwise attributed in the text is based on original research of R. E. Tulloss.
The cap is pale green-yellow to gray-brown to dark red-brown sometimes with olivaceous tints and often darkest in the center. It is 70-97 mm wide, and often lined with distinct dark radial fibers. Infrequently one half of the cap will be pale and the other half will be distinctly brown. The cap is broadly rounded, bell-shaped, convex or planar. The cap is dull when moist and shiny when dry. The flesh of the cap is white except for the pale cream center, and stains brown. The flesh is 4.5-9 mm above the stem and thins evenly to the smooth edge. The edge does not have remnants of the volva hanging from it. The volva is largely absent or if present appears as scattered warts or small patches on the cap, which are easily separated and orange-brown to dark brown in color.
The white to cream white gills are free from the stem to slightly attached, and crowded. The gills have a faint line and a small downward projection on the top of the stem. The gills stain brown and are 5.5- - 6+ mm broad. The short gills are not squarely cut off.
The stem is 90 - 108 × 12 - 15 mm and is cream white above and bellow the ring. Orange-brown stains develop on the lower half of the stem and on the bulb, which also is finely grooved vertically. The stem narrows upward, and flares briefly at the top. The bulb is subglobose to hemispheric and has multiple deep vertical clefts in it. The flesh of the stem is white, stuffed, and stains orange-brown where insects have tunneled into it. The cream-white, skirt-like ring is near the top of the stem and is sometimes streaked with brown. The ring's top surface is radially lined and its bottom surface is smooth. It can become brown with age. The white volval remnants become brown with age and sometimes form an incomplete short projection on the top rim of the bulb.
The odor of A. brunnescens suggests freshly dug potatoes, especially in stipe and bulb context when just sectioned. No taste is recorded.
The spores measure (7.0-) 8.0 - 9.2 (-9.5)
× (6.5-) 7.2 - 8.5 (-9.2) µm and are
globose to subglobose (occasionally broadly ellipsoid)
and amyloid. Clamps are absent from bases of basidia.
The present species is known from eastern North America. The reader may wish to compare A. brunnescens to the European species A. asteropus Sabo ex Romagn.
There is some question as to whether A. aestivalis Singer is a distinct species; however, Singer described it as pure white with or without some yellow over the disk and bruising much more slowly than does A. brunnescens. At present RET follows Jenkins (1986) in maintaining the two taxa as separate. Relevant molecular studies have not been completed to date.
Amanita brunnescens var. pallida L. Krieg. is originally described in such a way as to include both pale citrin collections of A. brunnescens and A. aestivalis. There is no holotype for var. pallida, but there are a number of candidates for a lectotype.
Note that the cap color in A. brunnescens is very variable; the specimens in photograph 2 (above) were found in close proximity and include cap colors ranging from pale citrin to brown. One cap (second from the viewer's right) is half citrin and half brown.—R. E. Tulloss and N. Goldman
G. F. Atk. 1918. Proc. Amer. Philos. Soc. 57: 354.
"Brown American Star-Footed Amanita"
≡Amanitina brunnescens (G. F. Atk.) E.-J. Gilbert. 1940. Iconogr. Mycol. (Milan) 27, suppl.1: 78, tabs. 35-27 (numerous figs.). [Based on his naming of Amanita brunnescens f. straminea, it is clear that Gilbert included Amanita citrina sensu auct. amer. in Amanita brunnescens. Hence, many of the spores depicted in the cited figures of the present work may not be from specimens of the latter species.]
=Amanita brunnescens var. pallida L. Krieg. 1927. Mycologia 19: 308.
=Amanita phalloides var. fuliginea Ferry. 1911. Études Amanites. Amanites Mort.: 94, pl. 4 (figs. 1-4).
≡Amanita phalloides f. fuliginea ("Peck") E.-J. Gilbert. 1918. Gen. Amanita Pers.: 49. [Superfluous name.]
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.
The following text may make multiple use of each data field.
The field may contain magenta text presenting data from a type study
and/or revision of other original material cited in the protolog of the present taxon.
Macroscopic descriptions in magenta are a combination of data from the protolog and
additional observations made on the exiccata during revision of the cited original
The same field may also contain black text, which is data from a revision of the present
taxon (including non-type material and/or material not cited in the protolog).
Paragraphs of black text will be labeled if further subdivision of
this text is appropriate.
Olive text indicates a specimen that has not been
thoroughly examined (for example, for microscopic details) and marks other places in the text
where data is missing or uncertain.
The following material that is not otherwise attributed in the text is based on original research of R. E. Tulloss.
Up to 70 - 97 mm wide, pale citrin (e.g., yellower than 1A2, paler that 1A3) to olivaceous brown to dark gray-brown or dark red-brown, often distinctly virgate with pigment at least partially in radial fibrils, often darker over disc [e.g., in pallid citrin cap, pale sordid yellow (e.g., ca. 5B2 or browner than 5Y 8/3)], infrequently with ca. one-half of cap pallid and one-half of cap much darker brown, broadly rounded conic to broadly campanulate to convex to planar, viscid to tacky to dry, dull when moist, shiny when dry; context white except for pale cream in upper part of disc, staining brown, 4.5 - 9 mm thick above stipe, thinning evenly to margin; margin nonstriate, nonappendiculate, slightly incurved at first; universal veil absent or present as scattered warts and small patches, submembranous at least at first, friable, detersile, whitish of off-white at first, becoming orange-brown then dark brown on exposure.
free to very narrowly adnate with dense faint lines (10× lens) or small decurrent tooth (10× lens) on upper stipe, crowded, white to cream in mass, very pale cream to pale cream in side view, staining brown, 5.5- - 6+ mm broad, broadest at midlength; lamellulae subtruncate to subattenuate to attenuate, unevenly distributed, plentiful, of diverse lengths.
90 - 108 × 12- 15 mm, entirely whitish or pale cream above partial veil and white below, with orange-brown to brown stains developing in lower half of stipe and on bulb, narrowing upward, barely flaring at apex, finely striatulate, fibrillose-squamulose in lower half; bulb 22 - 28 × 30- 34 mm, abrupt, subglobose? to hemispheric to subhemispheric, with multiple deep vertical clefts; context stuffed (often limited to upper stipe) to solid, white, staining like stipe surface, pale orange-brown to orange-brown in larva tunnels, with 2 mm wide central cylinder with densely packed white fibrillose stuffing material; partial veil superior to subapical, cream, with infrequent brown radial streaks, sometimes becoming sordid with age, brown-staining especially on edge, membranous, skirt-like, striate above, smooth below; universal veil originally pallid, becoming brown, forming incomplete very short limb on bulb rim or absent.
Odor of freshly dug potatoes especially in stipe and bulb context when just sectioned. Taste not recorded.
Spot test for tyrosinase (L-tyrosine) - negative on stipe surface and context (only areas tested). Spot test for laccase (syringaldazine) - negative thoughout basidiome or slowly positive on cut edges of lamellae, slowly positive (intensifying over 15 min.) in spots in stipe context and pileus context near top of stipe and upper center of bulb or base of bulb. Spot test with 2-furaldehyde ("furfural") - negative throughout basidiome. 10% KOH - on lamella, pale yellow; on cap of pallid specimen, pale yellow. Conc. HNO3 - on cap context, instantly orange, becoming yellow; on lamellae instantly yellow-orange, becoming yellow; on stipe context yellow; on bulb context, yellow; on pileus surface of pallid specimen, instantly pink, soon fading to pale yellow. Test vouchers: Tulloss 8-15-85-B. 11-25-85-A, 8-4-95-B, 9-22=96=F. [Note: Compare 2-furaldehye spot test with those of similar taxa A. aestivalis and A. asteropus.—ed.]
Jenkins (1982): [-/-/1] 7.8 - 9.4 × 7.8 - 8.6 μm, (Q = 1.0 - 1.10; Q' = 1.07),
hyaline, thin-walled amyloid, globose to subglobose; apiculus sublateral, cylindric; contents guttulate;
color in deposit not recorded.
Composite data from all material revised by RET: [35/2/2] (7.0-) 7.5 - 9.2 (-9.5) × (6.5-) 7.0 - 8.5 (-9.2) µm, (L = 8.2 - 8.7 µm;
L’ = µm; W = 7.6 - 8.0 µm; W’ = µm; Q = (1.0-) 1.03 - 1.15 (-1.18); Q = 1.08; Q’ = 1.08), hyaline, colorless, thin-walled, amyloid, globose to subglobose, infrequently broadly ellipsoid; apiculus sublateral, ??; contents granular to monoguttulate; white in deposit.
Solitary to subgregarious to gregarious, rarely in "fairy rings." Missouri: In upland Quercus-Carya flatwoods. New Jersey: At 12 - 214 m elev. In sandy soil of Pinus-Quercus barrens (e.g., with P. rigida, Q. marilandica, Sassafras albidum, Smilax glauca, Vaccinium spp., etc.). New York: In loamy clay or with Tsuga canadensis in mixed woods including Pinus, Abies, Quercus, Liriodendron tulipifera, Betula, Fagus grandifolia, Populus, Acer, Fraxinus and other hardwoods. South Carolina: In sandy-clay under layer of loam and duff of Pinus-Quercus upland forest. Tennessee: At 805 m elev. West Virginia: With Pinus, T. canadensis, and Acer pensylvanicum.
Jenkins (1982): U.S.A.: Tompkins Co. - ca. Ithaca, Cayuga Lk., 30.vii.1917 Leva B. Walker 24227 (holotype, CUP).
CONNECTICUT—Middlesex Co. - East Haddam, Devil's Hopyard St. Pk., 3.ix.2011 Sandy Sheine s.n. [Tulloss 9-3-11-N] (RET 488-6, dark pileus variant); Salmon River St. For. (West) [41°32’58” N/ 72°27’01” W, 21 m], 22.ix.1996 Sandy Sheine s.n. [Tulloss 9-22-96-F] (RET 251-5, pale pileus variant). Tolland Co. - Gay City St. Pk. [41°43’23” N/ 72°26’38” W, 209 m], 31.viii.1997 R. E. Tulloss 8-31-97-C (RET 267-8, dark pileus variant); Somers, 29.ix.1986 Ellen Greer s.n. [Tulloss 9-29-86-EG9] (RET 224-2, pale pileus variant), s.n. [Tulloss 9-29-86-EG10 (RET 224-3, pale pileus variant), s.n. [Tulloss 9-29-86-EG11] (RET 224-4, pale pileus variant).
INDIANA—Lawrence Co. - Lake Monroe, Dean Wilderness [38.9762° N/ 86.3518° W, 210 m], 11.ix.2012 Stephen Russell s.n. [mushroomobserver #109416] (RET 529-10, nrITSs & nrLSU seq'd.).
MAINE—Penobscot Co. - Orono, Univ. of Maine, 10.viii.1991 NEMF 1991 participant s.n. [Tulloss 8-10-91-A] (in herb. F. Massart; RET 031-1, pale pileus variant).
MASSACHUSETTS—ca. border of Berkshire & Franklin Cos. - Mohawk Trail St. For., 16.viii.1981 M. A. King & R. E. Tulloss 8-16-81-C (RET 323-1, pale pileus variant), 8-16-81-D (RET 323-2, dark pileus variant).
MISSOURI—Camden Co. - ca. Camdenton, Ha Ha Tonka St. Pk., River Cave, 24.vi.2009 Sherry Kaye s.n. [Tulloss 6-24-09-H] (RET 441-3), ca. Camdenton, Ha Ha Tonka St. Pk., Turkey Pen Hollows [37º58.183’ N/ 092º45.263’ W], 28.vi.2009 Jay Justice s.n. [Tulloss 6-28-09-B] (RET 441-9). Ste. Genevieve Co. - W of Ste. Genevieve, Hawn St. Pk. [37.8337° N/ 90.2416° W, 262 m], 13.viii.2011 Patrick Harvey s.n. [mushroomobserver.org #73884] (RET 494-10).
NEW JERSEY—Burlington Co. - W of Chatsworth, Franklin Parker Preserve, woods by north gate [39°48.827’ N/ 74°32.857’ W, 29 m], 14.x.2012 Naomi Goldman, Cristina Rodríguez Caycedo & R. E. Tulloss [Tulloss] 10-14-12-B (RET 518-5); Penn St. For., Oswego Lk. [39°44’02” N/ 74°29’26” W, 16 m], 2.viii.1981 M. A. King & R. E. Tulloss 8-2-81-A (RET 161-8). Middlesex Co. - East Brunswick, 176 Tice's Lane [40°27'15.41" N/ 74°25'09.29" W, 12 m], 22.vi.1989 Dr. Eugene Varney s.n. [Tulloss 6-22-89-A] (RET 248-3); Jamesburg, ca. Helmetta, Jamesburg Mun. Pk. [40°23’07” N/ 74°25’48” W], 9.viii.1981 M. A. King, Aaron Norarevian & R. E. Tulloss 8-9-81-B (RET 109-5), A. Norarevian & R. E. Tulloss 8-9-81-I (RET 172-6, dark cap variant), 28.vii.1985 Mark H., David C. & R. E. Tulloss 7-28-85-C (RET 204-1, pale pileus variant), 7-28-85-D (RET 204-2, pale pileus variant), 30..viii.1987 D. C. Tulloss s.n. [Tulloss 8-30-87-D] (RET 018-3). Monmouth Co. - Shark River Co. Pk. [40°12’18” N/ 74°05’44” W, 16 m], 19.viii.2011 L. K., M. A., O. C. & R. E. Tulloss & C. Rodríguez Caycedo [Tulloss] 8-19-11-A (RET 485-1); Upper Freehold, ca. Imlaystown, Clayton Co. Pk., Bridges Tr., 19.viii.2012 M. A. & R. E. Tulloss 8-19-12-E (RET 508-5, pallid pileus); Upper Freehold Twp., Assunpink Wildlife Mgmt. Area, 3.viii.1981 M. A. King & R. E. Tulloss 7-3-81-F (RET 115-1, dark pileus variant with progressive spore prints from single basidiome). Morris Co. - Hackettstown Reservoir, 26.vii.1984 R. E. Tulloss, Jim Richards, Robert Peabody, Geoffrey Kibby [Tulloss 7-26-84-L] (RET 052-7, dark pilleus variant); Mendham, Meadowood Twp. Pk. [40°47'31" N/ 74°38'43" W, 214 m], 22.vii.1984 S. Hopkins, G. Kibby & R. E. Tulloss 7-22-84-K (RET 051-7).
NEW YORK—Onondaga Co. - NW of Beaver Lk., 4.viii.1995 R. Archambault s.n. [Tulloss 8-4-95-B] (RET 154-5, brown pileus variant). Otsego Co. - Oneonta, 15.viii.1985 Connie Long s.n. [Tulloss 8-15-85-B] (RET 099-8). Ulster Co. - Rochester Twp., Minnewaska St. Pk., 19.viii.1994 Stachula s.n. [Tulloss 8-19-94-C] (RET 129-10, pale pileus variant).
OHIO—Butler Co. - Oxford Twp., Hueston Woods St. Pk., High Banks, 3.vii.1992 Mary W. & Michael A. Vincent 5445 (RET 069-6).
PENNSYLVANIA—Allegheny Co. - McCandless Twp., North Park, unkn. loc., 21.ix.2013 unkn. coll. s.n. [WPMC 20130921-93] (RET 568-3).
SOUTH CAROLINA—Oconee Co. - Sumter Nat. For., Stumphouse Tunnel Pk., 16.vii.1983 M. A. King & R. E. Tulloss 7-16-83-K (RET 219-10, pale pileus variant), 7-16-83-L (RET 219-9, pale pileus variant), 7-16-83-M (RET 220-1, dark pileus variant); Seneca, 28.xi.1985 R. E. Tulloss 11-28-85-A (RET 131-3, dark pileus variant).
TENNESSEE—Sevier Co. - ca. Gatlingburg, GSMNP, Cherokee Orchard [35°40'30.88” N/ 83°29'09.04” W, 805 m], 12.vii.2004 Dr. R. H. Petersen s.n. [Tulloss 7-12-04-AB] (RET 378-1).
VERMONT—Bennington Co. - ca. Bennington, Bennington Fish Culture Stn. [42°51'10" N/ 73°10'12" W, 288 m], 24.viii.1980 Adair, M. A. King & R. E. Tulloss 8-24-80-H (RET 392-9), M. A. King & R. E. Tulloss 8-24-80-C (RET 392-8).
WEST VIRGINIA—Greenbrier Co. - woods above rest stop along U.S. Rte. 60 ca. 1.6 km W of Charmco, 9.viii.1990 M. A. Vincent 4384 (in herb. C. Lavorato; MU F38505; RET 148-5). Marion Co. - Mill Fall Run, 5.x.1993 S. L. Stephenson 93-02 (FWVA).
We have for a long time held with Pomerleau () that the proposed variety pallida of the present species was without taxonomic value—having found a fairy ring of A. brunnescens containing brown pilei, pale citrin pilei, and at least one pileus that was pale citrin on one side and brown on the other. Recent unpublished genetic studies find that the material with brown caps and material with pallid caps may have no differences in their nrITS sequences (the presently proposed "fungal barcode) at all and have not been found differing in more than 4 base pairs (bp).
The present species is not strongly differentiated from A. aestivalis by the shape and size of spores as seen in the following sporograph comparison:
Limited DNA evidence (unpublished nrITS data) suggests that the present taxon and A. aestivalis may not be separable by the currently proposed "fungal barcode." At present they are distinguished by cap color and difference in speed and color of their staining reactions.
—R. E. Tulloss
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can be found here.
G. F. Atk.
"Brown American Star-Footed Amanita"
1. Amanita brunnescens, Maine, U.S.A.
2. Amanita brunnescens, South Carolina, U.S.A.
3. Amanita brunnescens, New York [state], U.S.A.
4. Amanita brunnescens, South Carolina, U.S.A.
Dr. Samuel S. Ristich - (1) Maine, U.S.A.
RET - (2, 4) South Carolina, U.S.A.; (3) New York [state], U.S.A.
Each spore data set is intended to comprise a set of measurements from a single specimen made by a single observer;
and explanations prepared for this site talk about specimen-observer pairs associated with each data set.
Combining more data into a single data set is non-optimal because it obscures observer differences
(which may be valuable for instructional purposes, for example) and may obscure instances in which
a single collection inadvertently contains a mixture of taxa.